All investigated cases of habitual tool use in wild chimpanzees and capuchin monkeys include youngsters encountering durable artefacts, most often in a supportive social context. We propose that enduring artefacts associated with tool use, such as previously used tools, partly processed food items and residual material from previous activity, aid non-human primates to learn to use tools, and to develop expertise in their use, thus contributing to traditional technologies in non-humans. Therefore, social contributions to tool use can be considered as situated in the three dimensions of Euclidean space, and in the fourth dimension of time. This notion expands the contribution of social context to learning a skill beyond the immediate presence of a model nearby. We provide examples supporting this hypothesis from wild bearded capuchin monkeys and chimpanzees, and suggest avenues for future research.

Body mass is fundamental for understanding growth, health, and aspects of life history but records of body mass are rarely available for wild primates. We documented the body mass of all individuals in a group of wild bearded capuchin monkeys (Sapajus libidinosus) at annual intervals for seven consecutive years. Sexual dimorphism in body mass was more pronounced than reported in the literature for adults in this genus: females in our sample were relatively light (average 2.1 kg), while males had average body mass (3.5 kg). Three other notable differences between males and females were evident. First, males grew more rapidly and for a longer period than females. We estimate that males attained full body mass at 9.8 years of age and females at 7.5 years. Second, males showed greater inter-individual variability than females in growth rates and adult mass. Third, males gained about 20% above their baseline body mass upon becoming alpha, and lost that amount when they lost that status, but body mass in females was unrelated to social status. We also report preliminary data on mass and age of natal males at dispersal and mass and age at first reproduction for one female. The pattern of sexual dimorphism in ontogeny and inter-individual variability in body mass in bearded capuchins suggests different competitive risks in the two sexes commensurate with a mating system characterized by female choice of mates in multi-male, multi-female groups. Am. J. Primatol. 78:473-484, 2016. © 2015 Wiley Periodicals, Inc.

The habitual use of tools by wild capuchin monkeys presents a unique opportunity to study the maintenance and transmission of traditions. Young capuchins spend several years interacting with nuts before cracking them efficiently with stone tools. Using a two-observer method, we quantified the magnitude of the social influences that sustain this long period of practice. During five collection periods (over 26 months), one observer recorded the behavior of 16 immature monkeys, and another observer concurrently recorded behavior of group members in the focal monkey's vicinity. The two-observer method provides a means to quantify distinct social influences. Data show that immatures match the behavior of the adults in time and especially in space. The rate of manipulation of nuts by the immatures quadrupled when others in the group cracked and ate nuts, and immatures were ten times more likely to handle nuts and 40 times more likely to strike a nut with a stone when they themselves were near the anvils. Moreover, immature monkeys were three times more likely to be near an anvil when others were cracking. We suggest a model for social influence on nut-cracking development, based on two related processes: (1) social facilitation from observing group members engaged in nut-cracking, and (2) opportunity for practice provided by the anvils, hammer stones and nut shells available on and around the anvils. Nut-cracking activities by others support learning by drawing immatures to the anvils, where extended practice can take place, and by providing materials for practice at these places.

Culture extends biology in that the setting of development shapes the traditions that individuals learn, and over time, traditions evolve as occasional variations are learned by others. In humans, interactions with others impact the development of cognitive processes, such as sustained attention, that shape how individuals learn as well as what they learn. Thus, learning itself is impacted by culture. Here, we explore how social partners might shape the development of psychological processes impacting learning a tradition. We studied bearded capuchin monkeys learning a traditional tool-using skill, cracking nuts using stone hammers. Young monkeys practice components of cracking nuts with stones for years before achieving proficiency. We examined the time course of young monkeys' activity with nuts before, during, and following others' cracking nuts. Results demonstrate that the onset of others' cracking nuts immediately prompts young monkeys to start handling and percussing nuts, and they continue these activities while others are cracking. When others stop cracking nuts, young monkeys sustain the uncommon actions of percussing and striking nuts for shorter periods than the more common actions of handling nuts. We conclude that nut-cracking by adults can promote the development of sustained attention for the critical but less common actions that young monkeys must practice to learn this traditional skill. This work suggests that in nonhuman species, as in humans, socially specified settings of development impact learning processes as well as learning outcomes. Nonhumans, like humans, may be culturally variable learners.T raditions are behaviors shared among members of a group that are reliably learned by new individuals, in part, with social support (1). The concept of tradition highlights behavioral consistency over time and across individuals. Here, we emphasize the origin of traditions in learning. Learned behavior is one manifestation of developmental plasticity, and generation of novel behavior is another. Novel behavioral variants may be learned by others, thus modifying existing traditions or leading to new ones (2). In this way, developmental plasticity enables heritable modification of behavior through learning, with the result that culture may extend biology (3, 4). Traditions appear to be widespread in the animal kingdom (5-7), leading theorists to propose that cultural evolution, rather than being restricted to recent human history, has general evolutionary significance (8, 9). Apes (6, 10) and monkeys provide several examples of traditions. Among monkeys, for example, vervet monkeys learn their mothers' food-processing techniques (11) and white-faced capuchin monkeys learn odd social games, such as poking a finger in another monkey's eye, that become established intermittently in groups (2, 12).Given the essential role of learning in traditions, cognitive processes associated with learning that show developmental plasticity themselves, should be incorporated into our understanding of how culture can...