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Flack, J. C., de Waal, F. B. M., & Krakauer, D. C. (2005). Social structure, robustness, and policing cost in a cognitively sophisticated species. Am Nat, 165(5), E126–139.
Abstract: Conflict management is one of the primary requirements for social complexity. Of the many forms of conflict management, one of the rarest and most interesting is third-party policing, or intervening impartially to control conflict. Third-party policing should be hard to evolve because policers personally pay a cost for intervening, while the benefits are diffused over the whole group. In this study we investigate the incidence and costs of policing in a primate society. We report quantitative evidence of non-kin policing in the nonhuman primate, the pigtailed macaque. We find that policing is effective at reducing the intensity of or terminating conflict when performed by the most powerful individuals. We define a measure, social power consensus, that predicts effective low-cost interventions by powerful individuals and ineffective, relatively costly interventions by low-power individuals. Finally, we develop a simple probabilistic model to explore whether the degree to which policing can effectively reduce the societal cost of conflict is dependent on variance in the distribution of power. Our data and simple model suggest that third-party policing effectiveness and cost are dependent on power structure and might emerge only in societies with high variance in power.
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Johnstone, R. A. (2001). Eavesdropping and animal conflict. Proc. Natl. Acad. Sci. U.S.A., 98(16), 9177–9180.
Abstract: Fights between pairs of animals frequently take place within a wider social context. The displays exchanged during conflict, and the outcome of an encounter, are often detectable by individuals who are not immediately involved. In at least some species, such bystanders are known to eavesdrop on contests between others, and to modify their behavior toward the contestants in response to the observed interaction. Here, I extend Maynard Smith's well known model of animal aggression, the Hawk-Dove game, to incorporate the possibility of eavesdroppers. I show that some eavesdropping is favored whenever the cost of losing an escalated fight exceeds the value of the contested resource, and that its equilibrium frequency is greatest when costs are relatively high. Eavesdropping reduces the risk of escalated conflict relative to that expected by chance, given the level of aggression in the population. However, it also promotes increased aggression, because it enhances the value of victory. The net result is that escalated conflicts are predicted to occur more frequently when eavesdropping is possible.
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Flack, J. C., Girvan, M., de Waal, F. B. M., & Krakauer, D. C. (2006). Policing stabilizes construction of social niches in primates. Nature, 439(7075), 426–429.
Abstract: All organisms interact with their environment, and in doing so shape it, modifying resource availability. Termed niche construction, this process has been studied primarily at the ecological level with an emphasis on the consequences of construction across generations. We focus on the behavioural process of construction within a single generation, identifying the role a robustness mechanism--conflict management--has in promoting interactions that build social resource networks or social niches. Using 'knockout' experiments on a large, captive group of pigtailed macaques (Macaca nemestrina), we show that a policing function, performed infrequently by a small subset of individuals, significantly contributes to maintaining stable resource networks in the face of chronic perturbations that arise through conflict. When policing is absent, social niches destabilize, with group members building smaller, less diverse, and less integrated grooming, play, proximity and contact-sitting networks. Instability is quantified in terms of reduced mean degree, increased clustering, reduced reach, and increased assortativity. Policing not only controls conflict, we find it significantly influences the structure of networks that constitute essential social resources in gregarious primate societies. The structure of such networks plays a critical role in infant survivorship, emergence and spread of cooperative behaviour, social learning and cultural traditions.
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de Waal, F. B. (2000). Primates--A natural heritage of conflict resolution. Science, 289(5479), 586–590.
Abstract: The traditional notion of aggression as an antisocial instinct is being replaced by a framework that considers it a tool of competition and negotiation. When survival depends on mutual assistance, the expression of aggression is constrained by the need to maintain beneficial relationships. Moreover, evolution has produced ways of countering its disruptive consequences. For example, chimpanzees kiss and embrace after fights, and other nonhuman primates engage in similar “reconciliations.” Theoretical developments in this field carry implications for human aggression research. From families to high schools, aggressive conflict is subject to the same constraints known of cooperative animal societies. It is only when social relationships are valued that one can expect the full complement of natural checks and balances.
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Flack, J. C., Krakauer, D. C., & de Waal, F. B. M. (2005). Robustness mechanisms in primate societies: a perturbation study. Proc Biol Sci, 272(1568), 1091–1099.
