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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., & McGrew, W. C. (2001). Is this the first portrayal of tool use by a chimp? (Vol. 409).
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Whiten, A., & Boesch, C. (2001). The cultures of chimpanzees. Sci Am, 284(1), 60–67.
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Whiten, A., Goodall, J., McGrew, W. C., Nishida, T., Reynolds, V., Sugiyama, Y., et al. (1999). Cultures in chimpanzees. Nature, 399(6737), 682–685.
Abstract: As an increasing number of field studies of chimpanzees (Pan troglodytes) have achieved long-term status across Africa, differences in the behavioural repertoires described have become apparent that suggest there is significant cultural variation. Here we present a systematic synthesis of this information from the seven most long-term studies, which together have accumulated 151 years of chimpanzee observation. This comprehensive analysis reveals patterns of variation that are far more extensive than have previously been documented for any animal species except humans. We find that 39 different behaviour patterns, including tool usage, grooming and courtship behaviours, are customary or habitual in some communities but are absent in others where ecological explanations have been discounted. Among mammalian and avian species, cultural variation has previously been identified only for single behaviour patterns, such as the local dialects of song-birds. The extensive, multiple variations now documented for chimpanzees are thus without parallel. Moreover, the combined repertoire of these behaviour patterns in each chimpanzee community is itself highly distinctive, a phenomenon characteristic of human cultures but previously unrecognised in non-human species.
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Whiten, A. (1998). Imitation of the sequential structure of actions by chimpanzees (Pan troglodytes). J Comp Psychol, 112(3), 270–281.
Abstract: Imitation was studied experimentally by allowing chimpanzees (Pan troglodytes) to observe alternative patterns of actions for opening a specially designed “artificial fruit.” Like problematic foods primates deal with naturally, with the test fruit several defenses had to be removed to gain access to an edible core, but the sequential order and method of defense removal could be systematically varied. Each subject repeatedly observed 1 of 2 alternative techniques for removing each defense and 1 of 2 alternative sequential patterns of defense removal. Imitation of sequential organization emerged after repeated cycles of demonstration and attempts at opening the fruit. Imitation in chimpanzees may thus have some power to produce cultural convergence, counter to the supposition that individual learning processes corrupt copied actions. Imitation of sequential organization was accompanied by imitation of some aspects of the techniques that made up the sequence.
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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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Dunbar, R. I. M., McAdam, M. R., & O'connell, S. (2005). Mental rehearsal in great apes (Pan troglodytes and Pongo pygmaeus) and children. Behav. Process., 69(3), 323–330.
Abstract: The ability to rehearse possible future courses of action in the mind is an important feature of advanced social cognition in humans, and the “social brain” hypothesis implies that it might also be a feature of primate social cognition. We tested two chimpanzees, six orangutans and 63 children aged 3-7 years on a set of four puzzle boxes, half of which were presented with an opportunity to observe the box before being allowed to open it (“prior view”), the others being given without an opportunity to examine the boxes before handling them (“no prior view”). When learning effects are partialled out, puzzle boxes in the “prior view” condition were opened significantly faster than boxes given in the “no prior view” condition by the children, but not by either of the great apes. The three species differ significantly in the speed with which they opened boxes in the “no prior view” condition. The three species' performance on this task was a function of relative frontal lobe volume, suggesting that it may be possible to identify quantitative neuropsychological differences between species.
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Bard, K. A. (2007). Neonatal imitation in chimpanzees (Pan troglodytes) tested with two paradigms. Anim. Cogn., 10(2), 233–242.
Abstract: Primate species differ in their imitative performance, perhaps reflecting differences in imitative capacity. The developmentally earliest form of imitation in humans, neonatal imitation, occurs in early interactions with social partners, and may be a more accurate index of innate capacity than imitation of actions on objects, which requires more cognitive ability. This study assessed imitative capacity in five neonatal chimpanzees, within a narrow age range (7-15 days of age), by testing responses to facial and vocal actions with two different test paradigms (structured and communicative). Imitation of mouth opening was found in both paradigms. In the communicative paradigm, significant agreement was found between infant actions and demonstrations. Additionally, chimpanzees matched the sequence of three actions of the TC model, but only on the second demonstration. Newborn chimpanzees matched more modeled actions in the communicative test than in the structured paradigm. These performances of chimpanzees, at birth, are in agreement with the literature, supporting a conclusion that imitative capacity is not unique to the human species. Developmental histories must be more fully considered in the cross-species study of imitation, as there is a greater degree of innate imitative capacity than previously known. Socialization practices interact with innate and developing competencies to determine the outcome of imitation tests later in life.
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Tanaka, M. (2007). Recognition of pictorial representations by chimpanzees (Pan troglodytes). Anim. Cogn., 10(2), 169–179.
Abstract: In this study, I investigated chimpanzees' ability to recognize pictorial representations. Four adults and three juvenile chimpanzees were trained to choose images of photographs of flowers among 12 items belonging to four categories on a touch-sensitive monitor. As a generalization test, the following five types of images were presented: (1) novel photographs, (2) colored sketches (more realistic), (3) a colored clip art (cartoon-like images), (4) black-and-white line drawings, and (5) Kanji characters (as the control images). One adult and all three juvenile chimpanzees were able to choose any style of the nonphotographic images of flowers significantly above the chance level, whereas none could choose the correct Kanji characters corresponding to a flower significantly above the chance level. The other three adult chimpanzees' performance level did not exceed the chance level in terms of choosing nonphotographic images although they showed good transfer skills to novel photographs. The results revealed that not all chimpanzees could recognize pictures used by humans without training. The results also suggest “critical period” in acquisition of skill in recognizing pictures in chimpanzees. Only one adult chimpanzee, who had acquired skill in recognizing visual symbols, also recognized pictures aside from the juvenile chimpanzees. Her learning history might have aided her in acquiring this skill. The results of this study suggest a relationship between pictorial competence and symbolic one.
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Call, J. (2006). Inferences by exclusion in the great apes: the effect of age and species. Anim. Cogn., 9(4), 393–403.
Abstract: This study investigated the ability of chimpanzees, gorillas, orangutans, and bonobos to make inferences by exclusion using the procedure pioneered by Premack and Premack (Cognition 50:347-362, 1994) with chimpanzees. Thirty apes were presented with two different food items (banana vs. grape) on a platform and covered with identical containers. One of the items was removed from the container and placed between the two containers so that subjects could see it. After discarding this item, subjects could select between the two containers. In Experiment 1, apes preferentially selected the container that held the item that the experimenter had not discarded, especially if subjects saw the experimenter remove the item from the container (but without seeing the container empty). Experiment 3 in which the food was removed from one of the containers behind a barrier confirmed these results. In contrast, subjects performed at chance levels when a stimulus (colored plastic chip: Exp. 1; food item: Exp. 2 and Exp. 3) designated the item that had been removed. These results indicated that apes made inferences, not just learned to use a discriminative cue to avoid the empty container. Apes perceived and treated the item discarded by the experimenter as if it were the very one that had been hidden under the container. Results suggested a positive relationship between age and inferential ability independent of memory ability but no species differences.
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