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de Waal, F. B. M. (2005). A century of getting to know the chimpanzee. Nature, 437(7055), 56–59.
Abstract: A century of research on chimpanzees, both in their natural habitat and in captivity, has brought these apes socially, emotionally and mentally much closer to us. Parallels and homologues between chimpanzee and human behaviour range from tool-technology and cultural learning to power politics and intercommunity warfare. Few behavioural domains have remained untouched by this increased knowledge, which has dramatically challenged the way we view ourselves. The sequencing of the chimpanzee genome will no doubt bring more surprises and insights. Humans do occupy a special place among the primates, but this place increasingly has to be defined against a backdrop of substantial similarity.
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Whiten, A., Horner, V., & de Waal, F. B. M. (2005). Conformity to cultural norms of tool use in chimpanzees. Nature, 437(7059), 737–740.
Abstract: Rich circumstantial evidence suggests that the extensive behavioural diversity recorded in wild great apes reflects a complexity of cultural variation unmatched by species other than our own. However, the capacity for cultural transmission assumed by this interpretation has remained difficult to test rigorously in the field, where the scope for controlled experimentation is limited. Here we show that experimentally introduced technologies will spread within different ape communities. Unobserved by group mates, we first trained a high-ranking female from each of two groups of captive chimpanzees to adopt one of two different tool-use techniques for obtaining food from the same 'Pan-pipe' apparatus, then re-introduced each female to her respective group. All but two of 32 chimpanzees mastered the new technique under the influence of their local expert, whereas none did so in a third population lacking an expert. Most chimpanzees adopted the method seeded in their group, and these traditions continued to diverge over time. A subset of chimpanzees that discovered the alternative method nevertheless went on to match the predominant approach of their companions, showing a conformity bias that is regarded as a hallmark of human culture.
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Brosnan, S. F., & De Waal, F. B. M. (2003). Monkeys reject unequal pay. Nature, 425(6955), 297–299.
Abstract: During the evolution of cooperation it may have become critical for individuals to compare their own efforts and pay-offs with those of others. Negative reactions may occur when expectations are violated. One theory proposes that aversion to inequity can explain human cooperation within the bounds of the rational choice model, and may in fact be more inclusive than previous explanations. Although there exists substantial cultural variation in its particulars, this 'sense of fairness' is probably a human universal that has been shown to prevail in a wide variety of circumstances. However, we are not the only cooperative animals, hence inequity aversion may not be uniquely human. Many highly cooperative nonhuman species seem guided by a set of expectations about the outcome of cooperation and the division of resources. Here we demonstrate that a nonhuman primate, the brown capuchin monkey (Cebus apella), responds negatively to unequal reward distribution in exchanges with a human experimenter. Monkeys refused to participate if they witnessed a conspecific obtain a more attractive reward for equal effort, an effect amplified if the partner received such a reward without any effort at all. These reactions support an early evolutionary origin of inequity aversion.
Keywords: Aging; Animals; Cebus/*psychology; Choice Behavior; *Cooperative Behavior; Female; Male; *Reward; Social Justice
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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. M., & Davis, J. M. (2003). Capuchin cognitive ecology: cooperation based on projected returns. Neuropsychologia, 41(2), 221–228.
Abstract: Stable cooperation requires that each party's pay-offs exceed those available through individual action. The present experimental study on brown capuchin monkeys (Cebus apella) investigated if decisions about cooperation are (a) guided by the amount of competition expected to follow the cooperation, and (b) made instantaneously or only after a period of familiarization. Pairs of adult monkeys were presented with a mutualistic cooperative task with variable opportunities for resource monopolization (clumped versus dispersed rewards), and partner relationships (kin versus nonkin). After pre-training, each pair of monkeys (N=11) was subjected to six tests, consisting of 15 2 min trials each, with rewards available to both parties. Clumped reward distribution had an immediate negative effect on cooperation: this effect was visible right from the start, and remained visible even if clumped trials alternated with dispersed trials. The drop in cooperation was far more dramatic for nonkin than kin, which was explained by the tendency of dominant nonkin to claim more than half of the rewards under the clumped condition. The immediacy of responses suggests a decision-making process based on predicted outcome of cooperation. Decisions about cooperation thus take into account both the opportunity for and the likelihood of subsequent competition over the spoils.
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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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Bonnie, K. E., & de Waal, F. B. M. (2006). Affiliation promotes the transmission of a social custom: handclasp grooming among captive chimpanzees. Primates, 47(1), 27–34.
