Influence of COVID-19 on the emergence of stone-tool use behavior in a population of common long-tailed macaques (Macaca fascicularis fascicularis) in Thailand.

Stone tool use is a rare behavior across nonhuman primates. Here we report the first population of common long-tailed macaques (Macaca fascicularis fascicularis) who customarily used stone tools to open rock oysters (Saccostrea forskali) on a small island along the Thai Gulf in Koh Ped (KPE), eastern Thailand. We observed this population several times during the past 10 years, but no stone-tool use behavior was observed until our survey during the coronavirus disease 2019 (COVID-19) pandemic in July 2022. KPE is located in Pattaya City, a hotspot for tourism in Thailand. Tourists in this area frequently provided large amounts of food for the monkeys on KPE. During the COVID-19 curfew, however, tourists were not allowed to access the island, and monkeys began to face food scarcity. During this time, we observed stone-tool use behavior for the first time on KPE. Based on our observations, the first tool manipulation was similar to stone throwing (a known precursor of stone tool use). From our observations in March 2023, we found 17 subadult/adult animals performing the behavior, 15 of 17 were males and mostly solitary while performing the behavior. The M. f. fascicularis subspecies was confirmed by distribution, morphological characteristics, and mtDNA and SRY gene sequences. Taken together, we proposed that the stone tool use behavior in the KPE common long-tailed macaques emerged due to the COVID-19 food scarcity. Since traveling is no longer restricted many tourists have started coming back to the island, and there is a high risk for this stone tool-use behavior to disappear within this population of long-tailed macaques.

Simulation and social network analysis provide insight into the acquisition of tool behaviour in hybrid macaques.

The pathways through which primates acquire skills are a central focus of cultural evolution studies. The roles of social and genetic inheritance processes in skill acquisition are often confounded by environmental factors. Hybrid macaques from Koram Island (Thailand) provide an opportunity to examine the roles of inheritance and social learning to skill acquisition within a single ecological setting. These hybrids are a cross between tool-using Burmese long-tailed (Macaca fascicularis aurea) and non-tool-using common long-tailed macaques (Macaca fascicularis fascicularis). This population provides an opportunity to explore the roles of social learning and inheritance processes while being able to exclude underlying ecological factors. Here, we investigate the roles of social learning and inheritance in tool use prevalence within this population using social network analysis and simulation. Agent-based modelling (ABM) is used to generate expectations for how social/asocial learning and inheritance structure the patterning in a social network. The results of the simulation show that various transmission mechanisms can be differentiated based on associations between individuals in a social network. The results provide an investigative framework for discussing tool use transmission pathways in the Koram social network. By combining ABM, network analysis, and behavioural data from the field we can investigate the roles social learning and inheritance play in tool acquisition of wild primates.

Modeling Oldowan tool transport from a primate perspective.

Living nonhuman primates have long served as a referential framework for understanding various aspects of hominin biological and cultural evolution. Comparing the cognitive, social, and ecological contexts of nonhuman primate and hominin tool use has allowed researchers to identify key adaptations relevant to the evolution of hominin behavior. Although the Oldowan is often considered to be a major evolutionary milestone, it has been argued that the Oldowan is rather an extension of behaviors already present in the ape lineage. This is based on the fact that while apes move tools through repeated, unplanned, short-distance transport bouts, they produce material patterning often associated with long-distance transport, planning, and foresight in the Oldowan. Nevertheless, remain fundamental differences in how Oldowan core and flake technology and nonhuman primate tools are used. The goal of the Oldowan hominins is to produce sharp-edged flakes, whereas nonhuman primates use stone tools primarily as percussors. Here, we present an agent-based model that investigates the explanatory power of the ape tool transport model in light of these differences. The model simulates the formation of the Oldowan record under the conditions of an accumulated short-distance transport pattern, as seen in extant chimpanzees. Our results show that while ape tool transport can account for some of the variation observed in the archaeological record, factors related to use-life duration severely limit how far an Oldowan core can be moved through repeated short-distance transport bouts. Thus, the ape tool transport has limitations in its ability to explain patterns in the Oldowan. These results provide a basis for discussing adaptive processes that would have facilitated the development of the Oldowan.

A primate model for the origin of flake technology.

