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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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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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Preston, S. D., & de Waal, F. B. M. (2002). Empathy: Its ultimate and proximate bases. Behav Brain Sci, 25(1), 1–20; discussion 20–71.
Abstract: There is disagreement in the literature about the exact nature of the phenomenon of empathy. There are emotional, cognitive, and conditioning views, applying in varying degrees across species. An adequate description of the ultimate and proximate mechanism can integrate these views. Proximately, the perception of an object's state activates the subject's corresponding representations, which in turn activate somatic and autonomic responses. This mechanism supports basic behaviors (e.g., alarm, social facilitation, vicariousness of emotions, mother-infant responsiveness, and the modeling of competitors and predators) that are crucial for the reproductive success of animals living in groups. The Perception-Action Model (PAM), together with an understanding of how representations change with experience, can explain the major empirical effects in the literature (similarity, familiarity, past experience, explicit teaching, and salience). It can also predict a variety of empathy disorders. The interaction between the PAM and prefrontal functioning can also explain different levels of empathy across species and age groups. This view can advance our evolutionary understanding of empathy beyond inclusive fitness and reciprocal altruism and can explain different levels of empathy across individuals, species, stages of development, and situations.
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Brosnan, S. F., Schiff, H. C., & de Waal, F. B. M. (2005). Tolerance for inequity may increase with social closeness in chimpanzees. Proc Biol Sci, 272(1560), 253–258.
Abstract: Economic decision-making depends on our social environment. Humans tend to respond differently to inequity in close relationships, yet we know little about the potential for such variation in other species. We examine responses to inequity in several groups of chimpanzees (Pan troglodytes) in a paradigm similar to that used previously in capuchin monkeys (Cebus apella). We demonstrate that, like capuchin monkeys, chimpanzees show a response to inequity of rewards that is based upon the partner receiving the reward rather than the presence of the reward alone. However, we also found a great amount of variation between groups tested, indicating that chimpanzees, like people, respond to inequity in a variable manner, which we speculate could be caused by such variables as group size, the social closeness of the group (as reflected in length of time that the group has been together) and group-specific traditions.
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de Waal, F. B. M. (2004). Peace lessons from an unlikely source. PLoS. Biol., 2(4), E101.
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de Waal, F. B. M. (2003). Darwin's legacy and the study of primate visual communication. Ann N Y Acad Sci, 1000, 7–31.
Abstract: After Charles Darwin's The Expression of the Emotions in Man and Animals, published in 1872, we had to wait 60 years before the theme of animal expressions was picked up by another astute observer. In 1935, Nadezhda Ladygina-Kohts published a detailed comparison of the expressive behavior of a juvenile chimpanzee and of her own child. After Kohts, we had to wait until the 1960s for modern ethological analyses of primate facial and gestural communication. Again, the focus was on the chimpanzee, but ethograms on other primates appeared as well. Our understanding of the range of expressions in other primates is at present far more advanced than that in Darwin's time. A strong social component has been added: instead of focusing on the expressions per se, they are now often classified according to the social situations in which they typically occur. Initially, quantitative analyses were sequential (i.e., concerned with temporal associations between behavior patterns), and they avoided the language of emotions. I will discuss some of this early work, including my own on the communicative repertoire of the bonobo, a close relative of the chimpanzee (and ourselves). I will provide concrete examples to make the point that there is a much richer matrix of contexts possible than the common behavioral categories of aggression, sex, fear, play, and so on. Primate signaling is a form of negotiation, and previous classifications have ignored the specifics of what animals try to achieve with their exchanges. There is also increasing evidence for signal conventionalization in primates, especially the apes, in both captivity and the field. This process results in group-specific or “cultural” communication patterns.
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Parr, L. A., Matheson, M. D., Bernstein, I. S., & De Waal, F. B. M. (1997). Grooming down the hierarchy: allogrooming in captive brown capuchin monkeys, Cebus apella. Anim. Behav., 54(2), 361–367.
Abstract: Observations of captive female brown capuchin monkeys in five groups revealed that grooming is primarily the occupation of dominant females at both the individual and dyadic levels. When categorized according to rank class, alpha females were the only class to perform significantly more grooming than they received. These results are inconsistent with reports on vervets, baboons and macaques, and suggest that grooming in capuchin monkeys may have different functions from those reported for cercopithecine primates. A dyadic analysis revealed, however, that grooming occurred more often between closely ranked females, similar to what is seen in several Old World monkey species. Therefore, some aspects of grooming in capuchins are similar to that seen in Old World monkeys, but the way they distribute grooming is different, which may prompt a re-evaluation of current theories regarding the social function of allogrooming in non-human primates.
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Weaver, A., & de Waal, F. B. M. (2002). An index of relationship quality based on attachment theory. J Comp Psychol, 116(1), 93–106.
Abstract: Two measures are reported of the nature or quality of a mother-offspring (MO) relationship during development using brown capuchin monkeys (Cebus apella) as models. One is a qualitative classification of MO relationships as secure, resistant, or avoidant attachments. The other is an empirical ratio of relative affiliation to agonism called the MO relationship quality, or MORQ, Index. The two methods tapped similar relationship features so relationships high or low of a median split of MORQ values were heuristically labeled secure (n = 22) or insecure (n = 16), respectively. A comparison revealed extensive behavioral differences between secure and insecure MO relationships and suggested MORQ provided an objective, continuous measure of attachment security.
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