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.     

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.     

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).     

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.     

Post‐conflict third‐party affiliation in chimpanzees: what's in it for the third party?

Affiliative behavior after conflicts between conflict participants and other group members is common in many primate species. The proposed functions for such triadic interactions are numerous, mostly concerning the benefit for the former conflict opponents. We investigated post‐conflict third‐party affiliation (TPA) in captive chimpanzees (Pan troglodytes) with the aim of assessing what the affiliating third parties may gain from affiliation. Specifically, we tested whether third‐party‐initiated affiliation protects the third parties from further aggression by conflict opponents. We found support for this “self‐protection hypothesis,” in that third parties selectively directed affiliation to those opponents who more often gave further aggression to them, and affiliation effectively decreased their chance of receiving aggression from these opponents. However, a subset of affiliation, provided to conflict victims by their own kin, appeared to not be self‐protective and the function of it remained open. We conclude that chimpanzee third‐party‐initiated affiliation is a more heterogeneous behavior than thus far assumed. Am. J. Primatol. 71:409–418, 2009. © 2009 Wiley‐Liss, Inc.     

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.     

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.     

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.     

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.     

Post-conflict Social Events in Wild Mountain Gorillas. II. Redirection,Side Direction,and Consolation

Inevitably, members of primate groups sometimes have aggressive contests with each other. Targets of aggression can engage in several types of interaction with third parties to ameliorate its adverse effects. They can redirect aggression, which may reduce the risk of further aggression from the initial opponents and reduce tension, or they can initiate affinitive interactions with third parties, in order to seek protection or reassurance. They can also receive reassuring acts from them (‘consolation’). Researchers have documented high levels of redirection in many primate species, but consolation is thus far known only in chimpanzees. Data on post-conflict social interactions between targets of aggression and third parties in two groups of wild mountain gorillas (Gorilla gorilla beringei) show that immatures and subordinate males, but not females, redirect aggression at high rates. Furthermore, immatures seek affinitive interactions with their mothers, and adult females seek them with adult males, at elevated rates. Affinitive interactions between females and males seem mainly to have a protective function and are associated with decreased risks of further aggression between female opponents. Redirection by females after female-female conflicts may be uncommon because targets commonly retaliate against aggressors. Males may offer females protection and consolation as services, and the results support the argument that such services are important when dominance relationships between females are often undecided and retaliation between opponents is common.     

Post-conflict Social Events in Wild Mountain Gorillas (Mammalia,Hominoidea). I. Social Interactions between Opponents

Researchers have documented elevated rates of affinitive interaction between opponents shortly after aggressive conflicts, or reconciliation, in many primate species. Reconciliation may ameliorate the immediate negative effects of aggression by reducing the chance of further aggression between opponents and thus reducing tension, and may avert or repair damage to long-term social relationships important to the animals' fitness. Data on post-conflict interactions between opponents in two groups of wild mountain gorillas (Gorilla gorilla beringei) fail to show reconciliation after conflicts between females, between males, or between immatures, but females seem to commonly reconcile themselves with males after receipt of male aggression. Females and subordinate males often avoid same-sex opponents after conflicts. Females commonly retaliate against female aggressors, and post-conflict rates of aggression between females are higher than baseline levels. Females may not need to achieve reconciliation with each other because relationships between co-resident relatives are resiliant, while those between non-relatives are mostly neutral to antagonistic. Males are, however, important social partners and protectors of females, and female transfer is common. Thus, the results strongly support the ‘important relationships’ hypothesis for the function of reconciliation.     

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.     

Aggression and Reconciliation in Cebus capucinus

Most data relating to aggressive and conciliatory behaviors are from Old World primates. We recorded agonistic interactions and post-conflict behaviors in a group of 12 white-faced capuchins (Cebus capucinus). After a conflict, we followed the aggressee as the focal individual during a 10-min postconflict period. We also conducted matched-control observations on the same individual. Conflicts involving physical contact were significantly bidirectional, and conflicts without physical contact were preferably unidirectional. Reconciliation was not be demonstrated at the group level. However, reconciliation occurred in kin and non-kin male/female dyads: their conciliatory tendencies were 48.1% and 21.2%, respectively. White-faced capuchins reconciled mainly during the first minutes after the end of the conflict. In kin and non-kin male/female dyads, selective attraction occurred, and aggressors were more likely to initiate affiliative contacts than aggressees. Hold-bottom and mount while emitting loud vocalizations were the most characteristic behaviors of reconciliation. Possible links may exist between aggressive and conciliatory patterns and other social variables.     

