Boesch, C. (2002). Cooperative hunting roles among Tai chimpanzees. Human Nature-an Interdisciplinary Biosocial Perspective, 13(1), 27-46.

All known chimpanzee populations have been observed to hunt small mammals for meat. Detailed observations have shown, however, that hunting strategies differ considerably between populations, with some merely collecting prey that happens to pass by while others hunt in coordinated groups to chase fast- moving prey. Of all known populations, Tai chimpanzees exhibit the highest level of cooperation when hunting. Some of the group hunting roles require elaborate coordination with other hunters as well as precise anticipation of the movements of the prey. The meat-sharing rules observed in this community guarantee the largest share of the meat to hunters who perform the most important roles leading to a capture. The learning time of such hunting roles is sometimes especially long. Tai chimpanzee males begin hunting monkeys at about age 10. The hunters' progress in learning the more sophisticated hunting roles is clearly correlated with age; only after 20 years of practice are they able to perform them reliably. This lengthy learning period has also been shown in some hunter-gatherer societies and confirms the special challenge that hunting represents.

Boysen, S.T. and Himes, G.T. (1999) Current issues and emerging theories in animal cognition. Annual Review of Psychology, 50, 683-705.

Comparative cognition is an emerging interdisciplinary field with contributions from comparative psychology, cognitive/experimental and developmental psychology, animal learning, and ethology, and is poised to move toward greater understanding of animal and human information-processing, reasoning, memory, and the phylogenetic emergence of mind. This chapter highlights some current issues and discusses four areas within comparative cognition that are yielding new approaches and hypotheses for studying basic conceptual capacities in nonhuman species. These include studies of imitation, tool use, mirror self- recognition, and the potential for attribution of mental states by nonhuman animals. Though a very old question in psychology, the study of imitation continues to provide new avenues for examining the complex relationships among and between the levels of imitative behaviors exhibited by many species. Similarly, recent work in animal tool use, mirror self-recognition (with all its contentious issues), and recent attempts to empirically study the potential for attributional capacities in nonhumans, all continue to provide fresh insights and novel paradigms for addressing the defining characteristics of these complex phenomena.

Brauer, J., Call, J., & Tomasello, M. (2005). All great ape species follow gaze to distant locations and around barriers. Journal of Comparative Psychology, 119(2), 145-154.

Following the gaze direction of conspecifics is an adaptive skill that enables individuals to obtain useful information about the location of food, predators, and group mates. In the current study, the authors compared the gaze-following skills of all 4 great ape species. In the 1st experiment, a human either looked to the ceiling or looked straight ahead. Individuals from all species reliably followed the human's gaze direction and sometimes even checked back when they found no target. In a 2nd experiment, the human looked behind some kind of barrier. Results showed that individuals from all species reliably put themselves in places from which they could see what the experimenter was looking at behind the barrier. These results support the hypothesis that great apes do not just orient to a target that another is oriented to, but they actually attempt to take the visual perspective of the other.

Call, J. (2001). Chimpanzee social cognition. Trends in Cognitive Sciences, 5(9), 388-393.

In the late 1970s, Premack and Woodruff asked whether chimpanzees had a theory of mind. The answer to this question has remained elusive. Whereas some authors argue that chimpanzees are capable of mental state attribution, others maintain that they simply learn certain cues in ertain situations. Recent studies challenge both views. On the one hand, chimpanzees know much more about seeing than cue-based explanations suggest; on the other hand, this knowledge does not necessarily entail understanding of the mental states of others. The hypothesis I put forward here is that chimpanzees learn cues in social situations but that they are also capable of knowledge abstraction to solve novel problems.

Cantalupo, C., & Hopkins, W. D. (2001). Asymmetric Broca's area in great apes - A region of the ape brain is uncannily similar to one linked with speech in humans. Nature, 414(6863), 505-505.

Brodmann's area 44 delineates part of Broca's area within the inferior frontal gyrus of the human brain and is a critical region for speech production, being larger in the left hemisphere than in the right - an asymmetry that has been correlated with language dominance. Here we show that there is a similar asymmetry in this area, also with left-hemisphere dominance, in three great ape species (Pan troglodytes, Pan paniscus and Gorilla gorilla). Our findings suggest that the neuroanatomical substrates for left-hemisphere dominance in speech production were evident at least five million years ago and are not unique to hominid evolution.

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Crockford et al. (2004). Wild chimpanzees produce group-specific calls: a case for vocal learning? Ethology, 110(3), 221-243.

Vocal learning, where animals can modify the structure of their vocalizations as a result of experience, has been found in a range of birds and mammals. Although vocal learning is a fundamental aspect of developing spoken language, there is as yet little evidence that vocal learning occurs in primates. Here we examine whether vocal learning may occur in chimpanzees. We analysed whether wild male chimpanzees, Pan troglodytes verus, of four communities living in a similar habitat in the Tai Forest, Cote d'Ivoire, developed community specific pant hoots. If so, we expected males of three contiguous communities to have distinct pant hoots, while pant hoots of males from a fourth, distant community, located 70 km away, should only differ from those of the contiguous communities by chance. Our analysis confirmed these expectations. In addition, the acoustic distances between the pant hoots of pairs of individuals did not correlate with the genetic relatedness of those pairs, where genetic relatedness was determined using nuclear DNA analysis. Thus, neither habitat nor genetic differences accounted for the observation that there were acoustic differences in the pant hoot structure of males living in neighbouring communities, but not in those of males from a distant community. This suggests that chimpanzees may actively modify pant hoots to be different from their neighbours, providing support for the vocal learning hypothesis.

