Mapping nut-cracking in a new population of wild capuchin monkeys (Sapajus libidinosus) at Ubajara National Park, Brazil.

Populations of bearded capuchin monkeys (Sapajus libidinosus) vary in their tool use behaviors, with some of this divergence regarded as culturally determined. The use of stone tools, primarily to crack open encased foods, is widespread among bearded capuchins living in dry habitats (Caatinga and Cerrado). Significant diversity in targets, processed foods, material, and size of tools is observed across populations. However, so far, only a few sites have been systematically studied, and we are still distant from a representative picture of the range of variation in capuchins' culture. In this study, we did a systematic assessment of stone tool use sites in the Ubajara National Park (UNP), in the Caatinga region of Ceará, Brazil, recording and measuring stone tools, processed foods, and available lithic resources as part of an extensive comparative research, the CapCult project. We found indirect and direct evidence that capuchin monkeys at UNP customarily use hammerstones and anvils to process at least two species of palm nuts, macauba (Acrocomia aculeata) and the harder babaçu (Attalea speciosa). Most of the anvils were rock surfaces and had leftovers of only one palm nut species. The hammerstones used to process both palm nuts were not significantly different in weight, although the ones used for Ac. aculeata were longer. We found a higher frequency of nut-cracking sites in the drier lowland area of the park, reflecting differences in the density of the most common palm species, Ac. aculeata, and availability of raw stone material. The stone tool use observed in UNP is within the scope of previously reported in savannah capuchin populations. Our study widens the knowledge of stone tool-use diversity in wild capuchin monkeys, which could contribute to shaping conservation policy, including cultural traits.

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.

Western chimpanzees (Pan troglodytes verus) access a nutritionally balanced, high energy, and abundant food, baobab (Adansonia digitata) fruit, with extractive foraging and reingestion.

Intrinsic to several hypotheses explaining the evolution of foraging behavior complexity, such as proto-tool use, is the assumption that more complex ingestive behaviors are adaptations allowing individuals to access difficult to procure but nutritionally or energetically rewarding foods. However, nutritional approaches to understanding this complexity have been underutilized. The goal of this study was to evaluate potential nutritional determinants of two unusual foraging behaviors, fruit cracking with anvils and seed reingestion, by adult male western chimpanzees (Pan troglodytes verus) at Fongoli, Senegal during the baobab (Adansonia digitata) fruit season. We examined these behaviors in relation to nutrient and energy intake, and compared macronutrient and energy concentrations found in baobab fruits to other plant foods. Adult males ingested at least 31 distinct foods from 23 plant species. Baobab fruit comprised the majority of daily energy intake (68 ± 34%, range: 0%-98%). The energetic concentration of baobab fruit varied by phenophase and part ingested, with ripe and semi-ripe fruit ranking high in energy return rate. Males preferred ripe and semi-ripe baobab fruit but unripe fruit intake was higher overall. The seed kernels were high in protein and fat relative to fruit pulp, and these kernels were easier to access during the unripe stage. During the ripe stage, seed kernels were accessible by reingestion, after the seed coat was softened during gut passage. In addition to providing macronutrients and energy, baobab fruit was a relatively abundant food source. We conclude that baobab pulp and seed are high quality foods at Fongoli during the baobab season because they are nutritionally balanced, high in energy, and relatively abundant in the environment. These nutritional and abundance characteristics may explain, in part, why these chimpanzees use anvils and reingestion to access a mechanically challenging food.

Bili-Uéré: A Chimpanzee Behavioural Realm in Northern Democratic Republic of Congo.

To improve our understanding of the evolutionary origins of culture and technology in humans, it is vital that we document the full extent of behavioural diversity in our great ape relatives. About half of the world's remaining chimpanzees (Pan spp.) live in the Democratic Republic of Congo (DRC), yet until now we have known almost nothing about their behaviour. Here we describe the insect-related tool technology of Bili-Uéré chimpanzees (Pan troglodytes schweinfurthii) inhabiting an area of at least a 50,000-km2 area in northern DRC, as well as their percussive technology associated with food processing. Over a 12-year period, we documented chimpanzee tools and artefacts at 20 survey areas and gathered data on dung, feeding remains and sleeping nests. We describe a new chimpanzee tool kit: long probes used to harvest epigaeic driver ants (Dorylus spp.), short probes used to extract ponerine ants and the arboreal nests of stingless bees, wands to dip for D. kohli and stout digging sticks used to access underground meliponine bee nests. Epigaeic Dorylus tools were significantly longer than the other tool types, and D. kohli tools were significantly thinner. Tools classified as terrestrial honey-digging sticks were a significant predictor for brushed and blunted tool ends, consistent with their presumed use. We describe two potential new tool types, an "ant scoop" and a "fruit hammer." We document an extensive percussive technology used to process termite mounds of Cubitermes sp. and Thoracotermes macrothorax and hard-shelled fruits such as Strychnos, along with evidence of the pounding open of African giant snails and tortoises. We encountered some geographic variation in behaviour: we found honey-digging tools, long driver ant probes and fruit-pounding sites only to the north of the Uele River; there were more epigaeic Dorylus tools to the north and more ponerine ant tools to the south. We found no evidence of termite-fishing, despite the availability of Macrotermes muelleri mounds throughout the region. This lack of evidence is consistent with the results of dung washes, which revealed a substantial proportion of driver ants, but no evidence of Macrotermes or other termites. Our results allow us to describe a new chimpanzee behavioural complex, characterised by a general similarity of multiple behaviours across a large, ecologically diverse region but with subtle differences in prey choice and techniques. We propose that this widespread and related suite of behaviours be referred to as the Bili-Uéré Chimpanzee Behavioural Realm. Possible explanations for this pattern are a recent chimpanzee expansion across the region and the interconnectedness of this population across at least the entirety of northern DRC.

