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Platyrrhines consume many species of arthropods in the order Orthoptera. Some species of orthopterans can produce chemical defenses that render them toxic or unpalatable and thus act as predator deterrents. These species include the stick grasshoppers (family Proscopiidae), which are widely distributed in the Caatinga biome in northeastern Brazil, which comprises part of the distribution of capuchin monkeys. Capuchin monkeys are omnivores and consume a wide variety of foods, including unpleasant-tasting, potentially toxic items, which they need to learn how to process. We describe the processing of stick grasshoppers (Stiphra sp.) by wild capuchin monkeys (Sapajus libidinosus) that live in Serra da Capivara National Park, Brazil, and compare how individuals of different age classes handle these potentially toxic food items. S. libidinosus predominantly avoided consuming the digestive tract, which contains toxic compounds, when feeding on stick grasshoppers. Immatures took longer than adults to process the stick grasshoppers, indicating that capuchins need to learn how to process the toxic digestive tract of these prey to avoid consuming it.

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Sapajus libidinosus members of the Pedra Furada group, living in the Serra da Capivara National Park, use stone tools in a wider variety of behaviors than any other living animal, except humans. To rescue the evolutionary history of the Caatinga S. libidinosus and identify factors that may have contributed to the emergence and maintenance of their tool-use culture, we conducted fieldwork seasons to obtain biological samples of these capuchin monkeys. UsingCYTBsequences, we show a discrete but constant population growth from the beginning of the Holocene to the present, overlapping the emergence of the Caatinga biome. Our habitat suitability reconstruction reports the presence of plants whose hard fruits, seeds, or roots are processed by capuchins using tools. TheS. libidinosusindividuals in the Caatinga were capable of dynamically developing and maintaining their autochthonous culture thanks to: a) cognitive capacity to generate and execute innovation under selective pressure; b) tolerance favoring learning and cultural inheritance; c) an unknown genetic repertoire that underpins the adaptive traits; d) a high degree of terrestriality; e) presence and abundance of natural resources, which makes some places "hot spots" for innovation, and cultural diversification within a relatively short time.

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Terrestriality was an essential factor in human evolution. Hominins' extensive use of the ground allowed exploring a new range of environments and their objects, including new resources and potential tool raw materials. Capuchin monkeys are primarily arboreal primates but are also the most prolific tool users among platyrrhines, customarily using stone tools on the ground due to physical limitations and material availability. OBJECTIVES: Our goals were to (1) measure the terrestriality levels of a group of capuchin monkeys and (2) test the hypothesis that terrestriality has a positive effect on the stone tool use variability because the increased time on the ground would offer more opportunities to interact with the available stones, leading to more innovations of tool use behaviors. We predict a more diversified use of stone tools in the population with a higher degree of terrestriality. MATERIALS AND METHODS: This study was on a group of capuchin monkeys (Sapajus libidinosus) at Serra da Capivara National Park (SCNP), Brazil. Scan sampling was done for 23 months, recording the behavior and substrate used by the individuals. RESULTS: The ground use rate was 41%, with no sex difference but an age difference: infants were less terrestrial than juveniles and adults. DISCUSSION: Compared to a population with a more limited tool use repertoire (Fazenda Boa Vista), SCNP adult capuchins are more terrestrial (43% v. 27%). Stone tool use diversity and terrestriality in capuchins appear to be positively correlated. Our results support this hypothesis and provide terrestriality measurements of the wild capuchin population with the most complex stone tool kit.

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Examining interactions among sympatric primate species can provide interesting information about competition, cooperation, and avoidance between those species. Those interactions can be neutral, positive, or negative for the species involved. Capuchin monkeys are medium-sized primates that can encounter both larger and smaller primates in their varied habitats. Gracile capuchins (Cebus) are reported to present different types of interactions with other primates. Interactions with howler monkeys frequently include physical aggression, while interactions with spider monkeys are mostly threats and chases. Moreover, interaction types are not consistent across populations. Among robust capuchins (Sapajus spp.), however, no reports have been published. Here we describe and classify encounters of Sapajus libidinosus and S. nigritus with Alouatta caraya, A. guariba, Brachyteles arachnoides, and Callithrix jacchus in three sites in the environments of Cerrado, Caatinga (savannah-like), and Atlantic forest, and compare the interaction patterns among sites and different group sizes. The latter is a factor that can influence the outcome, and we expected capuchins in larger groups to be more aggressive toward other primates. Our results of 8421 h of total contact with the capuchin groups show that, indeed, capuchins in sites with larger groups presented aggressive interactions with higher frequency. However, the other species' body size also seems important as smaller primates apparently avoided capuchins, and interactions with the larger muriquis were mostly neutral for the capuchin. Capuchins showed neutral or aggressive behaviors toward howler monkeys, with differences between the rainforest and savannah groups. We found that robust capuchins can present aggressive interactions even to primates larger than themselves and that aggressive behavior was the most common response in populations living in larger groups and drier environments.

