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de Waal, F. B. (1997). Food transfers through mesh in brown capuchins. J Comp Psychol, 111(4), 370–378.
Abstract: Capuchin monkeys (Cebus apella) share food even if their partner is behind a mesh restraint. Pairs of adult capuchins were moved into a test chamber in which 1 monkey received cucumber pieces for 20 min and the other received apple slices during the following 20 min. Tolerant transfers of food occurred reciprocally among females: The rate of transfer from Female B to A in the second test phase varied with the rate from Female A to B in the first test phase. Several social mechanisms may explain this reciprocity. Whereas this study does not contradict cognitively complex explanations (e.g., mental record keeping of given and received food), the results are consistent with a rather simple explanation: that food sharing reflects a combination of affiliative tendency and high tolerance. The study suggests that sharing mechanisms may be different for adult male capuchins, with males sharing food more readily and less discriminatingly than females.
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Manson, J. H., Perry, S., & Stahl, D. (2005). Reconciliation in wild white-faced capuchins (Cebus capucinus). Am. J. Primatol., 65(3), 205–219.
Abstract: 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.
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de Waal, F. B., & Berger, M. L. (2000). Payment for labour in monkeys. Nature, 404(6778), 563.
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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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Weaver, A., & de Waal, F. B. M. (2003). The mother-offspring relationship as a template in social development: reconciliation in captive brown capuchins (Cebus apella). J Comp Psychol, 117(1), 101–110.
Abstract: Mother-offspring (MO) relationship quality was investigated to determine its influence on the development of reconciliation--affiliation between opponents shortly after a fight--because it influenceswhat distressed youngsters learn about calming down. Data were longitudinal and cross-sectional observational samples of 38 MO pairs of monkeys across 24 months. An MO relationship quality index (RQI) classified each pair as secure or insecure. Reconciliation emerged in infancy.Secure youngsters had an appeasing conciliatory style, and insecure youngsters had an agitated conciliatory style. Conclusions are that reconciliation develops from the attachment behavior system and MO RQI is related to the particular conciliatory style youngsters develop by affecting how aroused they are by conflict and the subsequent socializing they seek to calm down.
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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.
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Brosnan, S. F., Freeman, C., & De Waal, F. B. M. (2006). Partner's behavior, not reward distribution, determines success in an unequal cooperative task in capuchin monkeys. Am. J. Primatol., 68(7), 713–724.
Abstract: It was recently demonstrated that capuchin monkeys notice and respond to distributional inequity, a trait that has been proposed to support the evolution of cooperation in the human species. However, it is unknown how capuchins react to inequitable rewards in an unrestricted cooperative paradigm in which they may freely choose both whether to participate and, within the bounds of their partner's behavior, which reward they will receive for their participation. We tested capuchin monkeys with such a design, using a cooperative barpull, which has been used with great success in the past. Contrary to our expectations, the equity of the reward distribution did not affect success or pulling behavior. However, the behavior of the partner in an unequal situation did affect overall success rates: pairs that had a tendency to alternate which individual received the higher-value food in unequal reward situations were more than twice as successful in obtaining rewards than pairs in which one individual dominated the higher-value food. This ability to equitably distribute rewards in inherently biased cooperative situations has profound implications for activities such as group hunts, in which multiple individuals work together for a single, monopolizable reward.
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Brosnan, S. F., & de Waal, F. B. M. (2004). Socially learned preferences for differentially rewarded tokens in the brown capuchin monkey (Cebus apella). J Comp Psychol, 118(2), 133–139.
Abstract: Social learning is assumed to underlie traditions, yet evidence indicating social learning in capuchin monkeys (Cebus apella), which exhibit traditions, is sparse. The authors tested capuchins for their ability to learn the value of novel tokens using a previously familiar token-exchange economy. Capuchins change their preferences in favor of a token worth a high-value food reward after watching a conspecific model exchange 2 differentially rewarded tokens, yet they fail to develop a similar preference after watching tokens paired with foods in the absence of a conspecific model. They also fail to learn that the value of familiar tokens has changed. Information about token value is available in all situations, but capuchins seem to pay more attention in a social situation involving novel tokens.
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Brosnan, S. F., & de Waal, F. B. M. (2004). A concept of value during experimental exchange in brown capuchin monkeys, Cebus apella. Folia Primatol (Basel), 75(5), 317–330.
Abstract: We evaluated the response of brown capuchin monkeys to two differentially valued tokens in an experimental exchange situation akin to a simple barter. Monkeys were given a series of three tests to evaluate their ability to associate tokens with food, then their responses were examined in a barter situation in which tokens were either limited or unlimited. Capuchins did not perform barter in the typical sense, returning the tokens which were associated with the reward. However, females, but not males, showed a different response, preferring the higher-value token. This may indicate that they learned to prefer one token over the other rather than to associate the tokens with their specific rewards. This sex difference parallels previous findings of greater reciprocity in female brown capuchins than in males.
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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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