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

Use-wear and residue analysis of pounding tools used by wild capuchin monkeys (Sapajus libidinosus) from Serra da Capivara (Piauí, Brazil)

Bearded capuchin monkeys (Sapajus libidinosus) from Serra da Capivara National Park (Brazil), perform the widest range of activities using stone tools of all the non-human tool-using primates. The behaviours behind this range of tool-use have been closely documented, but little is known about the characteristics of the tools themselves. Here we redress this imbalance and adopt an archaeological perspective to the analysis of capuchin pounding tools. We apply, for the first time, systematic microscopic techniques to the analysis of capuchin stone tools used for digging, cracking cashew nuts and seed processing to characterise their damage patterns combined with residue spatial distribution and micro-remains analysis. This work presents a standardized methodology for future primate archaeological use-wear studies as well as forming a reference collection which can be used to identify different activities within the primate archaeological record. Furthermore, understanding the archaeologically visible traces of primate percussive behaviours represents an initial step in developing a methodology to investigate if similar activities were practiced by early hominins and to help identify these activities in the Plio-Pleistocene archaeological record

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

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

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

Object manipulation and tool use in Nicobar long-tailed macaques (Macaca fascicularis umbrosus)

This is an accepted manuscript of an article published by Springer in International Journal of Primatology on 08/09/2020, available online: https://link.springer.com/article/10.1007%2Fs10764-020-00141-y The accepted version of the publication may differ from the final published version.Object manipulation and tool use by non-human primates have received considerable attention from primatologists and anthropologists, because of their broad implications for understanding the evolution of tool use in humans. To date, however, most of the studies on this topic have focused on apes, given their close evolutionary relationship with humans. In contrast, fewer studies on tool use and object manipulation have been conducted on monkeys. Documenting and studying object manipulation and tool use in species that are more distantly related to humans can provide a broader perspective on the evolutionary origins of this behaviour. We present a detailed description of toolaided behaviours and object manipulation by Nicobar long-tailed macaques ( Macaca fascicularis umbrosus ) living along the coastlines of Great Nicobar Island. We made observations from December 2018 to March 2019, using ad libitum and focal sampling methods. We observed behaviours related to object manipulation and tool use in six different behavioural contexts (foraging, hygiene, communication, play, selfdirected and self-hygiene behaviour) involving eight different types of objects, namely resonance rod, play object, rolling platform, scraping tool, dental groom, pounding substrate, leaves as grip pads and wipers, and stimulation tool. We observed that males were involved in tool use and object manipulation more frequently than females. Our results add to existing records of object manipulation, tool-use behaviour and tool variants displayed by non-human primates, showing that Nicobar macaques perform multiple and diverse tool-aided behaviours

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