We describe our recent studies of imitation and cultural transmission in chimpanzees and children, which question late twentieth-century characterizations of children as imitators, but chimpanzees as emulators. As emulation entails learning only about the results of others' actions, it has been thought to curtail any capacity to sustain cultures. Recent chimpanzee diffusion experiments have by contrast documented a significant capacity for copying local behavioural traditions. Additionally, in recent 'ghost' experiments with no model visible, chimpanzees failed to replicate the object movements on which emulation is supposed to focus. We conclude that chimpanzees rely more on imitation and have greater cultural capacities than previously acknowledged. However, we also find that they selectively apply a range of social learning processes that include emulation. Recent studies demonstrating surprisingly unselective 'over-imitation' in children suggest that children's propensity to imitate has been underestimated too. We discuss the implications of these developments for the nature of social learning and culture in the two species. Finally, our new experiments directly address cumulative cultural learning. Initial results demonstrate a relative conservatism and conformity in chimpanzees' learning, contrasting with cumulative cultural learning in young children. This difference may contribute much to the contrast in these species' capacities for cultural evolution.

Evolutionary theory predicts that natural selection will fashion cognitive biases to guide when, and from whom, individuals acquire social information, but the precise nature of these biases, especially in ecologically valid group contexts, remains unknown. We exposed four captive groups of chimpanzees (Pan troglodytes) to a novel extractive foraging device and, by fitting statistical models, isolated four simultaneously operating transmission biases. These include biases to copy (i) higher-ranking and (ii) expert individuals, and to copy others when (iii) uncertain or (iv) of low rank. High-ranking individuals were relatively un-strategic in their use of acquired knowledge, which, combined with the bias for others to observe them, may explain reports that high innovation rates (in juveniles and subordinates) do not generate a correspondingly high frequency of traditions in chimpanzees. Given the typically low rank of immigrants in chimpanzees, a ‘copying dominants’ bias may contribute to the observed maintenance of distinct cultural repertoires in neighboring communities despite sharing similar ecology and knowledgeable migrants. Thus, a copying dominants strategy may, as often proposed for conformist transmission, and perhaps in concert with it, restrict the accumulation of traditions within chimpanzee communities whilst maintaining cultural diversity.

One effective method for measuring personality in primates is to use personality trait ratings to distill the experience of people familiar with the individual animals. Previous rating instruments were created using either top-down or bottom-up approaches. Top-down approaches, which essentially adapt instruments originally designed for use with another species, can unfortunately lead to the inclusion of traits irrelevant to chimpanzees or fail to include all relevant aspects of chimpanzee personality. Conversely, because bottom-up approaches derive traits specifically for chimpanzees, their unique items may impede comparisons with findings in other studies and other species. To address the limitations of each approach, we developed a new personality rating scale using a combined top-down/bottom-up design. Seventeen raters rated 99 chimpanzees on the new 41-item scale, with all but one item being rated reliably. Principal components analysis, using both varimax and direct oblimin rotations, identified six broad factors. Strong evidence was found for five of the factors (Reactivity/Undependability, Dominance, Openness, Extraversion, and Agreeableness). A sixth factor (Methodical) was offered provisionally until more data are collected. We validated the factors against behavioral data collected independently on the chimpanzees. The five factors demonstrated good evidence for convergent and predictive validity, thereby underscoring the robustness of the factors. Our combined top-down/ bottom-up approach provides the most extensive data to date to support the universal existence of these five personality factors in chimpanzees. This framework, which facilitates cross-species comparisons, can also play a vital role in understanding the evolution of personality and can assist with husbandry and welfare efforts.

