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The earliest evidence for complex tool use in the archaeological record dates to 3.3 Ma. While wooden tools may have been used by our earliest ancestors, the evidence is absent due to poor preservation. However, insights into possible early hominin wooden tool use can be gained from observing the tool-use practices of our closest living relatives, chimpanzees (Pan troglodytes). By using stone hammers used to crack various nuts, chimpanzees leave a durable material signature comprised of formal tools and associated diagnostic fragments. While the archaeological evidence of chimpanzee wooden tool use is temporary, the combination of stone hammers and wooden anvils can create a more enduring lithic record. This study explores the lithic assemblages associated with wooden and stone anvil use at nut-cracking sites in Taï National Park, Côte d'Ivoire, using technological and use-wear analyses. Our results indicate clear differences in density, fracture patterns, and use-wear in the lithic records between wooden anvil and stone anvil sites. New archaeological excavations at six chimpanzee nut-cracking sites reveal that the anvils' material directly influences the visibility of nut-cracking evidence in the archaeological record. By examining the nature of the lithic signatures associated with wooden anvil and stone anvil use by chimpanzees, we can formulate hypotheses about the probability of such behaviors being preserved and identifiable in the Plio-Pleistocene hominin archaeological record. The variability in material signatures from nut-cracking on different anvils suggests that stone anvils leave a clear archaeological record. Evidence for wooden anvil use is likely underrepresented due to the more ephemeral nature of the associated percussive damage and material signature. It may, however, still be possible, albeit challenging, to identify wooden anvil use in the archaeological record. [ABSTRACT FROM AUTHOR]

Ecology is fundamental in the development, transmission, and perpetuity of primate technology. Previous studies on tool site selection have addressed the relevance of targeted resources and raw materials for tools, but few have considered the broader foraging landscape. In this landscape-scale study of the ecological contexts of wild chimpanzee (Pan troglodytes verus) tool use, we investigated the conditions required for nut-cracking to occur and persist in discrete locations at the long-term field site of Bossou, Guinea. We examined this at three levels: selection, frequency of use, and inactivity. We collected data on plant foods, nut trees, and raw materials using transect and quadrat methods, and conducted forest-wide surveys to map the location of nests and watercourses. We analysed data at the quadrat level (n = 82) using generalised linear models and descriptive statistics. We found that, further to the presence of a nut tree and availability of raw materials, abundance of food-providing trees as well as proximity to nest sites were significant predictors of nut-cracking occurrence. This suggests that the spatial distribution of nut-cracking sites is mediated by the broader behavioural landscape and is influenced by non-extractive foraging of perennial resources and non-foraging activities. Additionally, the number of functional tools was greater at sites with higher nut-cracking frequency, and was negatively correlated with site inactivity. Our research indicates that the technological landscape of Bossou chimpanzees shares affinities with the 'favoured places' model of hominin site formation, providing a comparative framework for reconstructing landscape-scale patterns of ancient human behaviour. A French translation of this abstract is provided in theelectronic supplementary information: EMS 2. [ABSTRACT FROM AUTHOR]

The origin of tool use is a central question in human evolutionary studies. Plio-Pleistocene core and flake technologies represent the earliest evidence of tool use in the human lineage. Some suggest this form of tool use is probably pre-dated by a phase of percussive tool use. However, there is currently no evidence for such a record. The archaeological signature of solely percussive behaviours is not as well understood as that associated with cores and flakes. The durable nature of primate percussive stone tools and their by-products provide an opportunity to investigate what such a record looks like. Here, we present a landscape-scale study of the chimpanzee ( Pan troglodytes verus ) material culture from the Djouroutou Chimpanzee Project, Taï Forest, Cote d'Ivoire. This study explores the interplay between behavioural and environmental factors in shaping the stone record of nut cracking. Through a survey of nut-cracking sites, the available nut species, and raw materials, we show how resource availability influences the resulting material signature of nut cracking. These results also reveal the diversity of material signatures associated with a purely percussive material record. We gain insight into the range of signatures that may be associated with a pre-core and flake archaeological record, providing new expectations for an earlier record of tool use.

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.

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.

Abstract Archaeological recovery of chimpanzee Panda oleosa nut cracking tools at the Panda 100 (P100) and Noulo sites in the Taï Forest, Côte d'Ivoire, showed that this behavior is over 4000 years old, making it the oldest known evidence of non-human tool use. In 2002, the first report on the lithic material from P100 was directly compared to early hominin stone tools, highlighting their similarities and proposing the name 'Pandan' for the chimpanzee material. Here we present an expanded and comprehensive technological, microscopic, and refit analysis of the late twentieth century lithic assemblage from P100. Our re-analysis provides new data and perspectives on the applicability of chimpanzee nut cracking tools to our understanding of the percussive behaviors of early hominins. We identify several new refit sets, including the longest (>17 m) hammerstone transport seen in the chimpanzee archaeological record. We provide detailed evidence of the fragmentation sequences of Panda nut hammerstones, and characterize the percussive damage on fragmented material from P100. Finally, we emphasize that the chimpanzee lithic archaeological record is dynamic, with the preservation of actual hammerstones being rare, and the preservation of small broken pieces more common. P100 – the first archaeological chimpanzee nut cracking lithic assemblage – provides a valuable comparative sample by which to identify past chimpanzee behavior elsewhere, as well as similar hominin percussive behavior in the Early Stone Age. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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 nut-cracking has been frequently inferred.

