Distribution of potential suitable hammers and transport of hammer tools and nuts by wild capuchin monkeys

Selection and transport of objects to use as tools at a distant site are considered to reflect planning. Ancestral humans transported tools and tool-making materials as well as food items. Wild chimpanzees also transport selected hammer tools and nuts to anvil sites. To date, we had no other examples of selection and transport of stone tools among wild nonhuman primates. Wild bearded capuchins (Cebus libidinosus) in Boa Vista (Piauí, Brazil) routinely crack open palm nuts and other physically well-protected foods on level surfaces (anvils) using stones (hammers) as percussive tools. Here we present indirect evidence, obtained by a transect census, that stones suitable for use as hammers are rare (study 1) and behavioral evidence of hammer transport by twelve capuchins (study 2). To crack palm nuts, adults transported heavier and harder stones than to crack other less resistant food items. These findings show that wild capuchin monkeys selectively transport stones of appropriate size and hardness to use as hammers, thus exhibiting, like chimpanzees and humans, planning in tool-use activities.     

Wild capuchin monkeys (Cebus libidinosus) use anvils and stone pounding tools

We conducted an exploratory investigation in an area where nut-cracking by wild capuchin monkeys is common knowledge among local residents. In addition to observing male and female capuchin monkeys using stones to pound open nuts on stone "anvils," we surveyed the surrounding area and found physical evidence that monkeys cracked nuts on rock outcrops, boulders, and logs (collectively termed anvils). Anvils, which were identified by numerous shallow depressions on the upper surface, the presence of palm shells and debris, and the presence of loose stones of an appropriate size to pound nuts, were present even on the tops of mesas. The stones used to crack nuts can weigh >1 kg, and are remarkably heavy for monkeys that weigh <4 kg. The abundance of shell remains and depressions in the anvil surface at numerous anvil sites indicate that nut-cracking activity is common and long-enduring. Many of the stones found on anvils (presumably used to pound nuts) are river pebbles that are not present in the local area we surveyed (except on or near the anvils); therefore, we surmise that they were transported to the anvil sites. Ecologically and behaviorally, nut-cracking by capuchins appears to have strong parallels to nut-cracking by wild chimpanzees. The presence of abundant anvil sites, limited alternative food resources, abundance of palms, and the habit of the palms in this region to produce fruit at ground level all likely contribute to the monkeys' routine exploitation of palm nuts via cracking them with stones. This discovery provides a new reference point for discussions regarding the evolution of tool use and material culture in primates. Routine tool use to exploit keystone food resources is not restricted to living great apes and ancestral hominids.     

Stone tool use by adult wild bearded capuchin monkeys (Cebus libidinosus). Frequency, efficiency and tool selectivity

Chimpanzees have been the traditional referential models for investigating human evolution and stone tool use by hominins. We enlarge this comparative scenario by describing normative use of hammer stones and anvils in two wild groups of bearded capuchin monkeys (Cebus libidinosus) over one year. We found that most of the individuals habitually use stones and anvils to crack nuts and other encased food items. Further, we found that in adults (1) males use stone tools more frequently than females, (2) males crack high resistance nuts more frequently than females, (3) efficiency at opening a food by percussive tool use varies according to the resistance of the encased food, (4) heavier individuals are more efficient at cracking high resistant nuts than smaller individuals, and (5) to crack open encased foods, both sexes select hammer stones on the basis of material and weight. These findings confirm and extend previous experimental evidence concerning tool selectivity in wild capuchin monkeys ( [Visalberghi et?al., 2009b] and [Fragaszy et?al., 2010b]).Male capuchins use tools more frequently than females and body mass is the best predictor of efficiency, but the sexes do not differ in terms of efficiency. We argue that the contrasting pattern of sex differences in capuchins compared with chimpanzees, in which females use tools more frequently and more skillfully than males, may have arisen from the degree of sexual dimorphism in body size of the two species, which is larger in capuchins than in chimpanzees. Our findings show the importance of taking sex and body mass into account as separate variables to assess their role in tool use.     

