Symbolic signal use in wild chimpanzee gestural communication?: A theoretical framework

Symbolic communication is not obvious in the natural communicative repertoires of our closest living relatives, the great apes. However, great apes do show symbolic competencies in laboratory studies. This includes the understanding and the use of human-provided abstract symbols. Given this evidence for the underlying ability, the apparent failure to make use of it in the wild is puzzling. We provide a theoretical framework for identifying basic forms of symbolic signal use in chimpanzee natural communication. In line with the laboratory findings, we concentrate on the most promising domain to investigate, namely gesture, and we provide a case study in this area. We suggest that evidence for basic symbolic signal use would consist of the presence of two key characteristics of symbolic communication, namely arbitrariness and conventionalization. Arbitrariness means that the linkage between the form of the gesture and its meaning shows no obvious logical or otherwise motivated connection. Conventionalization means that the gesture is shared at the group-level and is thus socially learned, not innate. Further, we discuss the emergence and transmission of these gestures. Demonstrating this basic form of symbolic signal use would indicate that the symbolic capacities revealed by laboratory studies also find their expression in the natural gestural communication of our closest living relatives, even if only to a limited extent. This theoretical article thus aims to contribute to our understanding of the developmental origins of great ape gestures, and hence, arguably, of human symbolic communication. It also has a very practical aim in that by providing clear criteria and by pointing out potential candidates for symbolic communication, we give fieldworkers useful prerequisites for identifying and analyzing signals which may demonstrate the use of great apes’ symbolic capacities in the wild

Modeling a primate technological niche

The ability to modify the environment through the transport of tools has been instrumental in shaping the evolutionary success of humans. Understanding the cause-and-effect relationships between hominin behavior and the environment ultimately requires understanding of how the archaeological record forms. Observations of living primates can shed light on these interactions by investigating how tool-use behaviors produce a material record within specific environmental contexts. However, this requires reconciling data derived from primate behavioral observations with the time-averaged nature of the Plio-Pleistocene archaeological record. Here, we use an agent-based model to investigate how repeated short-distance transport events, characteristic for primate tool use, can result in significant landscape-scale patterning of material culture over time. Our results illustrate the conditions under which accumulated short-distance transport bouts can displace stone tools over long distances. We show that this widespread redistribution of tools can also increase access to tool require resources over time. As such, these results elucidate the niche construction processes associated with this pattern of tool transport. Finally, the structure of the subsequent material record largely depends on the interaction between tool transport and environmental conditions over time. Though these results have implications for inferring hominin tool transports from hominin archaeological assemblages. Furthermore, they highlight the difficulties with connecting specific behavioral processes with the patterning in the archaeological record

Identifying functional and regional differences in chimpanzee stone tool technology

The earliest hominin archaeological sites preserve a record of stone tools used for cutting and pounding. Traditionally, sharp-edged flakes were seen as the primary means by which our earliest ancestors interacted with the world. The importance of pounding tools is increasingly apparent. In some cases, they have been compared with stone hammers and anvils used by chimpanzees for nut-cracking. However, there has been little focus on providing a robust descriptive and quantitative characterization of chimpanzee stone tools, allowing for meaningful comparisons between chimpanzee groups and with archaeological artefacts. Here we apply a primate archaeological approach to characterize the range of chimpanzee nut-cracking stone tools from Djouroutou in the Taï National Park. By combining a techno-typological analysis, and two- and three-dimensional measures of damage, we identify clear differences in the location and extent of damage between nut-cracking hammerstones and anvils used at Djouroutou and when compared with other wild chimpanzee populations. Furthermore, we discuss these results in relation to interpretations of Plio-Pleistocene percussive technology. We highlight potential difficulties in identifying the underlying function of percussive artefacts based on morphological or techno-typological attributes alone. The material record from Djouroutou represents an important new datum of chimpanzee regional and material culture