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Emulation has been distinguished from imitation as a form of observational learning because it focuses not on the model's actions but on the action's environmental results. Whether a species emulates, imitates or displays only simpler observational learning is expected to have profound implications for its capacity for cultural transmission. Chimpanzees' observational learning has been suggested to be primarily emulative, but this is an inference largely based upon low fidelity copying in experiments when comparing chimpanzees with humans rather than direct testing. Here we test directly for emulation learning by chimpanzees and children using a 'ghost' condition in which a sliding door obscuring a reward was moved to left or right with no agent visible, a context associated with the only published evidence for emulation learning in a nonhuman species (pigeons). Both children and chimpanzees matched the observed direction of ghost door movement on their first test trial. This is the first evidence for emulation in a non-human primate in the restricted context of a ghost condition. However, only the children continued to match in later trials. Individuals of both species continued to match with 99% or better fidelity when viewing a conspecific model operates the door. We conclude that chimpanzees can and will display emulation learning when the task is as simple as the present one, which contrasts with a failure to do so in a more complex manipulative task tested earlier. However, even with a simple task, emulation alone creates only fleeting fidelity compared with the opportunity to copy a conspecific, when considerable conformity is displayed.

Chimpanzees remain fixed on a single strategy, even if a novel, more efficient, strategy is introduced. Previous studies reporting such findings have incorporated paradigms in which chimpanzees learn one behavioural method and then are shown a new one that the chimpanzees invariably do not adopt. This study provides the first evidence that chimpanzees show such conservatism even when the new method employs the identical required behaviour as the first, but for a different reward. Groups of chimpanzees could choose to exchange one of two types of inedible tokens, with each token type being associated with a different food reward: one type was rewarded with a highly preferred food (grape) and the other type was rewarded with a less preferred food (carrot). Individuals first observed a model chimpanzee from their social group trained to choose one of the two types of tokens. In one group, this token earned a carrot, while in the other, control, group the token earned a grape. In both groups, chimpanzees conformed to the trained model’s choice. This was especially striking for those gaining the pieces of carrot, the less favoured reward. This resulted in a population-level trend of food choices, even when counter to their original, individual, preferences. Moreover, the chimpanzees’ food preferences did not change over time, demonstrating that these results were not due to a simple shift in preferences. We discuss social factors apparent in the interactions and suggest that, despite seeming to be inefficient, in chimpanzees, conformity may benefit them, possibly by assisting with the maintenance of group relations.

Inferring the evolutionary history of cognitive abilities requires large and diverse samples. However, such samples are often beyond the reach of individual researchers or institutions, and studies are often limited to small numbers of species. Consequently, methodological and site-specific-differences across studies can limit comparisons between species. Here we introduce the ManyPrimates project, which addresses these challenges by providing a large-scale collaborative framework for comparative studies in primate cognition. To demonstrate the viability of the project we conducted a case study of short-term memory. In this initial study, we were able to include 176 individuals from 12 primate species housed at 11 sites across Africa, Asia, North America and Europe. All subjects were tested in a delayed-response task using consistent methodology across sites. Individuals could access food rewards by remembering the position of the hidden reward after a 0, 15, or 30-second delay. Overall, individuals performed better with shorter delays, as predicted by previous studies. Phylogenetic analysis revealed a strong phylogenetic signal for short-term memory. Although, with only 12 species, the validity of this analysis is limited, our initial results demonstrate the feasibility of a large, collaborative open-science project. We present the ManyPrimates project as an exciting opportunity to address open questions in primate cognition and behaviour with large, diverse datasets.

Network optimality has been described in genes, proteins and human communicative networks. In the latter, optimality leads to the efficient transmission of information with a minimum number of connections. Whilst studies show that differences in centrality exist in animal networks with central individuals having higher fitness, network efficiency has never been studied in animal groups. Here we studied 78 groups of primates (24 species). We found that group size and neocortex ratio were correlated with network efficiency. Centralisation (whether several individuals are central in the group) and modularity (how a group is clustered) had opposing effects on network efficiency, showing that tolerant species have more efficient networks. Such network properties affecting individual fitness could be shaped by natural selection. Our results are in accordance with the social brain and cultural intelligence hypotheses, which suggest that the importance of network efficiency and information flow through social learning relates to cognitive abilities.