Post-conflict Behaviour Research Articles

Reconciling conflicts in a one-male society: the case of geladas (Theropithecus gelada)

Animals derive benefits from living in social groups but sociality also has its costs in that animals must compete with others for resources and mating opportunities. To cope with the conflict aftermath and social damage caused by competitive aggression, several group-living species use a variety of peace-keeping strategies. The affinitive post-conflict reunion of former opponents, defined as reconciliation, is the primary peace-keeping mechanism. In this study, we provide evidence for the occurrence of reconciliation and test some hypotheses on this post-conflict mechanism in geladas (Theropithecus gelada), a species often neglected in the study of post-conflict dynamics. The conciliatory contacts were uniformly distributed across the different sex-class combinations. Different from baboons, geladas did not show any particular kind of affinitive reconciliation behaviour. Notwithstanding the presence of a linear hierarchy, the dominance relationships did not affect the reconciliation dynamics. According to the valuable relationship hypothesis, coalitionary support seems to be a good predictor for a high level of conciliatory contacts. Finally, at an immediate level reconciliation plays a role in reducing renewed attacks by aggressors, which sought conciliatory contact more frequently than victims. In conclusion, even though the study of post-conflict behaviour in geladas needs to be continued, the patchy nature of their social network is a good model for testing some of the theoretical assumptions about primate conflict resolution.

Postconflict Behaviour in Brown Capuchin Monkeys (Cebus apella)

Postconflict affiliation has been mostly studied in Old World primates, and we still lack comparative research to understand completely the functional value of reconciliation. Cebus species display great variability in social characteristics, thereby providing a great opportunity for comparative studies. We recorded 190 agonistic interactions and subsequent postconflict behaviour in a captive group of brown capuchin monkeys (Cebus apella). Only 26.8% of these conflicts were reconciled. Reconciliation was more likely to occur between opponents that supported each other more frequently and that spent more time together. Postconflict anxiety was mostly determined by conflict intensity, and none of the variables thought to measure relationship quality had a significant effect on postconflict stress.

Postconflict Affiliation of Aggressors in Papio hamadryas

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.

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.

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.

Peacemaking on treetops: first evidence of reconciliation from a wild prosimian ( Propithecus verreauxi)

Reconciliation is defined as the first postconflict affinitive contact between former opponents. While reconciliation in anthropoid primates has been widely investigated, few studies have focused on postconflict mechanisms in prosimians, and only in captivity. Unlike anthropoids, Malagasy prosimians show female dominance, lack of sexual dimorphism and seasonal breeding. However, they share features with anthropoids such as cohesive societies, female philopatry and individual recognition. Comparing social prosimians with anthropoids is crucial for understanding the evolution of reconciliation dynamics. Here we present the first study on reconciliation in a wild prosimian. We focused on the Propithecus verreauxi (sifaka) of the Berenty forest (southern Madagascar). We examined postconflict behaviour in the light of theoretical expectations based on potential costs and benefits of the individuals involved. Our results indicate that P. verreauxi can evaluate possible risks and benefits of engaging in postconflict reunions. Victims were most likely to interact affinitively with the aggressor after low-intensity aggression. Moreover, only the conflicts occurring outside the feeding context were reconciled. Such results are consonant with the fact that, in P. verreauxi, social dominance is translated more into feeding priority than into a framework of despotic relationships. In agreement with the valuable relationship hypothesis, P. verreauxi were more likely to reconcile with valuable partners: reconciliation preferentially occurred between subordinates and top-ranking individuals, and between animals sharing good relationships (high levels of affinitive behaviours). Over the short term, reconciliation in P. verreauxi seems to have an important role in reducing the probability of further attacks by the aggressor.