Objectives Chimpanzee termite fishing has been studied for decades, yet the selective processes preceding the manufacture of fishing tools remain largely unexplored. We investigate raw material selection and potential evidence of forward planning in the chimpanzees of Issa valley, western Tanzania. Materials and Methods Using traditional archaeological methods, we surveyed the location of plants from where chimpanzees sourced raw material to manufacture termite fishing tools, relative to targeted mounds. We measured raw material abundance to test for availability and selection. Statistics included Chi-Squared, two-tailed Wilcoxon, and Kruskall-Wallace tests. Results Issa chimpanzees manufactured extraction tools only from bark, despite availability of other suitable materials (e.g., twigs), and selected particular plant species as raw material sources, which they often also exploit for food. Most plants were sourced 1-16 m away from the mound, with a maximum of 33 m. The line of sight from the targeted mound was obscured for a quarter of these plants. Discussion The exclusive use of bark tools despite availability of other suitable materials indicates a possible cultural preference. The fact that Issa chimpanzees select specific plant species and travel some distance to source them suggests some degree of selectivity and, potentially, forward planning. Our results have implications for the reconstruction of early hominin behaviors, particularly with regard to the use of perishable tools, which remain archaeologically invisible. [ABSTRACT FROM AUTHOR]

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.

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. [ABSTRACT FROM AUTHOR]

Cláudia Sousa was a founding member of the newly emerging and interdisciplinary field of primate archaeology. This discipline employs an archaeological approach to the study of nonhuman primates through the observation of their material culture - i. e. objects produced during tool use and manipulation - and has significant implications in the study of human evolution. [ABSTRACT FROM AUTHOR]

The concept of extended or distributed cognition has been present in archaeology for some time, yet despite its inclusion of non-human hominin ancestors, it has remained distinctly anthropocentric in nature. Here, we suggest that the same concept may also be used to independently describe and interpret non-human animals within their own social and material networks. We illustrate this suggestion with examples from the tool use behaviour of wild monkeys and chimpanzees. Non-human primate social groups develop bodies of traditional knowledge, and we consider whether idiosyncratic expression of such knowledge may be viewed in terms of an individual's constructed social identity. At a micro-level, the performance of an individual tool use technique may be analogous to the idea of ‘personhood’ found in anthropological holistic or perspectivist theory; at a macro-level the physical and social distribution of primate technology is amenable to interpretation as an example of extended or distributed cognition. We conclude that combined consideration of extended cognition and niche construction offers a promising means for interpreting the material residues of non-human primate behaviour. [ABSTRACT FROM AUTHOR]

Recovering evidence of past human activities enables us to recreate behaviour where direct observations are missing. Here, we apply archaeological methods to further investigate cultural transmission processes in percussive tool use among neighbouring chimpanzee communities in the Taï National Park, Côte d'Ivoire, West Africa. Differences in the selection of nut-cracking tools between neighbouring groups are maintained over time, despite frequent female transfer, which leads to persistent cultural diversity between chimpanzee groups. Through the recovery of used tools in the suggested natal territory of immigrants, we have been able to reconstruct the tool material selection of females prior to migration. In combination with direct observations of tool selection of local residents and immigrants after migration, we uncovered temporal changes in tool selection for immigrating females. After controlling for ecological differences between territories of immigrants and residents our data suggest that immigrants abandoned their previous tool preference and adopted the pattern of their new community, despite previous personal proficiency of the same foraging task. Our study adds to the growing body of knowledge on the importance of conformist tendencies in animals. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The last common ancestor (LCA) shared by chimpanzees ( Pan troglodytes) and bonobos ( P. paniscus) was an Early Pleistocene African ape, which, based on the behavior of modern chimpanzees, may be assumed to be a tool-using animal. However, the character of tool use in the Pan lineage prior to the 20th century is largely unknown. Here, I use available data on wild bonobo tool use and emerging molecular estimates of demography during Pan evolution to hypothesise the plausible tool use behavior of the bonobo-chimpanzee LCA (or ' Pancestor') at the start of the Pleistocene, over 2 million years ago. This method indicates that the common ancestor of living Pan apes likely used plant tools for probing, sponging, and display, but it did not use stone tools. Instead, stone tool use appears to have been independently invented by Western African chimpanzees ( P. t. verus) during the Middle Pleistocene in the region of modern Liberia-Ivory Coast-Guinea, possibly as recently as 200,000-150,000 years ago. If this is the case, then the LCA of humans and chimpanzees likely also did not use stone tools, and this trait probably first emerged among hominins in Pliocene East Africa. This review also suggests that the consistently higher population sizes of Central African chimpanzees ( P. t. troglodytes) over the past million years may have contributed to the increased complexity of wild tool use seen in this sub-species today. Am. J. Primatol. 76:910-918, 2014. © 2014 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

