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.     

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.     

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.     

Reconciliation and third‐party affiliation in carrion crows

Conflicts are costly because they can damage social relationships. To buffer conflicts, various species use post‐conflict behaviour, such as reconciliation or third‐party affiliation. Both behaviours have predominantly been studied in non‐human primates. However, recently, studies revealed post‐conflict behaviour in other mammalian and some bird species (e.g., corvids). While third‐party affiliation has been reported in several corvid species, reconciliation has only rarely been observed. The social structure of the studied groups has been postulated as a reason for the absence of reconciliation. Here, we investigated whether post‐conflict behaviours in corvids indeed mirror the relationship structure. We studied the behaviour of a newly established group of juvenile carrion crows (Corvus corone corone), where pair bonds had not yet been established. We applied a combination of observations and food monopolisation experiments to quantify the use of post‐conflict behaviours. Provisioning food in one or two pieces induced different patterns of aggression during feeding and differently affected the affiliation patterns after feeding. Specifically, victims of severe aggression affiliated with third parties after conflicts in the two‐piece condition, while aggressors affiliated with victims of mild aggression in the one‐piece condition. We thus provide the first evidence that a corvid species, crows, flexibly engage in both third‐party affiliation and reconciliation.     

Post-Conflict Affiliation in Barbary Macaques is Influenced by Conflict Characteristics and Relationship Quality, but Does Not Diminish Short-Term Renewed Aggression

Many group living primates have evolved mechanisms to repair their social relationships after conflicts ('reconciliation'). We analysed the post-conflict behaviour of female Barbary macaques, Macaca sylvanus , living in the enclosure 'La Forêt des Singes' at Rocamadour, France. Based on a sample of 914 conflicts, we investigated whether relationship (kinship, rank, affiliation, support and sex) and conflict characteristics (conflict intensity, context and duration) affected the quality and frequency of affiliative post-conflict interactions. Thirty-two per cent of all conflicts were followed by post-conflict affiliation. Rates of socio-positive interactions and support were better predictors of post-conflict affiliation than kinship or rank. Short conflicts were followed by post-conflict affiliation relatively more frequently, after a shorter latency, but only briefly, and such interactions were initiated by both parties equally frequently. The majority of affiliative post-conflict interactions occurred immediately after the end of the conflict. In sum, female Barbary macaques invest more in post-conflict affiliation with valuable partners, and they modulate their post-conflict behaviour in relation to conflict characteristics. Remarkably, affiliative post-conflict interactions increased the short-term probability of renewed aggression by the former aggressor to 16% compared with 9% for conflicts that were not followed by affiliative behaviour. Such renewed aggression after post-conflict affiliation occurred particularly frequently among females and after conflicts over food, suggesting that post-conflict affiliation sometimes falsely lures the former victim to stay in the vicinity, even at the risk of receiving renewed aggression.     

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.     

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.     

Post‐Conflict Non‐Aggressive Behaviours may be neither Friendly nor Conciliatory: Conflict Management of Male Hamadryas Baboons

The relational model of conflict resolution predicts that after an aggressive conflict there should be a motivational shift from aggression to attraction. Most tests of the reconciliation hypothesis assume, however, that all non‐aggressive post‐conflict behaviours between former opponents are motivationally homogeneous and qualify as friendly reunions. In fact, although the hypothesis predicts an increased occurrence of friendly contacts after conflicts, in practice, however, post‐conflict reunions often include a mixture of contact and non‐contact behaviours. Most reconciliation studies either (often) assume a conciliatory function for post‐conflict reunions or (less often) test functional predictions. Finally, the valuable relationships hypothesis predicts that conciliatory rates should be relatively higher between friends and allies than between non‐friends/allies. In this paper, we use data on non‐aggressive interactions following conflicts between adult male hamadryas baboons that are neither friends nor allies to assess the implications of all these important but largely overlooked issues. The analyses of the rate and temporal relation of non‐contact greeting (NCTG) to anxiety‐related behaviours and side‐directed aggression as well as of the behaviours used during non‐aggressive interactions with male and female third‐parties suggest that the NCTG used by males after conflicts were neither motivationally friendly nor functionally conciliatory. We point out that the gestures exchanged during these post‐conflict NCTG can be interpreted as formalized signals of equal status and that the rate and form of the greetings used by male opponents are indicative of high relationship insecurity and incompatibility respectively. We conclude that although male hamadryas’ post‐conflict NCTG are not conciliatory they may serve to assess their opponents’ attitude and to negotiate the restoring of their pre‐conflict levels of peaceful but non‐amicable co‐existence.     

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 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.     

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 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.     

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.     

Post-conflict affiliation in common marmosets (Callithrix jacchus jacchus)

Post-conflict (PC) affiliation has been demonstrated in a number of Old World monkeys and apes, but very little is known about the occurrence of the phenomenon in New World monkeys. This study examined 282 PC interactions after spontaneous conflicts around feeding time in two family groups (N = 12) of captive common marmosets, Callithrix jacchus jacchus. We found an overall corrected conciliatory tendency of 31%. Selective attraction was seen: former opponents met significantly more often in PCs than in matched controls (MCs) (27% and 16%, respectively). There was no difference in the occurrence of PC affiliation between dyads consisting of parent-offspring constellations compared to offspring-offspring constellations. PC affiliative behaviors were seen in the first three minutes following conflict termination, and consisted mainly of proximity, play invitations, and food transfer. Notably, former opponents remained within arm's reach after 17% of conflicts. Affiliation was more likely to follow after conflicts involving play issues. The functional importance of the PC affiliation in marmosets remains to be examined.     

