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.

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.

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.

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.

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.