Reconciliation, dominance and cortisol levels in children and adolescents (7–15-year-old boys)

The stress reduction hypothesis of reconciliation (Aureli, 1997; Aureli & Smucny, 2000) is tested on hormonal level. This study explores the potential relationships between post-conflict behaviour and cortisol level in boys between 7 and 15 years of age observed in summer camp during their free-time playing. Data on 56 boys are presented. Cortisol levels were measured in the morning (fixed time of the day), about 15 min after the end of conflict (PC) and next day without conflict as a match control (MC). Post-conflict of two types were analyzed: those followed with affiliation between former opponents and without affiliation. Age and dominance status of boys were taken in account in analyses of relationships between cortisol and post-conflict behaviour. The differences of cortisol level according to the occurrence of reconciliation were demonstrated. The stress reduction hypothesis of reconciliation was confirmed at the physiological level in children and adolescents.

Postconflict behavior in captive western lowland gorillas (Gorilla gorilla gorilla)

Postconflict (PC) behaviors, including reconciliation and consolation, have been observed in many primate and several nonprimate species. Using the PC-matched control (MC) method, PC behavior was examined in two groups (n=13) of captive western lowland gorillas, a species for which no conflict resolution data have been published. Analyses of 223 conflicts showed significantly more affiliation between former opponents after a conflict when compared to control periods, indicating reconciliation. Results also showed significantly more affiliation between the victim and a third-party after a conflict, indicating consolation. Both solicited and unsolicited consolation were observed. The majority of the affiliative interactions observed for both reconciliation and consolation were social proximity, which suggests that unlike most nonhuman primates, proximity, rather than physical contact, may be the main mechanism for resolving conflicts in western lowland gorillas. PC behavior was not uniform throughout the groups, but rather varied according to dyad type.

Postconflict Behavior Among Male Japanese Macaques

Reconciliation was first described more than 20 years ago. Since then, it has been observed in many mammals (mainly primates) but data on postconflict behavior among males are still scarce because they usually aggressively compete for mating partners, rarely maintain amicable relationships with one another. Accordingly, reconciliation is expected to occur at low rates. Although this is true for Japanese macaque males, the subspecies on Yakushima Island (Macaca fuscata yakui) seems to represent an exception as grooming among males occurs often. We analyzed postconflict behavior among them and discuss the possible factors that may favor the occurrence of grooming and reconciliation. Selective attraction between former opponents—reconciliation—occurred soon after conflicts. Consolation—affiliative interactions between a focal animal and group members other than the former opponents occurring earlier in PCs than in MCs—was absent among males. Conciliatory tendency is higher for Yakushima macaque males (0.31) versus that in studies on the other subspecies Macaca fuscata. We discuss differences in the behavioral ecology of the 2 subspecies, the ecological and social factors that may favor the occurrence of reconciliation, and the possible benefits that males gain from grooming exchange and reconciliation.

Reconciliation in wild white-faced capuchins (Cebus capucinus)

The likelihood of reconciliation (defined as preferential peaceful contact among former opponents following conflicts) has been predicted to vary positively with relationship value and compatibility, and negatively with relationship security. Long-term data on wild white-faced capuchins (Cebus capucinus) indicate that dyads consisting of an adult female and an alpha male have high value and compatibility, but low security. Two studies of C. capucinus postconflict (PC) behavior were carried out at Lomas Barbudal Biological Reserve, Costa Rica. One study consisted of 30-min PC and matched control (MC) follows. The second study extracted PC and MC periods from long follows, yielding PC/MC periods averaging 105 min. In study 2, but not study 1, significantly more PC/MC pairs were attracted (former opponents affiliated with each other sooner in the PC period than in the MC period) than were dispersed (former opponents affiliated with each other sooner in the MC period than in the PC period). Reconciliation in study 2 could not be explained as a by-product of former opponents' tendency to seek affiliative contact with conspecifics generally, or of the spatial proximity of opponents following conflicts. Attempted reconciliation was less likely to be followed by renewed aggression when reconciliation attempts were delayed following conflicts. The data were insufficient for a formal test of differences in conciliatory tendency (the difference between the number of attracted and dispersed PC/MC pairs, divided by the total number of pairs) among dyad types to be conducted.

Conflict and post-conflict behavior in a small group of chimpanzees.