Abstract: Conflict management mechanisms have a direct, critical effect on system robustness because they mitigate conflict intensity and help repair damaged relationships. However, robustness mechanisms can also have indirect effects on system integrity by facilitating interactions among components. We explore the indirect role that conflict management mechanisms play in the maintenance of social system robustness, using a perturbation technique to 'knockout' components responsible for effective conflict management. We explore the effects of knockout on pigtailed macaque (Macaca nemestrina) social organization, using a captive group of 84 individuals. This system is ideal in addressing this question because there is heterogeneity in performance of conflict management. Consequently, conflict managers can be easily removed without disrupting other control structures. We find that powerful conflict managers are essential in maintaining social order for the benefit of all members of society. We show that knockout of components responsible for conflict management results in system destabilization by significantly increasing mean levels of conflict and aggression, decreasing socio-positive interaction and decreasing the operation of repair mechanisms.
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Sutton, J. E., & Shettleworth, S. J. (2005). Internal sense of direction and landmark use in pigeons (Columba livia). J Comp Psychol, 119(3), 273–284.
Abstract: The relative importance of an internal sense of direction based on inertial cues and landmark piloting for small-scale navigation by White King pigeons (Columba livia) was investigated in an arena search task. Two groups of pigeons differed in whether they had access to visual cues outside the arena. In Experiment 1, pigeons were given experience with 2 different entrances and all pigeons transferred accurate searching to novel entrances. Explicit disorientation before entering did not affect accuracy. In Experiments 2-4, landmarks and inertial cues were put in conflict or tested 1 at a time. Pigeons tended to follow the landmarks in a conflict situation but could use an internal sense of direction to search when landmarks were unavailable.
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Kutsukake, N., & Castles, D. L. (2004). Reconciliation and post-conflict third-party affiliation among wild chimpanzees in the Mahale Mountains, Tanzania. Primates, 45(3), 157–165.
Abstract: This study investigated post-conflict (PC) behavior among wild chimpanzees (Pan troglodytes) of the M-group in the Mahale Mountains, Tanzania, and examined what types of behavior characterize the PC situation in this group, and the factors that influence the occurrence of PC affiliation between opponents soon after the end of an aggressive conflict (i.e., reconciliation). We found that the opponents affiliated selectively soon after the end of aggression, suggesting that reconciliation occurred in this group. The mean individual corrected conciliatory tendency (CCT) (Veenema et al. 1994 in Behav Proc 31:29-38) was 14.4%, which is similar to or lower than frequencies observed in studies of captive and wild chimpanzees. The valuable relationship hypothesis predicts that the CCT is higher among individuals who share valuable relationships (e.g., males or affiliative dyads) than among individuals who do not (e.g., females or less-associative dyads). However, the analysis based on data for aggression between unrelated individuals (including one incident between an adult and non-adult) and aggression between unrelated adults, did not uncover this difference. Affiliation by a previously uninvolved individual with the victim (“consolation”) and with the aggressor (“appeasement”) occurred more frequently following aggression than in the control condition. The results are compared with previous studies of captive and wild chimpanzees.
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Gruter, C. C. (2004). Conflict and postconflict behaviour in captive black-and-white snub-nosed monkeys (Rhinopithecus bieti). Primates, 45(3), 197–200.
Abstract: 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.
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Koski, S. E., Koops, K., & Sterck, E. H. M. (2007). Reconciliation, relationship quality, and postconflict anxiety: testing the integrated hypothesis in captive chimpanzees. Am. J. Primatol., 69(2), 158–172.
Abstract: Reconciliation is a conflict resolution mechanism that is common to many gregarious species with individualized societies. Reconciliation repairs the damaged relationship between the opponents and decreases postconflict (PC) anxiety. The “integrated hypothesis” links the quality of the opponents' relationship to PC anxiety, since it proposes that conflicts among partners with high relationship quality will yield high levels of PC anxiety, which in turn will lead to an increased likelihood of reconciliation. We tested the integrated hypothesis in captive chimpanzees (Pan troglodytes) in the Arnhem Zoo, The Netherlands. We applied the standard PC/matched control (MC) method. Our results mostly support the integrated hypothesis, in that more valuable and compatible partners (i.e., males and frequent groomers) reconciled more often than less valuable and weakly compatible partners (i.e., females and infrequent groomers). In addition, PC anxiety was higher after conflicts among males than among females. Emotional arousal thus appears to be a mediator facilitating reconciliation. However, in contrast to the predictions derived from the integrated hypothesis, PC anxiety appeared only in aggressees, and not in aggressors, of conflicts. This suggests that while relationship quality determines PC anxiety, it is dependent on the role of the participants in the conflict.
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Mallavarapu, S., Stoinski, T. S., Bloomsmith, M. A., & Maple, T. L. (2006). Postconflict behavior in captive western lowland gorillas (Gorilla gorilla gorilla). Am. J. Primatol., 68(8), 789–801.
Abstract: 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.
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