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.

Despite major advances in the study of animal tool behaviour, researchers continue to debate how exactly certain behaviours are acquired. While specific mechanisms, such as genetic predispositions or action copying, are sometimes suspected to play a major role in behavioural acquisition, controlled experiments are required to provide conclusive evidence. In this opinion piece, we refer to classic ethological methodologies to emphasize the need for studying the relative contributions of different factors to the emergence of specific tool behaviours. We describe a methodology, consisting of a carefully staged series of baseline and social-learning conditions, that enables us to tease apart the roles of different mechanisms in the development of behavioural repertoires. Experiments employing our proposed methodology will not only advance our understanding of animal learning and culture, but as a result, will also help inform hypotheses about human cognitive, cultural and technological evolution. More generally, our conceptual framework is suitable for guiding the detailed investigation of other seemingly complex animal behaviours.

The field of primate cognition studies how primates, including humans, perceive, process, store, retrieve, and use information to guide decision making and other behavior. Much of this research is motivated by a desire to understand how these abilities evolved. Large and diverse samples from a wide range of species are vital to achieving this goal. In reality, however, primate cognition research suffers from small sample sizes and is often limited to a handful of species, which constrains the evolutionary inferences we can draw. We conducted a systematic review of primate cognition research published between 2014 and 2019 to quantify the extent of this problem. Across 574 studies, the median sample size was 7 individuals. Less than 15% of primate species were studied at all, and only 19% of studies included more than one species. Further, the species that were studied varied widely in how much research attention they received, partly because a small number of test sites contributed most of the studies. These results suggest that the generalizability of primate cognition studies may be severely limited. Publication bias, questionable research practices, and a lack of replication attempts may exacerbate these problems. We describe the ManyPrimates project as one approach to overcoming some of these issues by establishing an infrastructure for large-scale collaboration in primate cognition research. Building on similar initiatives in other areas of psychology, this approach has already yielded one of the largest and most diverse primate samples to date and enables us to ask many research questions that can only be addressed through collaboration.

It is hypothesized that tool-assisted excavation of plant underground storage organs (USOs) played an adaptive role in hominin evolution and was also once considered a uniquely human behavior. Recent data indicate that savanna chimpanzees also use tools to excavate edible USOs. However, those chimpanzees remain largely unhabituated and we lack direct observations of this behavior in the wild. To fill this gap in our knowledge of hominoid USO extractive foraging, we conducted tool-mediated excavation experiments with captive chimpanzees naïve to this behavior. We presented the chimpanzees with the opportunity to use tools in order to excavate artificially-placed underground foods in their naturally forested outdoor enclosure. No guidance or demonstration was given to the chimpanzees at any time. The chimpanzees used tools spontaneously in order to excavate the underground foods. They exhibited six different tool use behaviors in the context of excavation: probe, perforate, dig, pound, enlarge and shovel. However, they still excavated manually more often than they did with tools. Chimpanzees were selective in their choice of tools that we provided, preferring longer tools for excavation. They also obtained their own tools mainly from naturally occurring vegetation and transported them to the excavation site. They reused some tools throughout the study. Our new data provide a direction for the study of variables relevant to modeling USO extractive foraging by early hominins.

Short-term memory is implicated in a range of cognitive abilities and is critical for understanding primate cognitive evolution. To investigate the effects of phylogeny, ecology and sociality on short-term memory, we tested the largest and most diverse primate sample to date (421 non-human primates across 41 species) in an experimental delayed-response task. Our results confirm previous findings that longer delays decrease memory performance across species and taxa. Our analyses demonstrate a considerable contribution of phylogeny over ecological and social factors on the distribution of short-term memory performance in primates; closely related species had more similar short-term memory abilities. Overall, individuals in the branch of Hominoidea performed better compared to Cercopithecoidea, who in turn performed above Platyrrhini and Strepsirrhini. Interdependencies between phylogeny and socioecology of a given species presented an obstacle to disentangling the effects of each of these factors on the evolution of short-term memory capacity. However, this study offers an important step forward in understanding the interspecies and individual variation in short-term memory ability by providing the first phylogenetic reconstruction of this trait’s evolutionary history. The dataset constitutes a unique resource for studying the evolution of primate cognition and the role of short-term memory in other cognitive abilities.