Chimpanzee research plays a central role in the discussions of conflict negotiation. Reconciliation, or the attraction and affiliation of former opponents following conflict, has been proposed as a central element of conflict negotiation in chimpanzees and various other taxa. In an attempt to expand the database of chimpanzee conflict resolution, conflict and post-conflict behavior were recorded for a small group of socially housed chimpanzees at the Chimpanzee and Human Communication Institute, at Central Washington University. Data were collected over six 6-week periods between 1997 and 2000, for a total of 840 hours of observation, resulting in a substantial post-conflict (PC) and matched control (MC) data set. The data demonstrate this group's tendencies to maintain visual contact and closer proximity after conflicts. Dyadic corrected conciliatory tendencies ranged between 0 - 37.5% and averaged 17.25% across all dyads. Individual corrected conciliatory tendencies ranged between 5.8 and 32%. The results of this study combined with recent publications on captive and free-ranging chimpanzee post-conflict behavior suggest that variation in post-conflict behavior may be important to our understanding of chimpanzee conflict negotiation, and may also have implications for the design and management of captive chimpanzee enclosures and social groups, respectively.

Postconflict behaviour of wild Indian langur monkeys: avoidance of opponents but rarely affinity

Friendly postconflict (PC) interactions between former opponents have been described for a large number of primate species held in captivity. We investigated conflict management behaviour for wild Hanuman langurs, Semnopithecus (Presbytis) entellus entellus, in India, using a large sample of over 6000 agonistic and affinitive interactions recorded in one one-male/multifemale troop and three all-male bands. We compared dyadic PC affinity with the dyadic baseline affinity during the overall observation time to minimize biases of traditional matched-control samples. PC affinity was recorded during only 15% of all dyads (immature male–immature male 0%, adult male–adult male 0%, adult male–immature male 5%, adult male–adult female 0%, adult female–adult female 42%). PC affinity is probably absent amongst males because their dominance relationships are strongly asymmetrical, leaving little room for emotional insecurity. Dyads among females, on the other hand, reflect frequent rank changes and close kinship which probably corresponds to higher levels of emotional uncertainty and greater need for PC affinity. Overall, PC affinity reached only 26% of the randomly expected value. Thus, the vast majority of langur monkey opponents avoided each other after conflicts. Avoidance as a low-cost option for group-living animals to cope with conflicts is often not possible in captivity. This suggests that reports of high rates of reconciliation may be at least partly artefacts of captivity. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.

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.

Conflict and postconflict behaviour in two lemur species with different social organizations (Eulemur fulvus and Eulemur macaco): A study on captive groups

AbstractAlthough the brown lemur (Eulemur fulvus) and the black lemur (Eulemur macaco) share many life parameters and are genetically closed, they show considerable difference in social organization. Dominance relationships with no systematic effect of gender characterize the former, whereas the latter is based on female dominance. The present study was done on two captive groups of brown lemurs and one semi–free‐ranging group of black lemurs. To reveal links between the specific pattern of social organization and agonistic behaviors, agonistic interactions were analyzed for each species as for their context of occurrence, symmetry, initiation, and outcome. The effect of gender in the initiation of conflicts appeared as the only notable interspecific difference, aggression being mostly initiated by females in E. macaco and by males in E. fulvus. Conflict outcome was generally in favor of the initiator, regardless of gender in both species. The analysis of postconflict behaviors revealed conciliatory processes in the brown lemur, whereas reconciliation seemed to be absent in the black lemur, a characteristic shared with the ring‐tailed lemur (Lemur catta), another lemur species with female dominance. Aggr. Behav. 28:62–74, 2002. © 2002 Wiley‐Liss, Inc.

Reconciliation and variation in post-conflict stress in Japanese macaques (Macaca fuscata fuscata): testing the integrated hypothesis.