When and how human ancestors first used tools remains unknown, despite intense research into the origins of technology. It has been hypothesized that the evolutionary roots of stone flake technology has its origin in percussive behavior. Before intentional stone flaking, hominins potentially engaged in various percussive behaviors resulting in accidental flake detachments. We refer to this scenario as the 'by-product hypothesis.' In this scenario, repeated detachments of sharp stone fragments eventually resulted in intentional flake production. Here, we tested the circumstances of accidental flake production as a by-product of percussive foraging in wild capuchin monkeys (Sapajus libidinosus) from Brazil, the only nonhuman primate known to habitually produce sharp-edged flakes through a percussive behavior. We conducted field experiments where we tested the potential for accidental flake production during nut cracking. We provided three different types of stone with varied material properties as anvils to assess the circumstances in which accidental production of sharp-edged flakes occurs during nut cracking. A further freehand knapping experiment, with the raw material that exhibited accidental flake detachments, allows a direct comparison of flakes that have been intentionally produced by an experienced knapper and flakes produced during nut cracking by capuchin monkeys. Our results show that raw material quality and morphology significantly affect the rate of sharp-edged flake production as well as the resulting lithic signature of this behavior. In addition, accidental flakes produced during capuchin nut cracking on highly isotropic raw material are similar in many respects to intentionally produced flakes by a human knapper. Our field experiments highlight the fact that nut-cracking behavior can lead to the unintentional production of substantial quantities of sharp-edged flakes and therefore supports the 'by-product hypothesis' as a potential mechanism for the emergence of hominin flake technology.

Wild monkeys flake stone tools.

Our understanding of the emergence of technology shapes how we view the origins of humanity. Sharp-edged stone flakes, struck from larger cores, are the primary evidence for the earliest stone technology. Here we show that wild bearded capuchin monkeys (Sapajus libidinosus) in Brazil deliberately break stones, unintentionally producing recurrent, conchoidally fractured, sharp-edged flakes and cores that have the characteristics and morphology of intentionally produced hominin tools. The production of archaeologically visible cores and flakes is therefore no longer unique to the human lineage, providing a comparative perspective on the emergence of lithic technology. This discovery adds an additional dimension to interpretations of the human Palaeolithic record, the possible function of early stone tools, and the cognitive requirements for the emergence of stone flaking.

Sleep patterns, daytime predation, and the evolution of diurnal sleep site selection in lorisiforms.

OBJECTIVES: Synthesize information on sleep patterns, sleep site use, and daytime predation at sleep sites in lorisiforms of Asia and Africa (10 genera, 36 species), and infer patterns of evolution of sleep site selection. MATERIALS AND METHODS: We conducted fieldwork in 12 African and six Asian countries, collecting data on sleep sites, timing of sleep and predation during daytime. We obtained additional information from literature and through correspondence. Using a phylogenetic approach, we established ancestral states of sleep site selection in lorisiforms and traced their evolution. RESULTS: The ancestral lorisiform was a fur-clinger and used dense tangles and branches/forks as sleep sites. Use of tree holes and nests as sleep sites emerged ∼22 Mya (range 17-26 Mya) in Africa, and use of bamboo emerged ∼11 (7-14) Mya in Asia and later in Africa. Fur clinging and some sleep sites (e.g., tree holes, nests, but not bamboo or dense tangles) show strong phylogenetic signal. Nests are used by Galagoides, Paragalago, Galago and Otolemur; tree holes by Galago, Paragalago, Sciurocheirus and Perodicticus; tangles by Nycticebus, Loris, Galagoides, Galago, Euoticus, Otolemur, Perodicticus and Arctocebus; all but Sciurocheirus and Otolemur additionally sleep on branches/forks. Daytime predation may affect sleep site selection and sleep patterns in some species of Nycticebus, Galago, Galagoides, Otolemur and Perodicticus. Most lorisiforms enter their sleep sites around sunrise and leave around sunset; several are active during twilight or, briefly, during daytime. CONCLUSION: Variations in sleep behavior, sleep patterns and vulnerability to daytime predation provide a window into the variation that was present in sleep in early primates. Overall, lorisiforms use the daytime for sleeping and no species can be classified as cathemeral or polycyclic.

Analysis of sea almond (Terminalia catappa) cracking sites used by wild Burmese long-tailed macaques (Macaca fascicularis aurea).

Nut-cracking is shared by all non-human primate taxa that are known to habitually use percussive stone tools in the wild: robust capuchins (Sapajus spp.), western chimpanzees (Pan troglodytes verus), and Burmese long-tailed macaques (Macaca fascicularis aurea). Despite opportunistically processing nuts, Burmese long-tailed macaques predominantly use stone tools to process mollusks in coastal environments. Here, we present the first comprehensive survey of sea almond (Terminalia catappa) nut-cracking sites created by macaques. We mapped T. catappa trees and nut-cracking sites that we encountered along the intertidal zone and forest border on the coasts of Piak Nam Yai Island, Thailand. For each nut-cracking site, we measured the physical properties (i.e., size, weight, use-wear) of hammer stones and anvils. We found that T. catappa trees and nut-cracking sites primarily occurred on the western coast facing the open sea, and cracking sites clusters around the trees. We confirmed previous results that nut cracking tools are among the heaviest tools used by long-tailed macaques; however, we found our sample of T. catappa stone tools lighter than a previously collected sea almond sample that, unlike our sample, was collected immediately after use within the intertidal zone. The difference was likely the result of tidal influences on tool-use sites. We also found that tool accumulations above the intertidal region do not resemble those within them, possibly leading to incomplete assessments of macaque stone tools through archaeological techniques that would use these durable sites.