Wild capuchin monkeys adjust stone tools according to changing nut properties

Animals foraging in their natural environments need to be proficient at recognizing and responding to changes in food targets that affect accessibility or pose a risk. Wild bearded capuchin monkeys (Sapajus libidinosus) use stone tools to access a variety of nut species, including otherwise inaccessible foods. This study tests whether wild capuchins from Serra da Capivara National Park in Brazil adjust their tool selection when processing cashew (Anacardium spp.) nuts. During the ripening process of cashew nuts, the amount of caustic defensive substance in the nut mesocarp decreases. We conducted field experiments to test whether capuchins adapt their stone hammer selection to changing properties of the target nut, using stones of different weights and two maturation stages of cashew nuts. The results show that although fresh nuts are easier to crack, capuchin monkeys used larger stone tools to open them, which may help the monkeys avoid contact with the caustic hazard in fresh nuts. We demonstrate that capuchin monkeys are actively able to distinguish between the maturation stages within one nut species, and to adapt their foraging behaviour accordingly

Distance-decay effect in stone tool transport by wild chimpanzees

Stone tool transport leaves long-lasting behavioural evidence in the landscape. However, it remains unknown how large-scale patterns of stone distribution emerge through undirected, short-term transport behaviours. One of the longest studied groups of stone-tool-using primates are the chimpanzees of the Taï National Park in Ivory Coast, West Africa. Using hammerstones left behind at chimpanzee Panda nut-cracking sites, we tested for a distance-decay effect, in which the weight of material decreases with increasing distance from raw material sources. We found that this effect exists over a range of more than 2 km, despite the fact that observed, short-term tool transport does not appear to involve deliberate movements away from raw material sources. Tools from the millennia-old Noulo site in the Taï forest fit the same pattern. The fact that chimpanzees show both complex short-term behavioural planning, and yet produce a landscape-wide pattern over the long term, raises the question of whether similar processes operate within other stone-tool-using primates, including hominins. Where hominin landscapes have discrete material sources, a distance-decay effect, and increasing use of stone materials away from sources, the Taï chimpanzees provide a relevant analogy for understanding the formation of those landscapes

Chimpanzee wooden tool analysis advances the identification of percussive technology

The ability of humans to mediate environmental variation through tool use is likely the key to our success. However, our current knowledge of early cultural evolution derives almost exclusively from studies of stone tools and fossil bones found in the archaeological record. Tools made of plants are intrinsically perishable, and as such are almost entirely absent in the early record of human material culture. Modern human societies as well as nonhuman primate species use plant materials for tools far more often than stone, suggesting that current archaeological data are missing a substantial component of ancient technology. Here, we develop methods that quantify internal and external damage pattern in percussive wooden tools of living primates. Our work shows that the inflicted damage is irreversible, potentially persisting throughout fossilization processes. This research presents opportunities to investigate organic artifacts, a significant and highly neglected aspect of technological evolution within the Primate order

Analysis of wild macaque stone tools used to crack oil palm nuts

The discovery of oil palm (Elaeis guineensis) nut-cracking by wild long-tailed macaques (Macaca fascicularis) is significant for the study of non-human primate and hominin percussive behaviour. Up until now, only West African chimpanzees (Pan troglodytes verus) and modern human populations were known to use stone hammers to crack open this particular hard-shelled palm nut. The addition of non-habituated, wild macaques increases our comparative dataset of primate lithic percussive behaviour focused on this one plant species. Here, we present an initial description of hammerstones used by macaques to crack oil palm nuts, recovered from active nut-cracking locations on Yao Noi Island, Ao Phang Nga National Park, Thailand. We combine a techno-typological approach with microscopic and macroscopic use-wear analysis of percussive damage to characterize the percussive signature of macaque palm oil nut-cracking tools. These artefacts are characterized by a high degree of battering and crushing on most surfaces, which is visible at both macro and microscopic levels. The degree and extent of this damage is a consequence of a dynamic interplay between a number of factors, including anvil morphology and macaque percussive techniques. Beyond the behavioural importance of these artefacts, macaque nut-cracking represents a new target for primate archaeological investigations, and opens new opportunities for comparisons between tool using primate species and with early hominin percussive behaviour, for which nutcracking has been frequently inferred

Social network analysis shows direct evidence for social transmission of tool use in wild chimpanzees

