Experimental investigation of orangutans’ lithic percussive and sharp stone tool behaviours

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

Establishing an infrastructure for collaboration in primate cognition research

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

The evolution of primate short-term memory

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 shortterm 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

Captive great apes tend to innovate simple tool behaviors quickly.

Recent studies have highlighted the important role that individual learning mechanisms and different forms of enhancenment play in the acquisition of novel behaviors by naïve individuals. A considerable subset of these studies has focused on tool innovation by our closest living relatives, the great apes, to better undestand the evolution of technology in our own lineage. To be able to isolate the role that individual learning plays in great ape tool innovation, researchers usually employ what are known as baseline tests. Although these baselines are commonly used in behavioral studies in captivity, the length of these tests in terms of number of trials and duration remains unstandarized across studies. To address this methodological issue, we conducted a literature review of great ape tool innovation studies conducted in zoological institutions and compiled various methodological data including the timing of innovation. Our literature review revealed an early innovation tendency in great apes, which was particularly pronounced when simple forms of tool use were investigated. In the majority of experiments where tool innovation took place, this occurred within the first trial and/or the first hour of testing. We discuss different possible sources of variation in the latency to innovate such as testing setup, species and task. We hope that our literature review helps researchers design more data-informed, resource-efficient experiments on tool innovation in our closest living relatives

Bone-related behaviours of captive chimpanzees (Pan troglodytes) during two excavating experiments.

After stone tools, bone tools are the most abundant artefact type in the Early Pleistocene archaeological record. That said, they are still relatively scarce, which limits our understanding of the behaviours that led to their production and use. Observations of extant primates constitute a unique source of behavioural data with which to construct hypotheses about the technological forms and repertoires exhibited by our hominin ancestors. We conducted two different experiments to investigate the behavioural responses of two groups of captive chimpanzees (Pan troglodytes; n = 33 and n = 9) to disarticulated, defleshed, ungulate bones while participating in a foraging task aimed at eliciting excavating behaviour. Each chimpanzee group was provided with bone specimens with different characteristics, and the two groups differed in their respective experience levels with excavating plant tools. We found that several individuals from the inexperienced group used the provided bones as tools during the task. In contrast, none of the individuals from the experienced group used bones as excavating tools, but instead continued using plant tools. These chimpanzees also performed non-excavating bone behaviours such as percussion and tool-assisted extraction of organic material from the medullary cavity. Our findings serve as a proof-of-concept that chimpanzees can be used to investigate spontaneous bone tool behaviours such as bone-assisted excavation. Furthermore, our results raise interesting questions regarding the role that bone characteristics, as well as previous tool-assisted excavating experience with other raw materials, might have in the expression of bone tool-assisted excavation

Captivity and habituation to humans raise curiosity in vervet monkeys.

The cognitive mechanisms causing intraspecific behavioural differences between wild and captive animals remain poorly understood. Although diminished neophobia, resulting from a safer environment and more "free" time, has been proposed to underlie these differences among settings, less is known about how captivity influences exploration tendency. Here, we refer to the combination of reduced neophobia and increased interest in exploring novelty as "curiosity", which we systematically compared across seven groups of captive and wild vervet monkeys (Chlorocebus pygerythrus) by exposing them to a test battery of eight novel stimuli. In the wild sample, we included both monkeys habituated to human presence and unhabituated individuals filmed using motion-triggered cameras. Results revealed clear differences in number of approaches to novel stimuli among captive, wild-habituated and wild-unhabituated monkeys. As foraging pressure and predation risks are assumed to be equal for all wild monkeys, our results do not support a relationship between curiosity and safety or free time. Instead, we propose "the habituation hypothesis" as an explanation of why well-habituated and captive monkeys both approached and explored novelty more than unhabituated individuals. We conclude that varying levels of human and/or human artefact habituation, rather than the risks present in natural environments, better explain variation in curiosity in our sample of vervet monkeys

Precise tactile stimulation of worker ants by a robotic manipulator reveals that individual responses are density- and context-dependent

1. Ant workers are often specialized in specific tasks, and it is well-established that the main task an ant performs in the colony can be used to predict its sensitivity and responses to task-associated stimuli. An often-overlooked aspect of ants' task specialization is that individuals often switch tasks throughout the day and are not always engaged in functional tasks. Furthermore, the tasks individuals engage in are often correlated with other context-specific factors, such as worker density, which can independently influence individuals' behaviour. Given this intra-individual variation in task engagement and its correlation with density, it is currently unknown how these two factors interact to modulate ants' sensitivity and responses to stimuli.2. To address this question, we built a robotic manipulation system that allowed us to teleoperate a dummy inside ant colonies and to provide simulated antennations to ants when performing different tasks in areas with different worker densities. We coupled this manipulation system with a custom-built automated tracking system (FORT) that allowed us to track individual identities and locations as well as to record the ants' responses to the dummy stimulation.3. We found independent effects of task and worker density on ants' responsiveness and alarm towards the dummy. Ants were less likely to respond and be alarmed by the dummy when stimulated in areas with high worker density. Responsiveness but not alarm was further influenced by the task being performed, with ants doing broodcare being the least responsive. Our results suggest that ants' behaviour is density-dependent and that ants experience a process of habituation to tactile stimulation. Additionally, ants' responsiveness is modulated by the task they are performing at a given time, showing that sensitivity to stimuli is context-dependent.4. Our robotic set-up constitutes a valuable tool to systematically investigate social insect behaviour under unprecedented experimental control to unravel the individual-level behavioural rules that underpin the organization of social insect colonies. The integrated system presented here opens new research avenues to empirically investigate the effects of more complex stimuli on social insect behaviour and has the potential to significantly further our understanding of decentralized collective systems

Collaborative open science as a way to reproducibility and new insights in primate cognition research

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