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Hopkins, W. D., Taglialatela, J. P., & Leavens, D. A. (2007). Chimpanzees differentially produce novel vocalizations to capture the attention of a human. Animal Behaviour, 73, 281-286.

Chimpanzees, Pan troglodytes, produce numerous species-atypical signals when raised in captivity. We examined contextual elements of the use of two of these vocal signals, the 'raspberry' and the extended grunt. Our results demonstrate that these vocalizations are not elicited by the presence of food, but instead function as attention-getting signals. These findings reveal a heretofore underappreciated category of animal signals: attention-getting sounds produced in novel environmental circumstances. The invention and use of species-atypical signals, considered in relation to group differences in signalling repertoires in apes in their natural habitats, may index a generative capacity in these hominoid species without obvious corollary in other primate species.

Melis, A. P., Hare, B., & Tomasello, M. (2006). Chimpanzees recruit the best collaborators. Science, 311(5765), 1297-1300.

Humans collaborate with non-kin in special ways, but the evolutionary foundations of these collaborative skills remain unclear. We presented chimpanzees with collaboration problems in which they had to decide when to recruit a partner and which potential partner to recruit. In an initial study, individuals recruited a collaborator only when solving the problem required collaboration. In a second study, individuals recruited the more effective of two partners on the basis of their experience with each of them on a previous day. Therefore, recognizing when collaboration is necessary and determining who is the best collaborative partner are skills shared by both chimpanzees and humans, so such skills may have been present in their common ancestor before humans evolved their own complex forms of collaboration.

Menzel, E.W., Savage-Rumbaugh, E.S. and Lawson, J. (1985) Chimpanzee (Pan-troglodytes) spatial problem-solving with the use of mirrors and televised equivalents of mirrors. Journal of Comparative Psychology, 99, 211-217.

Two adult male chimpanzees reached through a hole in the wall of their home cage and, by tracking the images of their hands and of an otherwise hidden target object in a mirror or closed circuit tevelvision picture, moved their hands in whichever direction was necessary to make contact with the target object. They discriminated between tive video images and tapes and performed effectively when the target objects were presented in novels locations and when the video picture was presented at random in different orientations. There was thus no consistent relation between the location of images on the monitor and the location of their real-world counterparts. Comparable performances in monkeys and nonprimates seem unlikely.

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Pika, S., & Mitani, J. (2006). Referential gestural communication in wild chimpanzees (Pan troglodytes). Current Biology, 16(6), R191-R192.

Humans commonly use referential gestures, for example pointing, which direct the attention of recipients to particular aspects of the environment . The use of these gestures has been linked with cognitive capacities such as mental state attribution 2 and 3 because the recipient must infer the signaler's meaning. In our closest living relatives, the non-human primates, referential gestures have been reported only in captive chimpanzees interacting with their human experimenters and human-raised or language-trained apes . Here we provide the first evidence for the widespread use of a referential gesture by wild chimpanzees (Pan troglodytes)







Pollard, K. S., Salama, S. R., Lambert, N., Lambot, M. A., Coppens, S., Pedersen, J. S., et al. (2006). An RNA gene expressed during cortical development evolved rapidly in humans. Nature, 443(7108), 167-172.

The developmental and evolutionary mechanisms behind the emergence of human-specific brain features remain largely unknown. However, the recent ability to compare our genome to that of our closest relative, the chimpanzee, provides new avenues to link genetic and phenotypic changes in the evolution of the human brain. We devised a ranking of regions in the human genome that show significant evolutionary acceleration. Here we report that the most dramatic of these 'human accelerated regions', HAR1, is part of a novel RNA gene (HAR1F) that is expressed specifically in Cajal Retzius neurons in the developing human neocortex from 7 to 19 gestational weeks, a crucial period for cortical neuron specification and migration. HAR1F is co-expressed with reelin, a product of Cajal - Retzius neurons that is of fundamental importance in specifying the six-layer structure of the human cortex. HAR1 and the other human accelerated regions provide new candidates in the search for uniquely human biology.

Ponting, C., & Jackson, A. P. (2005). Evolution of primary microcephaly genes and the enlargement of primate brains. Current Opinion in Genetics & Development, 15(3), 241-248.