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.     

The Long-Term Effects of Reconciliation in Japanese Macaques Macaca fuscata

With one exception, all previous studies of reconciliation in non-human primates (friendly reunion between former opponents) have focused on demonstrating the immediate, short-term effects despite the widely held view that reconciliation has a long-term function of repairing social relationships following aggression. To investigate this long-term function I compared mean interaction rates between opponents during the 10 d following reconciled and non-reconciled conflicts to baseline levels of interaction. Aggression rates during the 10 d after non-reconciled conflicts were significantly higher than the baseline rate, whereas after reconciled conflicts aggression was minimal. Similarly, grooming, proximity and approach rates during the 10 d after non-reconciled conflicts were significantly lower than the baseline rate whereas grooming, proximity and approach rates in the 10 d after reconciled conflicts were restored to baseline levels. These results indicate that there are consequences to not reconciling with a former opponent and highlight the fact that these may be costly in terms of increased risk of long-term aggression and reduced affiliation. The data support predictions from the Relationship-Repair Hypothesis suggesting that reconciliation functions as a mechanism for the repair of social relationships damaged by aggression.     

Aggressive and neutral interventions in conflicts in captive groups of brown lemurs (Eulemur fulvus fulvus)

Third-party interventions in conflicts have revealed complexity in primate social relationships. This type of intervention has seldom been analyzed in prosimians, although many of these species exhibit complex (multimale/multifemale) social organizations. The present study on captive brown lemurs (Eulemur fulvus fulvus) shows that dominant individuals were more likely to intervene in conflicts. Both males and females intervened aggressively in conflicts. Female aggressive interventions occurred mainly on behalf of close kin, whereas males mainly intervened on behalf of juveniles. This study also provides the first record of neutral or peaceful interventions in lemurs. Although females intervened neutrally, almost all neutral interventions were by dominant males. Dominant males intervened in conflicts neutrally more often than aggressively, principally in conflicts between adults and juveniles or between juveniles. Neutral interventions by males always ended the conflicts and were often followed by affiliative contacts between participants (intervenors and opponents). In lemurs, female interventions can be explained by kin selection, while the nature of dominant males' interventions suggests a control role. Interventions by males on behalf of juveniles may increase the formers' fitness.     

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.     

The integration of dominance and social bonding in primates

Social dominance is usually viewed from the perspective of intragroup competition over access to limited resources. The present paper, while not denying the importance of such competition, discusses the dominance concept among monkeys and apes in the context of affiliative bonding, social tolerance, and the reconciliation of aggressive conflicts. Two basic proximate mechanisms are supposed to provide a link between dominance and interindividual affiliation, namely, formalization of the dominance relationship (i.e., unequivocal communication of status), and conditional reassurance (i.e., the linkage of friendly coexistence to formalization of the relationship). Ritualized submission is imposed upon losers of dominance struggles by winners; losers are offered a "choice" between continued hostility or a tolerant relationship with a clearly signalled difference in status. If these two social mechanisms are lacking, aggression is bound to have dispersive effects. In their presence, aggression becomes a well-integrated, even constructive component of social life. In some higher primates this process of integration has reached the stage where status differences are strongly attenuated. In these species, sharing and trading can take the place of overt competition. The views underlying this "reconciled hierarchy" model are only partly new, as is evident from a review of the ethological literature. Many points are illustrated with data on a large semi-captive colony of chimpanzees (Pan troglodytes), particularly data related to striving for status, reconciliation behavior, and general association patterns. These observations demonstrate that relationships among adult male chimpanzees cannot be described in terms of a dichotomy between affiliative and antagonistic tendencies. Male bonding in this species has not been achieved by an elimination of aggression, but by a set of powerful buffering mechanisms that mitigate its effects. Although female chimpanzees do exhibit a potential for bonding under noncompetitive conditions, they appear to lack the buffering mechanisms of the males.     

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.