The diversity of great ape diets requires behavioral flexibility. Consequently, the exploration of potentially novel food sources is supposedly beneficial, but simultaneously, apes show high neophobia to prevent harmful and poisonous food intake. Social information, such as presence of group members or observations of nonnaïve, experienced individuals have been demonstrated to affect the acceptance of novel food items in primates. Sociality may have evolutionary effects on the response of apes to novel foods. Here we assess the social information hypothesis, which predicts that selection favors higher neophobia in species where social information is abundant.We report the results from 134 great apes housed in multiple facilities from four closely related species that naturally differ in their degree of sociality: Pongo pygmaeus, Pongo abelii, Pan troglodytes and Pan paniscus. We examined individuals' reactions to novel foods when alone, which enabled us to detect any inherent differences and revealed significant distinctions between species. Chimpanzees and bonobos, that are naturally exposed to higher amounts of social information, were less likely to consume novel foods alone (showed higher neophobia) than the two more solitary orangutan species.Chimpanzees were especially cautious and showed higher explorative behaviors before tasting novel food than other species. Age influenced neophobia as younger individuals of all species took longer to taste novel foods than adults did. K E Y W O R D Sgreat apes, neophobia, novel food, sociality, social information

Early stone tools, and in particular sharp stone tools, arguably represent one of the most important technological milestones in human evolution. The production and use of sharp stone tools significantly widened the ecological niche of our ancestors, allowing them to exploit novel food resources. However, despite their importance, it is still unclear how these early lithic technologies emerged and which behaviours served as stepping-stones for the development of systematic lithic production in our lineage. One approach to answer this question is to collect comparative data on the stone tool making and using abilities of our closest living relatives, the great apes, to reconstruct the potential stone-related behaviours of early hominins. To this end, we tested both the individual and the social learning abilities of five orangutans to make and use stone tools. Although the orangutans did not make sharp stone tools initially, three individuals spontaneously engaged in lithic percussion, and sharp stone pieces were produced under later experimental conditions. Furthermore, when provided with a human-made sharp stone, one orangutan spontaneously used it as a cutting tool. Contrary to previous experiments, social demonstrations did not considerably improve the stone tool making and using abilities of orangutans. Our study is the first to systematically investigate the stone tool making and using abilities of untrained, unenculturated orangutans showing that two proposed pre-requisites for the emergence of early lithic technologies–lithic percussion and the recognition of sharp-edged stones as cutting tools–are present in this species. We discuss the implications that ours and previous great ape stone tool experiments have for understanding the initial stages of lithic technologies in our lineage.

Background: Despite substantial research on early hominin lithic technologies, the learning mechanisms underlying flake manufacture and use are contested. To draw phylogenetic inferences on the potential cognitive processes underlying the acquisition of both of these abilities in early hominins, we investigated if and how one of our closest living relatives, chimpanzees (Pan troglodytes), could learn to make and use flakes. Methods: Across several experimental conditions, we tested unenculturated, naïve chimpanzees from two independent populations (n=11) for their abilities to spontaneously make and use their own flakes as well as to use pre-made flakes made by a human experimenter. Results: Despite the fact that the chimpanzees demonstrated an understanding of the requirements of the task and that subjects were sufficiently motivated and had ample opportunities to develop these behaviours, none of the chimpanzees tested, made or used flakes in any of the experimental conditions. Conclusions: These results differ from all previous ape flaking experiments, which found flake manufacture and use in bonobos and one orangutan. However, these earlier studies tested human-enculturated apes and provided the test subjects with flake making and using demonstrations. The contrast between these earlier positive findings and our negative findings (despite using a much larger sample size) suggests that human enculturation and/or human demonstrations may be necessary for chimpanzees to acquire these abilities. The data obtained in this study are consistent with the hypothesis that flake manufacture and use might have evolved in the hominin lineage after the split between Homo and Pan 7 million years ago, a scenario further supported by the initial lack of flaked stone tools in the archaeological record after this split. We discuss possible evolutionary scenarios for flake manufacture and use in both non-hominin and hominin lineages.