Brain size, in relation to body size, has varied markedly during the evolution of mammals. In particular, a large cerebral cortex is a feature that distinguishes humans from our fellow primates. Such anatomical changes must have a basis in genetic alterations, but the molecular processes involved have yet to be defined. However, recent advances from the cloning of two human disease genes promise to make inroads in this important area. Microcephalin (MCPH1) and Abnormal spindle-like microcephaly associated (ASPM) are genes mutated in primary microcephaly, a human neurodevelopmental disorder. In this 'atavistic' condition, brain size is reduced in volume to a size comparable with that of early hominids. Hence, it has been proposed that these genes evolved adaptively with increasing primate brain size. Subsequent studies have lent weight to this hypothesis by showing that both genes have undergone positive selection during great ape evolution. Further functional characterisation of their proteins will contribute to an understanding of the molecular and evolutionary processes that have determined human brain size.

Rivas, E. (2005). Recent use of signs by chimpanzees (Pan troglodytes) in interactions with humans. Journal of Comparative Psychology, 119(4), 404-417.

In light of the controversy about the linguistic properties of chimpanzee signing behavior, the recent sign use of 5 chimpanzees (Pan troglodytes) with long histories of sign use was analyzed while they interacted with longtime human companions. Four corpora from 1992 to 1999 consisting of 3,448 sign utterances were examined. The chimpanzees predominantly used object and action signs. There was no evidence for semantic or syntactic structure in combinations of signs. Longer combinations showed repetition and stringing of object and action signs. The chimpanzees mostly signed with an acquisitive motivation. Requests for objects and actions were the predominant communicative intentions of the sign utterances, though naming and answering also occurred. This recent sign use shows multiple differences with (early) human language.

Tomasello, M., Carpenter, M., Call, J., Behne, T., & Moll, H. (2005). Understanding and sharing intentions: The origins of cultural cognition. Behavioral and Brain Sciences, 28(5), 675-+.

We propose that the crucial difference between human cognition and that of other species is the ability to participate with others in collaborative activities with shared goals and intentions: shared intentionality. Participation in such activities requires not only especially powerful forms of intention reading and cultural learning, but also a unique motivation to share psychological states with others and unique forms of cognitive representation for doing so. The result of participating in these activities is species-unique forms of cultural cognition and evolution, enabling everything from the creation and use of linguistic symbols to the construction of social norms and individual beliefs to the establishment of social institutions. In support of this proposal we argue and present evidence that great apes (and some children with autism) understand the basics of intentional action, but they still do not participate in activities involving joint intentions and attention (shared intentionality). Human children's skills of shared intentionality develop gradually during the first 14 months of life as two ontogenetic pathways intertwine: (1) the general ape line of understanding others as animate, goal-directed, and intentional agents; and (2) a species-unique motivation to share emotions, experience, and activities with other persons. The developmental outcome is children's ability to construct dialogic cognitive representations, which enable them to participate in earnest in the collectivity that is human cognition.

Warneken, F., & Tomasello, M. (2006). Altruistic helping in human infants and young chimpanzees. Science, 311(5765), 1301-1303.

Human beings routinely help others to achieve their goals, even when the helper receives no immediate benefit and the person helped is a stranger. Such altruistic behaviors (toward non-kin) are extremely rare evolutionarily, with some theorists even proposing that they are uniquely human. Here we show that human children as young as 18 months of age (prelinguistic or just-linguistic) quite readily help others to achieve their goals in a variety of different situations. This requires both an understanding of others' goals and an altruistic motivation to help. In addition, we demonstrate similar though less robust skills and motivations in three young chimpanzees.

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Wich, S. A., & de Vries, H. (2006). Male monkeys remember which group members have given alarm calls. Proceedings of the Royal Society B-Biological Sciences, 273(1587), 735-740.

Primates give alarm calls in response to the presence of predators. In some species, such as the Thomas langur (Presbytis thomasi), males only emit alarm calls if there is an audience. An unanswered question is whether the audience's behaviour influences how long the male will continue his alarm calling. We tested three hypotheses that might explain the alarm calling duration of male Thomas langurs: the fatigue, group size and group member behaviour hypotheses. Fatigue and group size did not influence male alarm calling duration. We found that males only ceased calling shortly after all individuals in his group had given at least one alarm call. This shows that males keep track of and thus remember which group members have called.

Wrangham, R. W., Wilson, M. L., & Muller, M. N. (2006). Comparative rates of violence in chimpanzees and humans. Primates, 47(1), 14-26.

This paper tests the proposal that chimpanzees (Pan troglodytes) and humans have similar rates of death from intraspecific aggression, whereas chimpanzees have higher rates of non-lethal physical attack (Boehm 1999, Hierarchy in the forest: the evolution of egalitarian behavior. Harvard University Press). First, we assembled data on lethal aggression from long-term studies of nine communities of chimpanzees living in five populations. We calculated rates of death from intraspecific aggression both within and between communities. Variation among communities in mortality rates from aggression was high, and rates of death from intercommunity and intracommunity aggression were not correlated. Estimates for average rates of lethal violence for chimpanzees proved to be similar to average rate for subsistence societies of hunter-gatherers and farmers. Second, we compared rates of non-lethal physical aggression for two populations of chimpanzees and one population of recently settled hunter-gatherers. Chimpanzees had rates of aggression between two and three orders of magnitude higher than humans. These preliminary data support Boehm's hypothesis.