Reconciliation in primates, a post-conflict affiliative interaction between former opponents, appears to have two functions: (1) to repair relationship damaged by aggression such that animals who share more valuable relationships are more likely to reconcile, and (2) to reduce the post-conflict uncertainty and stress of former combatants. The 'integrated hypothesis' of reconciliation links these functions by arguing that the disturbance of a valuable relationship by aggression should result in particularly high levels of stress, which in turn should facilitate efforts to reconcile and thus gain relief from post-conflict stress. A key prediction of the integrated hypothesis is that victims of aggression suffer more stress following conflicts with individuals with whom they share a valuable relationship. In this article, we test the integrated hypothesis by observing the post-conflict behaviour of victims among a free-ranging provisioned troop of Japanese macaques (Macaca fuscata fuscata) living in Shiga Heights, Nagano, Japan. In this troop, monkeys reconciled roughly one in seven conflicts. The only factor that we could significantly relate to the occurrence of reconciliation was kinship; kin reconciled more frequently than non-kin did. Receiving aggression increased and reconciliation reduced the probability of being re-attacked after aggressive interactions, supporting the hypothesis that reconciliation repairs relationships. Victims' self-directed behaviour (SDB) - a behavioural index of stress comprising increases in scratching, self-grooming, and body-shaking - was elevated following aggression but decreased rapidly following reconciliation, supporting the idea that reconciliation functions to reduce post-conflict stress. Post-conflict SDB varied as follows: (1) victims showed a higher level of stress following aggression with kin than with non-kin, and (2) juvenile victims were less distressed than adults. The level of post-conflict SDB performed by juveniles following conflicts with kin was indistinguishable from that performed by adults but was greatly reduced following attacks from non-kin. These results indicate that post-conflict SDB keenly reflects the value of relationships between opponents, and that the post-conflict behaviour of free-ranging Japanese macaques fits the predictions of the integrated hypothesis.

POST-CONFLICT BEHAVIOUR OF WILD CHIMPANZEES (PAN TROGLODYTES SCHWEINFURTHII) IN THE BUDONGO FOREST, UGANDA

AbstractSince de Waal & van Roosmalen (1979) first documented the occurrence of reconciliation between former opponents in captive chimpanzees, the study of the post-conflict behaviour of primates has provided valuable information about some of the details of primate social organisation. The vast majority of these studies have been carried out on captive subjects and it has been assumed that these findings are representative of wild primates. We set out to investigate whether this was true for the Sonso community of wild chimpanzees (Pan troglodytes schweinfurthii) of the Budongo Forest, Uganda, using controlled procedures comparable with those used in captive studies. We found that these chimpanzees were much less likely to reconcile than their captive counterparts. Only one dimension of relationship quality had an effect on the likelihood of reconciliation. Individuals which were highly compatible, in terms of time spent affiliating, reconciled conflicts more often than those with weak relationships. Captive chimpanzees have also been shown to 'console' one another (de Waal & van Roosmalen, 1979; de Waal & Aureli, 1996), where uninvolved bystanders initiate affiliative contacts with victims of aggression. This study did not confirm that consolatory behaviour was characteristic of wild chimpanzee post-conflict behaviour. Nor did these chimpanzees use explicit gestures during post-conflict interactions as they have been shown to do in two out of three captive studies. We conclude that the post-conflict behaviour of chimpanzees is more variable than has previously been thought and is likely to be dependent on the prevailing social environment.

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.

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.

Reconciliation in captive Guyanese squirrel monkeys (Saimiri sciureus).

The tendency for agonistic interaction to increase the probability of friendly interaction between social partners has been demonstrated across a range of Old World primates. While research on such post-conflict behavior proceeds into an hypothesis-testing phase, new comparative information must accumulate to provide full phylogenetic perspective on primate social behavior. Data from New World and prosimian primates are yet extremely limited. We studied captive squirrel monkeys (Saimiri sciureus) via post-conflict (PC) and matched control (MC) observations and analyzed results using both the PC-MC and time-rule methods. Former opponents maintaining affiliative relationships soon engaged in friendly interaction following large proportions of agonistic interactions, whereas non-affiliated individuals, including virtually all male-female pairs, reconciled conflicts rarely. Close-proximity approaching and huddling contact constituted the principal modes of post-conflict amicability. Agonistic interactions of relatively high intensity were most likely to be reconciled and most likely to be reconciled via physical contact. High vulnerability of Saimiri to predation may have favored this species' strong inclination to reconcile soon after agonistic interaction. Research on free-living populations of this and other primate species is needed to illuminate similarities and differences across taxa.

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.