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Tufted capuchin monkeys (Sapajus spp.) are the only Neotropical Primates that regularly use tools in the wild, but only one population of bearded capuchin monkeys (Sapajus libidinosus) is known to habitually use sticks as probes. In this population, males are typically the only sex to use stick tools, something unexpected, since there are no obvious physical constraints, and females do use stone tools in the wild and sticks in experimental conditions. We investigated the development of probe tool use in eight infants to clarify whether social influences on learning varied between the sexes, as tool use learning by capuchin monkeys is a socially biased process. We found that in the first 10 months of age, females manipulate sticks as much as males, but after 10-12 months of age, males begin to manipulate them at higher frequencies. We examined if social connections-as opportunities for social learning-could explain this difference and verified that, on close distance social networks, infant males and females have similar connections with older males. However, males observe probe tool use events more often than females when close to such events. The higher frequency of manipulation of sticks, as well as the higher rates of probe tool use observation, appear to be the key to understand why only males are probe tool users in this population. Since there are only male potential models of probe use, a sex motivational bias could explain the sex difference in observation; a bias in observation could explain the differences in manipulation-and manipulation rates would certainly influence the chances of individual, trial-and-error learning (a case of "local/stimulus enhancement").

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The human archaeological record changes over time. Finding such change in other animals requires similar evidence, namely, a long-term sequence of material culture. Here, we apply archaeological excavation, dating and analytical techniques to a wild capuchin monkey (Sapajus libidinosus) site in Serra da Capivara National Park, Brazil. We identify monkey stone tools between 2,400 and 3,000 years old and, on the basis of metric and damage patterns, demonstrate that capuchin food processing changed between ~2,400 and 300 years ago, and between ~100 years ago and the present day. We present the first example of long-term tool-use variation outside of the human lineage, and discuss possible mechanisms of extended behavioural change.

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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.

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Snakes present a hazard to primates, both as active predators and by defensive envenomation. This risk might have been a selective pressure on the evolution of primate visual and cognitive systems, leading to several behavioral traits present in human and non-human primates, such as the ability to quickly learn to fear snakes. Primates seldom prey on snakes, and humans are one of the few primate species that do. We report here another case, the wild capuchin monkey (Sapajus libidinosus), which preys on snakes. We hypothesized that capuchin monkeys, due to their behavioral plasticity, and cognitive and visual skills, would be capable of discriminating dangerous and non-dangerous snakes and behave accordingly. We recorded the behavioral patterns exhibited toward snakes in two populations of S. libidinosus living 320 km apart in Piauí, Brazil. As expected, capuchins have a fear reaction to dangerous snakes (usually venomous or constricting snakes), presenting mobbing behavior toward them. In contrast, they hunt and consume non-dangerous snakes without presenting the fear response. Our findings support the tested hypothesis that S. libidinosus are capable of differentiating snakes by level of danger: on the one hand they protect themselves from dangerous snakes, on the other hand they take opportunities to prey on non-dangerous snakes. Since capuchins and humans are both predators and prey of snakes, further studies of this complex relationship may shed light on the evolution of these traits in the human lineage.

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Capuchin monkeys at Serra da Capivara National Park (SCNP) usually forage on the ground for roots and fossorial arthropods, digging primarily with their hands but also using stone tools to loosen the soil and aid the digging process. Here we describe the stone tools used for digging by two groups of capuchins on SCNP. Both groups used tools while digging three main food resources: Thiloa glaucocarpa tubers, Ocotea sp roots, and trapdoor spiders. One explanation for the occurrence of tool use in primates is the "necessity hypothesis", which states that the main function of tool use is to obtain fallback food. We tested for this, but only found a positive correlation between plant food availability and the frequency of stone tools' use. Thus, our data do not support the fallback food hypothesis for the use of tools to access burrowed resources.

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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.

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