The physical characteristics and usage patterns of stone axe and pounding hammers used by long‐tailed macaques in the Andaman Sea region of Thailand

Stone hammering in natural conditions has been extensively investigated in chimpanzees and bearded capuchins. In contrast, knowledge of stone tool use in wild Old World monkeys has been limited to anecdotal reports, despite having known for over 120 years that Macaca fascicularis aurea use stone tools to process shelled foods from intertidal zones on islands in the Andaman Sea. Our report is the first scientific investigation to look at the stone tools used by these macaques. We observed they were skilled tool users and used stone tools daily. They selected tools with differing qualities for differing food items, and appeared to use at least two types of stone tools. Pounding hammers were used to crush shellfish and nuts on anvils and axe hammers were used to pick or chip at oysters attached to boulders or trees. We found significant physical differences between these two tools. Tools at oyster beds were smaller and exhibited scarring patterns focused more often on the points, whereas tools found at anvils were larger and showed more scarring on the broader surfaces. We also observed grip differences between the two tool types. Lastly, macaques struck targets with axe hammers more rapidly and over a wider range of motion than with pounding hammers. Both our behavioral and lithic data support that axe hammers might be used with greater control and precision than pounding hammers. Hand‐sized axe hammers were used for controlled chipping to crack attached oysters, and larger pounding hammers were used to crush nuts and unattached shellfish on anvils. In addition to stones, they also used hand‐sized auger shells (Turritella attenuata) as picks to axe attached oysters. Pound hammering appears similar to the stone tools used by chimpanzees and capuchins, but axe hammering has not yet been documented in other nonhuman primates in natural conditions. Am. J. Primatol. 71:594–608, 2009. © 2009 Wiley‐Liss, Inc.     

New data on the use of stone tools by chimpanzees in Guinea and Liberia

Two types of use of “hammers” for cracking nuts by wild-living chimpanzees have been distinguished: (1) Relatively small stones are used by the chimpanzee community at Bossou in Guinea to crack the nuts of oil palms growing on abandoned farmland, while no nuts of wild tree species are cracked. (2) Larger hammer stones (and, at some sites, wooden clubs) are used in a more sophisticated manner to crack the nuts of wild trees, but not of oil palms, in an area ranging from south-east Sierra Leone through Liberia to the south-west of the Ivory Coast. The first author (1986) has proposed that Type I has been copied by the chimpanzees, under pressure of food shortage, from the local human population. New data now indicate that, at Bossou, while habitat deterioration has continued, the number of hammer and anvil stones per utilized oil palm tree has approximately tripled in the last six years. The quantity of food obtained from oil palm nut kernels, however, amounts to only a few percent of the total diet. For the rest these apes depend to a large extent on many other agricultural products cultivated at Bossou which they are allowed freely to consume, including even cassava (manioc) roots and sweet potatoes dug by them from the ground. Some factors determining the chosen size of hammers were analyzed. Two abnormal hammers were found whose wear suggested a tentative, human-like manner of use. No evidence has been found to indicate the use of stone tools by chimpanzees in the adjoining chimpanzee-inhabited areas around the range of the Bossou community. Type II stone tool use was found, however, in a primary forest on a mountain≈13 km west of Bossou. This is especially intriguing because the site is separated by a wide belt of drier rain forest from the belt of very humid rain forest in the south where all the other known Type II sites are located. More research on the geographical distribution of the use of stone tools by chimpanzees and on the underlying ecological factors is recommended.     

Stone Anvil Damage by Wild Bearded Capuchins (Sapajus libidinosus) during Pounding Tool Use: A Field Experiment

We recorded the damage that wild bearded capuchin monkeys (Sapajus libidinosus) caused to a sandstone anvil during pounding stone tool use, in an experimental setting. The anvil was undamaged when set up at the Fazenda Boa Vista (FBV) field laboratory in Piauí, Brazil, and subsequently the monkeys indirectly created a series of pits and destroyed the anvil surface by cracking palm nuts on it. We measured the size and rate of pit formation, and recorded when adult and immature monkeys removed loose material from the anvil surface. We found that new pits were formed with approximately every 10 nuts cracked, (corresponding to an average of 38 strikes with a stone tool), and that adult males were the primary initiators of new pit positions on the anvil. Whole nuts were preferentially placed within pits for cracking, and partially-broken nuts outside the established pits. Visible anvil damage was rapid, occurring within a day of the anvil''s introduction to the field laboratory. Destruction of the anvil through use has continued for three years since the experiment, resulting in both a pitted surface and a surrounding archaeological debris field that replicate features seen at natural FBV anvils.     