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.

Cashew Nut Positioning during Stone Tool Use by Wild Bearded Capuchin Monkeys (Sapajus libidinosus).

Wild capuchin monkeys (Sapajus libidinosus) at Serra da Capivara National Park, Brazil, regularly use stone tools to break open cashew nuts (Anacardium spp.). Here we examine 2 approaches used by the capuchins to position the kidney-shaped cashew nuts on an anvil before striking with a stone tool. Lateral positioning involves placing the nut on its flatter, more stable side, therefore requiring less attention from the monkey during placement. However, the less stable and never previously described arched position, in which the nut is balanced with its curved side uppermost, requires less force to crack the outer shell. We observed cashew nut cracking in a field experimental setting. Only 6 of 20 adults, of both sexes, were observed to deliberately place cashew nuts in an arched position, which may indicate that the technique requires time and experience to learn. We also found that use of the arched position with dry nuts, but not fresh, required, in 63% of the time, an initial processing to remove one of the cashew nut lobes, creating a more stable base for the arch. This relatively rare behaviour appears to have a complex ontogeny, but further studies are required to establish the extent to which social learning is involved.

The extent of cultural variation between adjacent chimpanzee (Pan troglodytes verus) communities; a microecological approach.

Chimpanzees show cultural differences among populations across Africa but also between neighboring communities. The extent of these differences among neighbors, however, remains largely unknown. Comparing three neighboring chimpanzee community in the Taï National Park, Côte d'Ivoire, we found 27 putative cultural traits, including tool use, foraging, social interaction, communication and hunting behavior, exceeding by far previously known diversity. As foraging behavior is predominantly influenced by the environment, we further compared in detail ecological circumstances underlying insectivore feeding behavior to analyze whether foraging differences on Dorylus ants and Thoracotermes termites seen between neighboring chimpanzee communities were caused by environmental factors. Differences in the prey characteristics of Dorylus ants (aggression level, running speed, and nest structure) that could influence the behavior of chimpanzees were excluded, suggesting that the observed group-specific variation is not ecologically driven. Only one community preyed on Thoracotermes termites despite a similar abundance of termite mounds in all three territories, supporting the idea that this difference is also not shaped by the environment. Therefore, our study suggests that transmission of cultural knowledge plays a role in determining insectivory prey behavior. This behavioral plasticity, independent of ecological conditions, can lead to large numbers of cultural diversification between neighboring chimpanzee communities. These findings not only deepen our understanding of the cultural abilities of chimpanzees in the wild but also open up possible future comparisons of the origin of cultural diversification among humans and chimpanzees.

Tradition over trend: Neighboring chimpanzee communities maintain differences in cultural behavior despite frequent immigration of adult females.

The notion of animal culture has been well established mainly through research aiming at uncovering differences between populations. In chimpanzees (Pan troglodytes verus), cultural diversity has even been found in neighboring communities, where differences were observed despite frequent immigration of individuals. Female chimpanzees transfer at the onset of sexual maturity at an age, when the behavioral repertoire is fully formed. With immigrating females, behavioral variety enters the group. Little is known about the diversity and the longevity of cultural traits within a community. This study is building on previous findings of differences in hammer selection when nut cracking between neighboring communities despite similar ecological conditions. We now further investigated the diversity and maintenance of cultural traits within one chimpanzee community and were able to show high levels of uniformity in group-specific behavior. Fidelity to the behavior pattern did not vary between dispersing females and philopatric males. Furthermore, group-specific tool selection remained similar over a period of 25 years. Additionally, we present a study case on how one newly immigrant female progressively behaved more similar to her new group, suggesting that the high level of similarity in behavior is actively adopted by group members possibly even when originally expressing the behavior in another form. Taken together, our data support a cultural transmission process in adult chimpanzees, which leads to persisting cultural behavior of one community over time.

In search of animal normativity: a framework for studying social norms in non-human animals.