The impact of paternity on male–infant association in a primate with low paternity certainty

In multimale groups where females mate promiscuously, male–infant associations have rarely been studied. However, recent studies have shown that males selectively support their offspring during agonistic conflicts with other juveniles and that father's presence accelerates offspring maturation. Furthermore, it was shown that males invest in unrelated infants to enhance future mating success with the infant's mother. Hence, infant care might provide fitness gain for males. Here, we investigate male–infant associations in rhesus macaques (Macaca mulatta), a primate with low paternity certainty as females mate with multiple partners and males ensure paternity less efficiently through mate‐guarding. We combined behavioural data with genetic paternity analyses of one cohort of the semi‐free‐ranging population of Cayo Santiago (Puerto Rico) and recorded affiliative and aggressive interactions between focal subjects and adult males from birth to sexual maturation (0–4 years) of focal subjects. Our results revealed that 9.6% of all interactions of focal subjects involved an adult male and 94% of all male–infant interactions were affiliative, indicating the rareness of male–infant aggression. Second and most interestingly, sires were more likely to affiliate with their offspring than nonsires with unrelated infants. This preference was independent of mother's proximity and emphasized during early infancy. Male–infant affiliation rose with infant age and was pronounced between adult males and male rather than female focal subjects. Overall, our results suggest that male–infant affiliation is also an important component in structuring primate societies and affiliation directed towards own offspring presumably represent low‐cost paternal care.     

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.     

Postconflict behavior among female Sichuan snub-nosed monkeys Rhinopithecus roxellana within one-male units in the Qinling Mountains, China

For group-living primates, the information on postconflict management is crucial for understanding primate competition and cooperation. However, such information is poorly known for snub-nosed monkeys, especially for wild populations. In this study, from September 2007 to June 2008, we investigated postconflict behavior among adult females Sichuan snub-nosed monkeys Rhinopithecus roxellana within one-male units in a wild, provisioned group in the Qinling Mountains of China by means of the time-rule method and the PC-MC method. We obtained a total of 81 PC-MC pairs and each individual was involved in only 0.004 aggressive behavior per observation hour. The first affiliative behavior was more likely to occur within the first minute after a conflict. The postconflict affiliative behaviors most often seen were contact-sit, embrace and grooming. The affiliative contacts between adult females occur due to selective attraction, I.e. Reconciliation. The pattern of postconflict affiliation demonstrates that the R. Roxellana belongs to a tolerant species.     

Post‐birth separation affects the affiliative behaviour of kin in a pitviper with maternal attendance

The relatedness of individuals can have pronounced effects on behavioural interactions, as engaging in mutually beneficial behaviours with kin can increase inclusive fitness. Parental care can be particularly important for kin discrimination in birds and mammals, but similar studies have not been conducted on species exhibiting more rudimentary forms of care. Maternal attendance of young is ubiquitous among viviparous temperate pitvipers, but the adaptive value of this behaviour has received little attention. We sought to determine if being deprived of a maternal attendance period as neonates altered how cottonmouths (Agkistrodon piscivorus Lacépède), a common North American pitviper, responded to kin vs. non‐kin. We measured the affiliative behaviour of related and unrelated juvenile–juvenile and mother–juvenile pairs that had been allowed a maternal attendance period or had been separated since birth. We found that maternal attendance was not required for sibling or mother–offspring recognition, but did enhance female affiliative behaviour overall, and particularly that of sisters. In contrast, related juveniles that were separated at birth showed a reversal of the sex‐specific affiliative behaviour observed in maternally attended juveniles. Post‐birth separation had only a modest effect on mother–juvenile affiliative behaviour, and no effect on the strong affiliation between mothers and daughters. The patterns of affiliative behaviour observed in maternally attended snakes corresponded to patterns that have emerged from previous captive and field studies of pitvipers; however, the behaviour of juveniles separated at birth was atypical. Thus, it is possible that maternal attendance plays some role in the development of adaptive sex‐specific and kin‐directed affiliative behaviour in pitvipers.     

Affiliation in Four Echimyid Rodent Species based on Intrasexual Dyadic Encounters: Evolutionary Implications

In this work we explored the evolution of sociality in cursorial echimyids by comparing affiliation among three species of Trinomys and one species of Thrichomys. We captured specimens of Trinomys yonenagae, Trinomys albispinus minor, and Thrichomys apereoides in areas of the Brazilian semiarid Caatinga, and Trinomys iheringi denigratus in one area of Atlantic Forest. For each species, we recorded 12 intra‐sexual dyadic encounters in a neutral arena (six between males and six between females) in order to test the hypothesis that species and sex influence level of affiliation. This response variable was assessed based on an affiliation index, calculated as the proportion of the total number of affiliative behaviors to the total number of social behaviors exhibited by the dyad during each encounter. Hypothesis test was performed by means of a parametric two‐way anova . The test was able to detect significant differences only among species, not among sexes. Trinomys yonenagae was the most affiliative species, while T. apereoides and T. albispinus minor were the most agonistic ones. Trinomys iheringi denigratus showed an intermediate pattern. We suggest, based on out‐group comparison, that affiliation in Trinomys increased in the lineage containing T. iheringi denigratus and T. yonenagae and that higher affiliation in the last species can be adaptive to the life in the desert‐like habitat where it lives.