The authors are grateful to the Royal Zoological Society of Scotland for providing core funding for the Budongo Conservation Field Station. The fieldwork of CH was funded by the Leverhulme Trust, the Lucie Burgers Stichting, and the British Academy. TP was funded by the Canadian Research Chair in Continental Ecosystem Ecology, and received computational support from the Theoretical Ecosystem Ecology group at UQAR. The research leading to these results has received funding from the People Programme (Marie Curie Actions) and from the European Research Council under the European Union's Seventh Framework Programme (FP7/2007–2013) REA grant agreement n°329197 awarded to TG, ERC grant agreement n° 283871 awarded to KZ. WH was funded by a BBSRC grant (BB/I007997/1).Social network analysis methods have made it possible to test whether novel behaviors in animals spread through individual or social learning. To date, however, social network analysis of wild populations has been limited to static models that cannot precisely reflect the dynamics of learning, for instance, the impact of multiple observations across time. Here, we present a novel dynamic version of network analysis that is capable of capturing temporal aspects of acquisition-that is, how successive observations by an individual influence its acquisition of the novel behavior. We apply this model to studying the spread of two novel tool-use variants, "moss-sponging'' and "leaf-sponge re-use,'' in the Sonso chimpanzee community of Budongo Forest, Uganda. Chimpanzees are widely considered the most "cultural'' of all animal species, with 39 behaviors suspected as socially acquired, most of them in the domain of tool-use. The cultural hypothesis is supported by experimental data from captive chimpanzees and a range of observational data. However, for wild groups, there is still no direct experimental evidence for social learning, nor has there been any direct observation of social diffusion of behavioral innovations. Here, we tested both a static and a dynamic network model and found strong evidence that diffusion patterns of moss-sponging, but not leaf-sponge re-use, were significantly better explained by social than individual learning. The most conservative estimate of social transmission accounted for 85% of observed events, with an estimated 15-fold increase in learning rate for each time a novice observed an informed individual moss-sponging. We conclude that group-specific behavioral variants in wild chimpanzees can be socially learned, adding to the evidence that this prerequisite for culture originated in a common ancestor of great apes and humans, long before the advent of modern humans.Publisher PDFPeer reviewe

Primate Archaeology Evolves

Since its inception, archaeology has traditionally focused exclusively on humans and our direct ancestors. However, recent years have seen archaeological techniques applied to material evidence left behind by non-human animals. Here, we review advances made by the most prominent field investigating past non-human tool use: primate archaeology. This field combines survey of wild primate activity areas with ethological observations, excavations and analyses that allow the reconstruction of past primate behaviour. Because the order Primates includes humans, new insights into the behavioural evolution of apes and monkeys also can be used to better interrogate the record of early tool use in our own, hominin, lineage. This work has recently doubled the set of primate lineages with an excavated archaeological record, adding Old World macaques and New World capuchin monkeys to chimpanzees and humans, and it has shown that tool selection and transport, and discrete site formation, are universal among wild stone-tool-using primates. It has also revealed that wild capuchins regularly break stone tools in a way that can make them difficult to distinguish from simple early hominin tools. Ultimately, this research opens up opportunities for the development of a broader animal archaeology, marking the end of archaeology’s anthropocentric era

Food cleaning in gorillas: Social learning is a possibility but not a necessity

Food cleaning is widespread in the animal kingdom, and a recent report confirmed that (amongst other behaviours) wild western lowland gorillas also show food cleaning. The authors of this report conclude that this behaviour, based on its distribution patterns, constitutes a potential candidate for culture. While different conceptualisations of culture exist, some more and some less reliant on behavioural form copying, all of them assign a special role to social learning processes in explaining potentially cultural behaviours. Here we report the results of an experiment that tested to what extent food cleaning behaviour in a group of captive Western lowland gorillas (Gorilla gorilla gorilla) relies on social learning processes. Subjects were provided with clean and dirty apples. When they were provided with dirty apples, all subjects showed evidence of food cleaning in at least 75% of trials. Preferred cleaning techniques differed between individuals, four out of five of subjects expressed a behaviour analogous to that reported in wild conspecifics. Given this occurrence of food cleaning in a culturally unconnected population of gorillas, we conclude that social learning is unlikely to play a central role in the emergence of the food cleaning behavioural form in Western lowland gorillas; instead, placing a greater emphasis on individual learning of food cleaning’s behavioural form