Adult and juvenile bearded capuchin monkeys handle stone hammers differently during nut-cracking

Wild bearded capuchin monkeys (Sapajus libidinosus) habitually use stone hammers to crack open palm nuts and seeds on anvils. This activity requires strength, balance, and precise movement of a large stone with respect to the item placed on an anvil. We explored how well young monkeys cope with these challenges by examining their behavior and the behavior of adults while they cracked palm nuts using a stone. Using video records, we compared actions of six juvenile (2–5 years) and six adult (7+ years) wild monkeys during their first 20 strikes with one unfamiliar ellipsoid, quartzite stone (540 g), and the outcomes of these strikes. Compared with adults, juveniles cracked fewer nuts, performed a more diverse set of exploratory actions, and less frequently placed one or both hands on top of the stone on the downward motion. Adults and juveniles displayed similar low frequencies of striking with a slanted trajectory, missing the nut, and losing control over the nut or stone after striking. These findings indicate that young monkeys control the trajectory of a stone adequately but that is not sufficient to crack nuts as effectively as adults do. Compared with juveniles, adults more quickly perceive how to grip the stone efficiently, and they are able to adjust their grip dynamically during the strike. Young monkeys develop expertise in the latter aspects of cracking nuts over the course of several years of regular practice, indicating that perceptual learning about these aspects of percussion occurs slowly. Juvenile and adult humans learning to use stones to crack nuts also master these features of cracking nuts very slowly.     

Why Don't Chimpanzees in Gabon Crack Nuts?

Some populations of wild chimpanzees (Pan troglodytes) use hammers and anvils of stone or wood to crack open nuts for food. Others do not. The aim of this study was to ask why one non-nut-cracking population, in the Lopé Reserve, Gabon, lacks this useful form of tool use. We tested 10 hypotheses: (1) nuts are absent; (2) nuts are few; (3) nuts are unsuitable; (4) hammers are absent; (5) hammers are unsuitable; (6) anvils are absent; (7) anvils are unsuitable; (8) nuts are displaced by better food items; (9) intelligence is insufficient; and (10) knowledge is insufficient. All but the last are clearly falsified, leaving by exclusion the likelihood that Lopé's chimpanzees lack the technology—knowledge of appropriate technique—to exploit this resource. Thus, the behavioral differences across populations of these apes are cultural and not environmentally dictated. This explanation is congruent with the distribution of chimpanzee nut-cracking across Africa.     

Optional tool use: The case of wild bearded capuchins (Sapajus libidinosus) cracking cashew nuts by biting or by using percussors

Tool use in humans can be optional, that is, the same person can use different tools or no tool to achieve a given goal. Strategies to reach the same goal may differ across individuals and cultures and at the intra‐individual level. This is the first experimental study at the intra‐individual level on the optional use of a tool in wild nonhuman primates. We investigated optional tool use by wild bearded capuchins (Sapajus libidinosus) of Fazenda Boa Vista (FBV; Piauí, Brazil). These monkeys habitually succeed in cracking open the mesocarp of dry cashew nuts (Anacardium spp.) by pounding them with stones and/or by biting. We assessed whether availability of a stone and resistance of the nut affected capuchins' choice to pound or to bite the nuts and their rates of success. Sixteen capuchins (1–16 years) received small and large dry cashew nuts by an anvil together with a stone (Stone condition) or without a stone (No‐Stone condition). In the Stone conditions, subjects used it to crack the nut in 89.1% (large nuts) and 90.1% (small nut) of the trials. Nut size significantly affected the number of strikes used to open it. Availability of the stone significantly increased the average percent of success. In the No‐Stone conditions, monkeys searched for and used other percussors to crack the nuts in 54% of trials. In all conditions, age affects percentage of success and number of strikes to reach success. We argue that exclusive use of stones in other sites may be due to the higher abundance of stones at these sites compared with FBV. Since capuchins opened cashews with a tool 1–2 years earlier than they succeed at cracking more resistant palm nuts, we suggest that success at opening cashew nuts with percussors may support the monkeys' persistent efforts to crack palm nuts.     

