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.

Abstract: The transmission of tool use is a rare event in monkeys. Such an event arose in a group of semi-free-ranging Tonkean macaques (Macaca tonkeana) in which leaning a pole against the park's fence (branch leaning) appeared and spread to several males. This prompted us to test individual and social learning of this behavior in seven young males. In the first experiment, three males learned individually to obtain a food reward using a wooden pole as a climbing tool. They began using the pole to retrieve the reward only when they could alternatively experience acting on the object and reaching the target. In a second experiment, we first tested whether four other subjects could learn branch leaning after having observed a group-mate performing the task. Despite repeated opportunities to observe the demonstrator, they did not learn to use the pole as a tool. Hence we exposed the latter subjects to individual learning trials and they succeeded in the task. Tool use was not transmitted in the experimental situation, which contrasts with observations in the park. We can conclude that the subjects were not able to recognize the target as such. It is possible that they recognized it and learned the task individually when we alternated the opportunity to act upon the object and to reach the reward. This suggests that these macaques could then have associated the action they exercised upon the pole and the use of the pole as a means to reach the reward.

Abstract: In the local population of kea in Mount Cook Village, New Zealand, some keas open the lids of rubbish bins with their bill to obtain food scraps within. We investigated the extent to which this innovation has spread in the local population, and what factors limit the acquisition of bin opening. Only five males of 36 individually recognised birds were observed to have performed successful bin opening. With one exception there were always other keas present, watching successful bin opening. Seventeen additional individuals were seen to have benefitted from lid opening. Their foraging success was less than that of the bin openers. Social status of bin openers did not differ from scrounging males. Among the individuals that were regularly seen at the site of the bins but were not successful in bin opening, social status and the ratio of feeding directly from open bins correlated with the amount of opening attempts. We conclude that scrounging facilitated certain behavioural aspects of bin opening rather than inhibiting them. The fact that only 9% of opening attempts were successful, and the long period of time required to increase efficiency in lid opening shows that mainly individual experience, and to a lesser extent insight and social learning, play key roles in acquisition of the opening technique. The results indicate that the spread of innovative solutions of challenging mechanical problems in animals may be restricted to only a few individuals.

Abstract: This study focuses on the development of spontaneous object manipulation in three infant chimpanzees during their first 2 years of life. The three infants were raised by their biological mothers who lived among a group of chimpanzees. A human tester conducted a series of cognitive tests in a triadic situation where mothers collaborated with the researcher during the testing of the infants. Four tasks were presented, taken from normative studies of cognitive development of Japanese infants: inserting objects into corresponding holes in a box, seriating nesting cups, inserting variously shaped objects into corresponding holes in a template, and stacking up wooden blocks. The mothers had already acquired skills to perform these manipulation tasks. The infants were free to observe the mothers' manipulative behavior from immediately after birth. We focused on object-object combinations that were made spontaneously by the infant chimpanzees, without providing food reinforcement for any specific behavior that the infants performed. The three main findings can be summarized as follows. First, there was precocious appearance of object-object combination in infant chimpanzees: the age of onset (8-11 months) was comparable to that in humans (around 10 months old). Second, object-object combinations in chimpanzees remained at a low frequency between 11 and 16 months, then increased dramatically at the age of approximately 1.5 years. At the same time, the accuracy of these object-object combinations also increased. Third, chimpanzee infants showed inserting behavior frequently and from an early age but they did not exhibit stacking behavior during their first 2 years of life, in clear contrast to human data.

Abstract: This article explores the maternal role in the acquisition of tool-use behaviours by infant chimpanzees ( Pan troglodytes). A honey-fishing task, simulating ant/termite fishing found in the wild, was introduced to three dyads of experienced mother and naive infant chimpanzees. Four fishing sites and eight sets of 20 objects to be used as tools, not all appropriate, were available. Two of the mothers constantly performed the task, using primarily two kinds of tools; the three infants observed them. The infants, regardless of the amount of time spent observing, successfully performed the task around the age of 20-22 months, which is earlier than has been recorded in the wild. Two of the infants used the same types of tools that the adults predominantly used, suggesting that tool selectivity is transmitted. The results also show that adults are tolerant of infants, even if unrelated; infants were sometimes permitted to lick the tools, or were given the tools, usually without honey, as well as permitted to observe the adult performances closely.