Abstract: Handclasp grooming is a unique social custom, known to occur regularly among some, but not all populations of chimpanzees (Pan troglodytes). As with other cultural behaviors, it is assumed that this distinctive grooming posture is learned socially by one individual from another. However, statistical comparisons among factors thought to influence how a behavior spreads within a group have never, to our knowledge, been conducted. In the present study, the origination and spread of handclasp grooming in a group of captive chimpanzees was followed throughout more than 1,500 h of observation over a period of 12 years. We report on the frequency, bout duration, and number and demography of performers throughout the study period, and compare these findings to those reported for wild populations. We predicted that dyads with strong affiliative ties, measured by time spent in proximity to and grooming one another, were likely to develop a handclasp grooming partnership during the study period. A quadratic assignment procedure was used to compare correlations among observed frequencies of grooming and proximity with handclasp grooming in all possible dyads within the group. As predicted, the formation of new handclasp grooming dyads was positively correlated with the rate of overall grooming and proximity within a dyad. In addition, in nearly all dyads formed, at least one individual had been previously observed to handclasp groom. We concluded that affiliation and individual experience determines the transmission of handclasp grooming among captive chimpanzees.
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Brosnan, S. F., & de Waal, F. B. M. (2005). Responses to a simple barter task in chimpanzees, Pan troglodytes. Primates, 46(3), 173–182.
Abstract: Chimpanzees (Pan troglodytes) frequently participate in social exchange involving multiple goods and services of variable value, yet they have not been tested in a formalized situation to see whether they can barter using multiple tokens and rewards. We set up a simple barter economy with two tokens and two associated rewards and tested chimpanzees on their ability to obtain rewards by returning the matching token in situations in which their access to tokens was unlimited or limited. Chimpanzees easily learned to associate value with the tokens, as expected, and did barter, but followed a simple strategy of favoring the higher-value token, regardless of the reward proffered, instead of a more complex but more effective strategy of returning the token that matched the reward. This response is similar to that shown by capuchin monkeys in our previous study. We speculate that this response, while not ideal, may be sufficient to allow for stability of the social exchange system in these primates, and that the importance of social barter to both species may have led to this convergence of strategies.
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Horner, V., Whiten, A., Flynn, E., & de Waal, F. B. M. (2006). Faithful replication of foraging techniques along cultural transmission chains by chimpanzees and children. Proc. Natl. Acad. Sci. U.S.A., 103(37), 13878–13883.
Abstract: Observational studies of wild chimpanzees (Pan troglodytes) have revealed population-specific differences in behavior, thought to represent cultural variation. Field studies have also reported behaviors indicative of cultural learning, such as close observation of adult skills by infants, and the use of similar foraging techniques within a population over many generations. Although experimental studies have shown that chimpanzees are able to learn complex behaviors by observation, it is unclear how closely these studies simulate the learning environment found in the wild. In the present study we have used a diffusion chain paradigm, whereby a behavior is passed from one individual to the next in a linear sequence in an attempt to simulate intergenerational transmission of a foraging skill. Using a powerful three-group, two-action methodology, we found that alternative methods used to obtain food from a foraging device (“lift door” versus “slide door”) were accurately transmitted along two chains of six and five chimpanzees, respectively, such that the last chimpanzee in the chain used the same method as the original trained model. The fidelity of transmission within each chain is remarkable given that several individuals in the no-model control group were able to discover either method by individual exploration. A comparative study with human children revealed similar results. This study is the first to experimentally demonstrate the linear transmission of alternative foraging techniques by non-human primates. Our results show that chimpanzees have a capacity to sustain local traditions across multiple simulated generations.
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de Waal, F. B. M., Dindo, M., Freeman, C. A., & Hall, M. J. (2005). The monkey in the mirror: hardly a stranger. Proc. Natl. Acad. Sci. U.S.A., 102(32), 11140–11147.
Abstract: It is widely assumed that monkeys see a stranger in the mirror, whereas apes and humans recognize themselves. In this study, we question the former assumption by using a detailed comparison of how monkeys respond to mirrors versus live individuals. Eight adult female and six adult male brown capuchin monkeys (Cebus apella) were exposed twice to three conditions: (i) a familiar same-sex partner, (ii) an unfamiliar same-sex partner, and (iii) a mirror. Females showed more eye contact and friendly behavior and fewer signs of anxiety in front of a mirror than they did when exposed to an unfamiliar partner. Males showed greater ambiguity, but they too reacted differently to mirrors and strangers. Discrimination between conditions was immediate, and blind coders were able to tell the difference between monkeys under the three conditions. Capuchins thus seem to recognize their reflection in the mirror as special, and they may not confuse it with an actual conspecific. Possibly, they reach a level of self-other distinction intermediate between seeing their mirror image as other and recognizing it as self.
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