How bearded capuchin monkeys (Sapajus libidinosus) prepare to use a stone to crack nuts

Bearded capuchin monkeys crack nuts with naturally varying stone hammers, suggesting they may tune their grips and muscular forces to each stone. If so, they might use discrete actions on a stone before lifting and striking, and they would likely use these actions more frequently when the stone is larger and/or less familiar and/or when first initiating striking. We examined the behavior of (a) four monkeys (all proficient at cracking nuts) with two larger (1 kg) and two smaller (0.5 kg) stones, (b) 12 monkeys with one 1 kg stone, and (c) one monkey during its first 100 strikes with an initially unfamiliar 1 kg stone. Bearded capuchin monkeys used three discrete actions on the stone before striking, all more often with the larger stones than the smaller stones. We infer that the first discrete action (Spin) aided the monkey in determining where to grip the stone, the second (Flip) allowed it to position the stone on the anvil ergonomically before lifting it, and the third (Preparatory Lift) readied the monkey for the strenuous lifting action. The monkey that provided 100 strikes with one initially unfamiliar stone performed fewer Spins in later strikes but performed Flip and Preparatory Lift at consistent rates. The monkeys gripped the stone with both hands along the sides to lift it, but usually moved one or both hands to the top of the stone at the zenith of the lift for the downward strike. The findings highlight two new aspects of the capuchins’ nut‐cracking: (a) Anticipatory actions with the stone before striking, especially when the stone is larger or unfamiliar, and when initiating striking and (b) shifting grips on the stone during a strike. We invite researchers to investigate if other taxa use anticipatory actions and shift their grips during percussive activity.     

Kinematics and energetics of nut-cracking in wild capuchin monkeys (Cebus libidinosus) in Piauí, Brazil

Wild bearded capuchins (Cebus libidinosus, quadrupedal, medium-sized monkeys) crack nuts using large stones. We examined the kinematics and energetics of the nut-cracking action of two adult males and two adult females. From a bipedal stance, the monkeys raised a heavy hammer stone (1.46 and 1.32 kg, from 33 to 77% of their body weight) to an average height of 0.33 m, 60% of body length. Then, they rapidly lowered the stone by flexing the lower extremities and the trunk until the stone contacted the nut. A hit consisting of an upward phase and a downward phase averaged 0.74 s in duration. The upward phase lasted 69% of hit duration. All subjects added discernable energy to the stone in the downward phase. The monkeys exhibited individualized kinematic strategies, similar to those of human weight lifters. Capuchins illustrate that human-like bipedal stance and large body size are unnecessary to break tough objects from a bipedal position. The phenomenon of bipedal nut-cracking by capuchins provides a new comparative reference point for discussions of percussive tool use and bipedality in primates.     

First GIS Analysis of Modern Stone Tools Used by Wild Chimpanzees (Pan troglodytes verus) in Bossou,Guinea, West Africa

Stone tool use by wild chimpanzees of West Africa offers a unique opportunity to explore the evolutionary roots of technology during human evolution. However, detailed analyses of chimpanzee stone artifacts are still lacking, thus precluding a comparison with the earliest archaeological record. This paper presents the first systematic study of stone tools used by wild chimpanzees to crack open nuts in Bossou (Guinea-Conakry), and applies pioneering analytical techniques to such artifacts. Automatic morphometric GIS classification enabled to create maps of use wear over the stone tools (anvils, hammers, and hammers/ anvils), which were blind tested with GIS spatial analysis of damage patterns identified visually. Our analysis shows that chimpanzee stone tool use wear can be systematized and specific damage patterns discerned, allowing to discriminate between active and passive pounders in lithic assemblages. In summary, our results demonstrate the heuristic potential of combined suites of GIS techniques for the analysis of battered artifacts, and have enabled creating a referential framework of analysis in which wild chimpanzee battered tools can for the first time be directly compared to the early archaeological record.     