Some chimpanzee populations prey upon army ants, usually with stick tools. However, how their prey's subterranean nesting and nomadic lifestyle influence the apes' harvesting success is still poorly understood. This is particularly true for chimpanzees ( Pan troglodytes ellioti) at Gashaka/Nigeria, which consume army ants ( Dorylus rubellus) with much higher frequency than at other sites. We assessed various harvesting and search options theoretically available to the apes. For this, we reconstructed annual consumption patterns from feces and compared the physical characteristics of exploited ant nests with those that were not targeted. Repeated exploitation of a discovered nest is viable only in the short term, as disturbed colonies soon moved to a new site. Moreover, monitoring previously occupied nest cavities is uneconomical, as ants hardly ever re-used them. Thus, the apes have to detect new nests regularly, although colony density is relatively low (1 colony/1.3 ha). Surprisingly, visual search cues seem to be of limited importance because the probability of a nest being exploited was independent of its conspicuousness (presence of excavated soil piles, concealing leaf-litter or vegetation). However, chimpanzees preferentially targeted nests in forests or at the base of food trees, that is, where the apes spend relatively more time and/or where ant colony density is highest. Taken together, our findings suggest that, instead of employing a search strategy based on visual cues or spatial memory, chimpanzee predation on army ants contains a considerable opportunistic element. Am. J. Primatol. 75:1231-1244, 2013. © 2013 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

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., (© 2022 The Authors.)

Abstract: Savanna chimpanzees are known to re-use areas of the landscape for sleep, and patterns of chimpanzee sleeping site re-use are proposed as a referential model for early hominin archaeological site formation. We recorded the prevalence of deformed but healed branches and remnants of dead branches found around fresh nests at the savanna site of Issa in Ugalla, Tanzania. These old nest scars were found in 79% of 112 beds. We also randomly selected potential nesting locations for a subset of 32 beds within the same trees, and found nest scars in only 19% of these “control” locations. We then monitored 275 nests for up to 19 months for decay, regeneration of new branches, and re-use. Of these 275 nest locations, 24% were re-used within the first nine months of monitoring, and most re-use occurred when the nest had already decayed and was not easily visible from the ground. After 18 months, the proportion of specific nest positions re-used increased to 48%. This fidelity is likely a result of the creation of ideally-shaped support structures and supple new growth for mattress material with successive use of nest locations. We propose that specific nest site re-use may not be a direct product of environmental determination, but a result of “niche construction” through formation of good building sites within trees. Environmental modification through construction behaviour may have influenced both chimpanzee and early hominin ranging, and thus leaves behind recognisable patterns of artefact deposition across the landscape. [Copyright &y& Elsevier]

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.

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., Competing Interests: LL, MG, TP, MS, LK, SM No competing interests declared, (© 2019, Luncz et al.)

Chimpanzees are well‐studied, but raw material acquisition for tool use is still poorly understood as sources are difficult to trace. This study pioneers the use of information that can be gleaned from plant scars made by chimpanzees while they source vegetation parts to manufacture termite fishing tools. Source plant species, raw material types and locations relative to targeted termite mounds were recorded for populations at Gombe, Issa, and Mahale in western Tanzania. Recovered bark, twig, and vine tools were traced to 29 plant species, while grass sources were indeterminable. Bark extraction scars remained detectable for months, and thus possibly for as long as the plant is alive, while twig and vine scars preserved for a few weeks only. Scars preserve better than tools, given that twice as many plant species could be linked to the former than to the latter. Some source species were exploited across all sites for the same type of tool material, while two species were sourced for different types. Compared to apes at Gombe and Mahale, Issa chimpanzees carried material from twice as far away, perhaps because the Issa habitat is more open and dry, which entails greater distances between suitable raw material sources and targeted mounds. Site‐specific tools were based on different raw materials, in two cases sourced from the same species, which could suggest learned preferences for particular tool material. "Archaeology of the perishable" as pioneered in this study broadens the methodological approach of the wider field of primate archaeology to include reconstructions of past animal behavior associated with the production of plant based tools. The sourcing of raw material for termite fishing tools by chimpanzees leaves a record on the source plant – scars – that can be studied long after the time of their procurement. Different chimpanzee populations living in western Tanzania used varied tool materials, suggesting preferences for types of tool material used. Apes living in a more open, drier environment, carried raw material for almost double the distance compared to apes living in closer and denser environments. [ABSTRACT FROM AUTHOR]

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 nut-cracking has been frequently inferred., Competing Interests: We declare we have no competing interests.