Revisiting Panda 100, the first archaeological chimpanzee nut-cracking site.

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.

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.

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.

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.

Stone tool use by wild capuchin monkeys (Sapajus libidinosus) at Serra das Confusões National Park, Brazil.

Capuchin monkeys (Sapajus spp.) are proficient tool users, and the use of stone tools occurs in several populations, mostly to crack open encased foods. Two well-studied Brazilian populations of Sapajus libidinosus inhabit Fazenda Boa Vista and Serra da Capivara National Park and present different behavioral sets regarding tool use. Serra das Confusões National Park (SCoNP) lies between those sites, but little is known about the capuchin monkey population that lives there. To begin unraveling the capuchin behavior in this area, we conducted a brief survey for tool use sites. We found indirect evidence that capuchin monkeys at SCoNP use stone hammers to crack open at least four species of seeds and fruits. Plant reproductive parts there are processed with stone tools in a similar pattern to the other sites. Further study is needed to directly observe tool use by capuchin monkeys at SCoNP, verify the occurrence of other possible types of tool use in this population, and thus fully compare their tool use repertoire to that of other populations.

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 nut-cracking has been frequently inferred.

How similar are nut-cracking and stone-flaking? A functional approach to percussive technology.

Various authors have suggested similarities between tool use in early hominins and chimpanzees. This has been particularly evident in studies of nut-cracking which is considered to be the most complex skill exhibited by wild apes, and has also been interpreted as a precursor of more complex stone-flaking abilities. It has been argued that there is no major qualitative difference between what the chimpanzee does when he cracks a nut and what early hominins did when they detached a flake from a core. In this paper, similarities and differences between skills involved in stone-flaking and nut-cracking are explored through an experimental protocol with human subjects performing both tasks. We suggest that a 'functional' approach to percussive action, based on the distinction between functional parameters that characterize each task and parameters that characterize the agent's actions and movements, is a fruitful method for understanding those constraints which need to be mastered to perform each task successfully, and subsequently, the nature of skill involved in both tasks.

Experimental studies illuminate the cultural transmission of percussive technologies in Homo and Pan.

The complexity of Stone Age tool-making is assumed to have relied upon cultural transmission, but direct evidence is lacking. This paper reviews evidence bearing on this question provided through five related empirical perspectives. Controlled experimental studies offer special power in identifying and dissecting social learning into its diverse component forms, such as imitation and emulation. The first approach focuses on experimental studies that have discriminated social learning processes in nut-cracking by chimpanzees. Second come experiments that have identified and dissected the processes of cultural transmission involved in a variety of other force-based forms of chimpanzee tool use. A third perspective is provided by field studies that have revealed a range of forms of forceful, targeted tool use by chimpanzees, that set percussion in its broader cognitive context. Fourth are experimental studies of the development of flint knapping to make functional sharp flakes by bonobos, implicating and defining the social learning and innovation involved. Finally, new and substantial experiments compare what different social learning processes, from observational learning to teaching, afford good quality human flake and biface manufacture. Together these complementary approaches begin to delineate the social learning processes necessary to percussive technologies within the Pan-Homo clade.

Limestone percussion tools from the late Early Pleistocene sites of Barranco León and Fuente Nueva 3 (Orce, Spain).

In recent years, there is growing interest in the study of percussion scars and breakage patterns on hammerstones, cores and tools from Oldowan African and Eurasian lithic assemblages. Oldowan stone toolkits generally contain abundant small-sized flakes and their corresponding cores, and are characterized by their structural dichotomy of heavy- and light-duty tools. This paper explores the significance of the lesser known heavy-duty tool component, providing data from the late Lower Pleistocene sites of Barranco León and Fuente Nueva 3 (Orce, Spain), dated 1.4-1.2 Myr. Using quantitative and qualitative data from the large-sized limestone industries from these two major sites, we present a new methodology highlighting their morpho-technological features. In the light of the results, we discuss the shortfalls of extant classificatory methods for interpreting the role of percussive technology in early toolkits. This work is rooted in an experimental program designed to reproduce the wide range of percussion marks observed on the limestone artefacts from these two sites. A visual and descriptive reference is provided as an interpretative aid for future comparative research. Further experiments using a variety of materials and gestures are still needed before the elusive traces yield the secrets of the kinds of percussive activities carried out by hominins at these, and other, Oldowan sites.

The development of plant food processing in the Levant: insights from use-wear analysis of Early Epipalaeolithic ground stone tools.

In recent years, the study of percussive, pounding and grinding tools has provided new insights into human evolution, more particularly regarding the development of technology enabling the processing and exploitation of plant resources. Some of these studies focus on early evidence for flour production, an activity frequently perceived as an important step in the evolution of plant exploitation. The present paper investigates plant food preparation in mobile hunter-gatherer societies from the Southern Levant. The analysis consists of a use-wear study of 18 tools recovered from Ohalo II, a 23 000-year-old site in Israel showing an exceptional level of preservation. Our sample includes a slab previously interpreted as a lower implement used for producing flour, based on the presence of cereal starch residues. The use-wear data we have obtained provide crucial information about the function of this and other percussive tools at Ohalo II, as well as on investment in tool manufacture, discard strategies and evidence for plant processing in the Late Pleistocene. The use-wear analysis indicates that the production of flour was a sporadic activity at Ohalo II, predating by thousands of years the onset of routine processing of plant foods.