Wild Bearded Capuchin Monkeys (Sapajus libidinosus) Strategically Place Nuts in a Stable Position during Nut-Cracking

Humans can use hand tools smoothly and effectively in varying circumstances; in other words, skillfully. A few other species of primates crack encased foods using hammer tools and anvils. Are they skilled? Positioning the food on the anvil so that it does not fall off when struck is a component of skilled cracking. We discovered that bearded capuchin monkeys deliberately place palm nuts in a relatively stable position on the anvil before striking them. In the first experiment, we marked the meridians of palm nuts where they stopped when rolled on a flat surface (“Stop meridian”). We videotaped monkeys as they cracked these nuts on an anvil. In playback we coded the position of the Stop meridian prior to each strike. Monkeys typically knocked the nuts on the anvil a few times before releasing them in a pit. They positioned the nuts so that the Stop meridian was within 30 degrees of vertical with respect to gravity more often than expected, and the nuts rarely moved after the monkeys released them. In the second experiment, 14 blindfolded people (7 men) asked to position marked nuts on an anvil as if to crack them reliably placed them with the Stop meridian in the same position as the monkeys did. In the third experiment, two people judged that palm nuts are most bilaterally symmetric along a meridian on, or close to, the Stop meridian. Thus the monkeys reliably placed the more symmetrical side of the nuts against the side of the pit, and the nuts reliably remained stationary when released. Monkeys apparently used information gained from knocking the nut to achieve this position. Thus, monkeys place the nuts skillfully, strategically managing the fit between the variable nuts and pits in the anvil, and skilled placement depends upon information generated by manual action.     

The evolution and cultural transmission of percussive technology: integrating evidence from palaeoanthropology and primatology

We bring together the quite different kinds of evidence available from palaeoanthropology and primatology to better understand the origins of Plio-Pleistocene percussive technology. Accumulated palaeoanthropological discoveries now document the Oldowan Complex as the dominant stone tool making culture between 2.6–1.4 Ma, the earlier part of this contemporaneous with pre-Homo hominins. The principal types of artefacts and other remains from 20 Early Stone Age (Oldowan and earliest Acheulean) localities in Africa and elsewhere are reviewed and described. To better understand the ancestral behavioural foundations of this early lithic culture, we examine a range of recent findings from primatology. In particular, we attempt to identify key shared characteristics of Homo and Pan that support inferences about the preparedness of our common ancestor for the innovation of stone tool culture. Findings of particular relevance include: (i) the discovery of an expanding repertoire of percussive and other tool use based on directed use of force among wild chimpanzees, implicating capacities that include significant innovatory potential and appreciation of relevant causal factors; (ii) evidence of material cultural diversity among wild chimpanzees, indicating a readiness to acquire and transmit tool use innovations; and (iii) experimental studies of social learning in chimpanzees and bonobos that now encompass the acquisition of nut cracking through observation of skilled use of hammers and anvils by conspecifics, the diffusion within and between groups of alternative styles of tool use, and the adoption of free-hand stone-to-stone percussion to create useful flakes for cutting to gain access to food resources. We use the distributions of the inferred cultural traits in the wild to assess how diffusion relates to geographic distances, and find that shared traits drop by 50% from the approximately eight characteristic of close neighbours over a distance of approximately 700 km. This pattern is used to explore the implications of analogous processes operating in relation to Early Stone Age sites.     

Mental map in wild chimpanzees: An analysis of hammer transports for nut cracking

The mental map of wild chimpanzees is analyzed in the context of their transports of clubs and stones used for cracking two species of nuts of different hardness,Coula edulis andPanda oleosa, in the Tai National Park (Ivory Coast). For the harderPanda nuts, they transport the harder hammers, i.e., almost exclusively stones, hammers of greater weight, and the transports are longer than forCoula nuts. The analysis made for the transports forPanda nuts shows that they seem to remember the location of stones and to choose the stones so as to keep the transport distance minimal. The chimpanzees seem to possess an Euclidian space, which allows them to somehow measure and remember distances; to compare several such distances so as to choose the stone with the shortest distance to a goal tree; to correctly locate a new stone location with reference to different trees; and to change their reference point so as to measure the distance to eachPanda tree from any stone location. They also combine the weight and the distance. The wild chimpanzees of the Tai National Park seem to possess concrete operation abilities in spatial representation.     

