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Hostetter, A. B., Cantero, M., & Hopkins, W. D. (2001). Differential use of vocal and gestural communication by chimpanzees (Pan troglodytes) in response to the attentional status of a human (Homo sapiens). J. Comp. Psychol., 115(4), 337–343.
Abstract: This study examined the communicative behavior of 49 captive chimpanzees (Pan troglodytes), particularly their use of vocalizations, manual gestures, and other auditory- or tactile-based behaviors as a means of gaining an inattentive audience's attention. A human (Homo sapiens) experimenter held a banana while oriented either toward or away from the chimpanzee. The chimpanzees' behavior was recorded for 60 s. Chimpanzees emitted vocalizations faster and were more likely to produce vocalizations as their 1st communicative behavior when a human was oriented away from them. Chimpanzees used manual gestures more frequently and faster when the human was oriented toward them. These results replicate the findings of earlier studies on chimpanzee gestural communication and provide new information about the intentional and functional use of their vocalizations.
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Horowitz, A. C. (2003). Do humans ape? Or do apes human? Imitation and intention in humans (Homo sapiens) and other animals. J Comp Psychol, 117(3), 325–336.
Abstract: A. Whiten, D. M. Custance, J.-C. Gomez, P. Teixidor, and K. A. Bard (1996) tested chimpanzees' (Pan troglodytes) and human children's (Homo sapiens) skills at imitation with a 2-action test on an “artificial fruit.” Chimpanzees imitated to a restricted degree; children were more thoroughly imitative. Such results prompted some to assert that the difference in imitation indicates a difference in the subjects' understanding of the intentions of the demonstrator (M. Tomasello, 1996). In this experiment, 37 adult human subjects were tested with the artificial fruit. Far from being perfect imitators, the adults were less imitative than the children. These results cast doubt on the inference from imitative performance to an ability to understand others' intentions. The results also demonstrate how any test of imitation requires a control group and attention to the level of behavioral analysis.
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Whiten, A., Custance, D. M., Gomez, J. C., Teixidor, P., & Bard, K. A. (1996). Imitative learning of artificial fruit processing in children (Homo sapiens) and chimpanzees (Pan troglodytes). J Comp Psychol, 110(1), 3–14.
Abstract: Observational learning in chimpanzees and young children was investigated using an artificial fruit designed as an analog of natural foraging problems faced by primates. Each of 3 principal components could be removed in 2 alternative ways, demonstration of only one of which was watched by each subject. This permitted subsequent imitation by subjects to be distinguished from stimulus enhancement. Children aged 2-4 years evidenced imitation for 2 components, but also achieved demonstrated outcomes through their own techniques. Chimpanzees relied even more on their own techniques, but they did imitate elements of 1 component of the task. To our knowledge, this is the first experimental evidence of chimpanzee imitation in a functional task designed to simulate foraging behavior hypothesized to be transmitted culturally in the wild.
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Biro, D., Inoue-Nakamura, N., Tonooka, R., Yamakoshi, G., Sousa, C., & Matsuzawa, T. (2003). Cultural innovation and transmission of tool use in wild chimpanzees: evidence from field experiments. Anim. Cogn., 6(4), 213–223.
Abstract: Chimpanzees (Pan troglodytes) are the most proficient and versatile users of tools in the wild. How such skills become integrated into the behavioural repertoire of wild chimpanzee communities is investigated here by drawing together evidence from three complementary approaches in a group of oil-palm nut- ( Elaeis guineensis) cracking chimpanzees at Bossou, Guinea. First, extensive surveys of communities adjacent to Bossou have shown that population-specific details of tool use, such as the selection of species of nuts as targets for cracking, cannot be explained purely on the basis of ecological differences. Second, a 16-year longitudinal record tracing the development of nut-cracking in individual chimpanzees has highlighted the importance of a critical period for learning (3-5 years of age), while the similar learning contexts experienced by siblings have been found to result in near-perfect (13 out of 14 dyads) inter-sibling correspondence in laterality. Third, novel data from field experiments involving the introduction of unfamiliar species of nuts to the Bossou group illuminates key aspects of both cultural innovation and transmission. We show that responses of individuals toward the novel items differ markedly with age, with juveniles being the most likely to explore. Furthermore, subjects are highly specific in their selection of conspecifics as models for observation, attending to the nut-cracking activities of individuals in the same age group or older, but not younger than themselves. Together with the phenomenon of inter-community migration, these results demonstrate a mechanism for the emergence of culture in wild chimpanzees.
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Okamoto, S., Tomonaga, M., Ishii, K., Kawai, N., Tanaka, M., & Matsuzawa, T. (2002). An infant chimpanzee (Pan troglodytes) follows human gaze. Anim. Cogn., 5(2), 107–114.
Abstract: The ability of non-human primates to follow the gaze of other individuals has recently received much attention in comparative cognition. The aim of the present study was to investigate the emergence of this ability in a chimpanzee infant. The infant was trained to look at one of two objects, which an experimenter indicated by one of four different cue conditions: (1) tapping on the target object with a finger; (2) pointing to the target object with a finger; (3) gazing at the target object with head orientation; or (4) glancing at the target object without head orientation. The subject was given food rewards independently of its responses under the first three conditions, so that its responses to the objects were not influenced by the rewards. The glancing condition was tested occasionally, without any reinforcement. By the age of 13 months, the subject showed reliable following responses to the object that was indicated by the various cues, including glancing alone. Furthermore, additional tests clearly showed that the subject's performance was controlled by the “social” properties of the experimenter-given cues but not by the non-social, local-enhancing peripheral properties.
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de Waal, F. B., & Seres, M. (1997). Propagation of handclasp grooming among captive chimpanzees. Am. J. Primatol., 43(4), 339–346.
Abstract: A grooming posture previously reported for two wild chimpanzee (Pan troglodytes) communities developed spontaneously in a captive group of the same species. This offered a unique opportunity to follow the propagation of a new social custom. The posture consists of two partners grasping hands--either both right hands or both left hands--and raising the arms in an A-frame above their heads while mutually grooming with their free hands. The propagation of this pattern was followed over a 5 year period. In the beginning, handclasps were always initiated by the same adult female. This female initiated the posture mainly with her adult female kin. In subsequent years, these relatives became frequent participants in the posture with each other as well as with nonrelatives. Over the years the posture increased in frequency and duration and spread to the majority of adults and also to a few adolescents and older juveniles. The pattern persisted after removal of the apparent originator.
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Aureli, F., & de Waal, F. B. (1997). Inhibition of social behavior in chimpanzees under high-density conditions. Am. J. Primatol., 41(3), 213–228.
Abstract: This is the first study to investigate the short-term effects of high population density on captive chimpanzees (Pan troglodytes). Subjects of the study were 45 chimpanzees living in five different groups at the Yerkes Regional Primate Research Center. The groups were observed under two conditions: 1) when they had access to both the indoor and outdoor sections of their enclosures; 2) during cold days when they were locked into the indoor runs, which reduced the available space by more than half. Under the high-density condition, allogrooming and submissive greetings decreased, but juvenile play increased. Remarkably, the rate of various forms of agonistic behavior, such as aggression, bluff charge, bluff display, and hooting, occurred less frequently under the high-density condition. This general decrease in adult social activity, including agonistic behavior, can be interpreted as an inhibition strategy to reduce opportunities for conflict when interindividual distances are reduced. This strategy is probably effective only in the short run, however. Behavioral indicators of anxiety, such as rough scratching and yawning, showed elevated rates, suggesting increased social tension under the high-density condition.
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Cohen, J. (2007). Animal behavior. The world through a chimp's eyes (Vol. 316).
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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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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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