Social norms - rules governing which behaviours are deemed appropriate or inappropriate within a given community - are typically taken to be uniquely human. Recently, this position has been challenged by a number of philosophers, cognitive scientists, and ethologists, who have suggested that social norms may also be found in certain non-human animal communities. Such claims have elicited considerable scepticism from norm cognition researchers, who doubt that any non-human animals possess the psychological capacities necessary for normative cognition. However, there is little agreement among these researchers about what these psychological prerequisites are. This makes empirical study of animal social norms difficult, since it is not clear what we are looking for and thus what should count as behavioural evidence for the presence (or absence) of social norms in animals. To break this impasse, we offer an approach that moves beyond contested psychological criteria for social norms. This approach is inspired by the animal culture research program, which has made a similar shift away from heavily psychological definitions of 'culture' to become organised around a cluster of more empirically tractable concepts of culture. Here, we propose an analogous set of constructs built around the core notion of a normative regularity, which we define as a socially maintained pattern of behavioural conformity within a community. We suggest methods for studying potential normative regularities in wild and captive primates. We also discuss the broader scientific and philosophical implications of this research program with respect to questions of human uniqueness, animal welfare and conservation.

Emergent technological variation in archaeological landscapes: a primate perspective.

Archaeological evidence informs our understanding of the evolution of hominin behaviour. Such evidence is traditionally used to reconstruct hominin activities and intentions. In the Plio-Pleistocene, the presence or absence of specific tools and variation in artefact density is often used to infer foraging strategies, cognitive traits and functional activities. However, the Plio-Pleistocene archaeological record is known to be time-averaged and forms through the aggregation of repeated behavioural events over time. Thus, archaeological patterns do not reflect discrete episodes of activity, but rather the interaction of behaviour with environmental factors over time. However, little is known about how such interactions produce archaeological variation diversity. Primate archaeology can help address this research gap by providing the opportunity to observe how behaviour produces material patterns in a natural setting. This study, thus, examines how varying the material properties of stone and resource availability influence the artefactual signature of nut-cracking in a population of long-tailed macaques from Lobi Bay, Yao Noi island, Thailand. Results show that these interactions can produce a structured and diverse material signature in terms of artefact density and frequency of specific artefact types. These findings demonstrate how material patterns can emerge from long-term interactions between behaviour and environmental factors.

Wild macaques challenge the origin of intentional tool production.

Intentionally produced sharp-edged stone flakes and flaked pieces are our primary evidence for the emergence of technology in our lineage. This evidence is used to decipher the earliest hominin behavior, cognition, and subsistence strategies. Here, we report on the largest lithic assemblage associated with a primate foraging behavior undertaken by long-tailed macaques (Macaca fascicularis). This behavior results in a landscape-wide record of flaked stone material, almost indistinguishable from early hominin flaked pieces and flakes. It is now clear that the production of unintentional conchoidal sharp-edged flakes can result from tool-assisted foraging in nonhominin primates. Comparisons with Plio-Pleistocene lithic assemblages, dating from 3.3 to 1.56 million years ago, show that flakes produced by macaques fall within the technological range of artifacts made by early hominins. In the absence of behavioral observations, the assemblage produced by monkeys would likely be identified as anthropogenic in origin and interpreted as evidence of intentional tool production.

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.

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.

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.

Three-dimensional surface morphometry differentiates behaviour on primate percussive stone tools.

The Early Stone Age record preserves a rich behavioural signature of hominin stone tool making and use. The role of percussive technology in the daily subsistence strategies of our earliest ancestors has seen renewed focus recently. Studies of modern primate tool use highlight the diverse range of behaviours potentially associated with percussive technology. This has prompted significant methodological developments to characterize the associated damage marks (use-wear) on hammerstones and anvils. Little focus has, however, been paid to identifying whether these techniques can successfully differentiate between the damage patterns produced by specific and differing percussive behaviours. Here, we present a novel workflow drawing on the strengths of visual identification and three-dimensional (3D) surface quantification of use-wear. We apply this methodology firstly to characterize macaque percussive use-wear and test the efficacy of 3D surface quantification techniques in differentiating between percussive damage and natural surface topography. Secondly, we use this method to differentiate between use-wear associated with various wild macaque percussive behaviours. By combining analyst-directed, 3D surface analysis and use-wear dimensional analysis, we show that macaque percussive behaviours create specific diagnostic signatures and highlight a means of quantifiably recording such behavioural signatures in both primate and hominin contexts.

Group-specific archaeological signatures of stone tool use in wild macaques.

Stone tools in the prehistoric record are the most abundant source of evidence for understanding early hominin technological and cultural variation. The field of primate archaeology is well placed to improve our scientific knowledge by using the tool behaviours of living primates as models to test hypotheses related to the adoption of tools by early stone-age hominins. Previously we have shown that diversity in stone tool behaviour between neighbouring groups of long-tailed macaques (Macaca-fascicularis) could be explained by ecological and environmental circumstances (Luncz et al., 2017b). Here however, we report archaeological evidence, which shows that the selection and reuse of tools cannot entirely be explained by ecological diversity. These results suggest that tool-use may develop differently within species of old-world monkeys, and that the evidence of material culture can differ within the same timeframe at local geographic scales and in spite of shared environmental and ecological settings.