Physical properties of palm fruits processed with tools by wild bearded capuchins (Cebus libidinosus)

Habitually, capuchin monkeys access encased hard foods by using their canines and premolars and/or by pounding the food on hard surfaces. Instead, the wild bearded capuchins (Cebus libidinosus) of Boa Vista (Brazil) routinely crack palm fruits with tools. We measured size, weight, structure, and peak-force-at-failure of the four palm fruit species most frequently processed with tools by wild capuchin monkeys living in Boa Vista. Moreover, for each nut species we identify whether peak-force-at-failure was consistently associated with greater weight/volume, endocarp thickness, and structural complexity. The goals of this study were (a) to investigate whether these palm fruits are difficult, or impossible, to access other than with tools and (b) to collect data on the physical properties of palm fruits that are comparable to those available for the nuts cracked open with tools by wild chimpanzees. Results showed that the four nut species differ in terms of peak-force-at-failure and that peak-force-at-failure is positively associated with greater weight (and consequently volume) and apparently with structural complexity (i.e. more kernels and thus more partitions); finally for three out of four nut species shell thickness is also positively associated with greater volume. The finding that the nuts exploited by capuchins with tools have very high resistance values support the idea that tool use is indeed mandatory to crack them open. Finally, the peak-force-at-failure of the piassava nuts is similar to that reported for the very tough panda nuts cracked open by wild chimpanzees; this highlights the ecological importance of tool use for exploiting high resistance foods in this capuchin species.     

Transport of Functionally Appropriate Tools by Capuchin Monkeys (Cebus apella)

Capuchin monkeys (Cebus sp.) are notable among New World monkeys for their widespread use of tools. Like chimpanzees, they use both hammer tools and insertion tools in the wild to acquire food that would be unobtainable otherwise. Recent evidence indicates that capuchins transport stones to anvil sites and use the most functionally efficient stones to crack nuts. We further investigated capuchins’ assessment of functionality by testing their ability to select a tool that was appropriate for two different tool‐use tasks: A stone for a hammer task and a stick for an insertion task. To select the appropriate tools, the monkeys investigated a baited tool‐use apparatus (insertion or hammer), traveled to a location in their enclosure where they could no longer see the apparatus, made a selection between two tools (stick or stone), and then could transport the tool back to the apparatus to obtain a walnut. We incorporated tool transport and the lack of a visual cue into the design to assess willingness to transport the tools and the monkeys’ memory for the proper tool. Six brown capuchins (Cebus apella) were first trained to select and use the appropriate tool for each apparatus. Four animals completed training and were then tested by allowing them to view a baited apparatus and then travel to a location 8 m distant where they could select a tool while out of view of the apparatus. All four monkeys chose the correct tool significantly more than expected and transported the tools back to the apparatus. Results confirm capuchins’ propensity for transporting tools, demonstrate their capacity to select the functionally appropriate tool for two different tool‐use tasks, and indicate that they can retain the memory of the correct choice during a travel time of several seconds.     

Percussive tool use by Ta? Western chimpanzees and Fazenda Boa Vista bearded capuchin monkeys: a comparison

Percussive tool use holds special interest for scientists concerned with human origins. We summarize the findings from two field sites, Taï and Fazenda Boa Vista, where percussive tool use by chimpanzees and bearded capuchins, respectively, has been extensively investigated. We describe the ecological settings in which nut-cracking occurs and focus on four aspects of nut-cracking that have important cognitive implications, namely selection of tools, tool transport, tool modification and modulation of actions to reach the goal of cracking the nut. We comment on similarities and differences in behaviour and consider whether the observed differences reflect ecological, morphological, social and/or cognitive factors. Both species are sensitive to physical properties of tools, adjust their selection of hammers conditionally to the resistance of the nuts and to transport distance, and modulate the energy of their strikes under some conditions. However, chimpanzees transport hammers more frequently and for longer distances, take into account a higher number of combinations of variables and occasionally intentionally modify tools. A parsimonious interpretation of our findings is that morphological, ecological and social factors account for the observed differences. Confirmation of plausible cognitive differences in nut-cracking requires data not yet available.