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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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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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Hayashi, M. (2007). Stacking of blocks by chimpanzees: developmental processes and physical understanding. Anim. Cogn., 10(2), 89–103.
Abstract: The stacking-block task has been used to assess cognitive development in both humans and chimpanzees. The present study reports three aspects of stacking behavior in chimpanzees: spontaneous development, acquisition process following training, and physical understanding assessed through a cylindrical-block task. Over 3 years of longitudinal observation of block manipulation, one of three infant chimpanzees spontaneously started to stack up cubic blocks at the age of 2 years and 7 months. The other two infants began stacking up blocks at 3 years and 1 month, although only after the introduction of training by a human tester who rewarded stacking behavior. Cylindrical blocks were then introduced to assess physical understanding in object-object combinations in three infant (aged 3-4) and three adult chimpanzees. The flat surfaces of cylinders are suitable for stacking, while the rounded surface is not. Block manipulation was described using sequential codes and analyzed focusing on failure, cause, and solution in the task. Three of the six subjects (one infant and two adults) stacked up cylindrical blocks efficiently: frequently changing the cylinders' orientation without contacting the round side to other blocks. Rich experience in stacking cubes may facilitate subjects' stacking of novel, cylindrical shapes from the beginning. The other three subjects were less efficient in stacking cylinders and used variable strategies to achieve the goal. Nevertheless, they began to learn the effective way of stacking over the course of testing, after about 15 sessions (75 trials).
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Neiworth, J. J., Hassett, J. M., & Sylvester, C. J. (2007). Face processing in humans and new world monkeys: the influence of experiential and ecological factors. Anim. Cogn., 10(2), 125–134.
Abstract: This study tests whether the face-processing system of humans and a nonhuman primate species share characteristics that would allow for early and quick processing of socially salient stimuli: a sensitivity toward conspecific faces, a sensitivity toward highly practiced face stimuli, and an ability to generalize changes in the face that do not suggest a new identity, such as a face differently oriented. The look rates by adult tamarins and humans toward conspecific and other primate faces were examined to determine if these characteristics are shared. A visual paired comparison (VPC) task presented subjects with either a human face, chimpanzee face, tamarin face, or an object as a sample, and then a pair containing the previous stimulus and a novel stimulus was presented. The stimuli were either presented all in an upright orientation, or all in an inverted orientation. The novel stimulus in the pair was either an orientation change of the same face/object or a new example of the same type of face/object, and the stimuli were shown either in an upright orientation or in an inverted orientation. Preference to novelty scores revealed that humans attended most to novel individual human faces, and this effect decreased significantly if the stimuli were inverted. Tamarins showed preferential looking toward novel orientations of previously seen tamarin faces in the upright orientation, but not in an inverted orientation. Similarly, their preference to look longer at novel tamarin and human faces within the pair was reduced significantly with inverted stimuli. The results confirmed prior findings in humans that novel human faces generate more attention in the upright than in the inverted orientation. The monkeys also attended more to faces of conspecifics, but showed an inversion effect to orientation change in tamarin faces and to identity changes in tamarin and human faces. The results indicate configural processing restricted to particular kinds of primate faces by a New World monkey species, with configural processing influenced by life experience (human faces and tamarin faces) and specialized to process orientation changes specific to conspecific faces.
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Shoshani, J., Kupsky, W. J., & Marchant, G. H. (2006). Elephant brain. Part I: gross morphology, functions, comparative anatomy, and evolution. Brain Res Bull, 70(2), 124–157.
Abstract: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior.
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Hostetter, A. B., Russell, J. L., Freeman, H., & Hopkins, W. D. (2007). Now you see me, now you don't: evidence that chimpanzees understand the role of the eyes in attention. Anim. Cogn., 10(1), 55–62.
Abstract: Chimpanzees appear to understand something about the attentional states of others; in the present experiment, we investigated whether they understand that the attentional state of a human is based on eye gaze. In all, 116 adult chimpanzees were offered food by an experimenter who engaged in one of the four experimental manipulations: eyes closed, eyes open, hand over eyes, and hand over mouth. The communicative behavior of the chimpanzees was observed. More visible behaviors were produced when the experimenter's eyes were visible than when the experimenter's eyes were not visible. More vocalizations were produced when the experimenter's eyes were closed than when they were open, but there were no differences in other attention getting behaviors. There was no effect of age or rearing history. The results suggest that chimpanzees use the presence of the eyes as a cue that their visual gestures will be effective.
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