Chimpanzees predict that a competitor's preference will match their own

The ability to predict how another individual will behave is useful in social competition. Chimpanzees can predict the behaviour of another based on what they observe her to see, hear, know and infer. Here we show that chimpanzees act on the assumption that others have preferences that match their own. All subjects began with a preference for a box with a picture of food over one with a picture of nothing, even though the pictures had no causal relation to the contents. In a back-and-forth food competition, chimpanzees then avoided the box with the picture of food when their competitor had chosen one of the boxes before them—presumably on the assumption that the competitor shared their own preference for it and had already chosen it. Chimpanzees predicted that their competitor''s preference would match their own and adjusted their behavioural strategies accordingly.     

Chimpanzees help conspecifics obtain food and non-food items

Chimpanzees (Pan troglodytes) sometimes help both humans and conspecifics in experimental situations in which immediate selfish benefits can be ruled out. However, in several experiments, chimpanzees have not provided food to a conspecific even when it would cost them nothing, leading to the hypothesis that prosociality in the food-provisioning context is a derived trait in humans. Here, we show that chimpanzees help conspecifics obtain both food and non-food items--given that the donor cannot get the food herself. Furthermore, we show that the key factor eliciting chimpanzees' targeted helping is the recipients' attempts to either get the food or get the attention of the potential donor. The current findings add to the accumulating body of evidence that humans and chimpanzees share the motivation and skills necessary to help others in situations in which they cannot selfishly benefit. Humans, however, show prosocial motives more readily and in a wider range of contexts.     

Use of tool sets by chimpanzees for multiple purposes in Moukalaba-Doudou National Park,Gabon

We report our recent findings on the use of tool sets by chimpanzees in Moukalaba-Doudou National Park, Gabon. Direct observations and evidences left by chimpanzees showed that chimpanzees used sticks as pounders, enlargers, and collectors to extract honey from beehives of stingless bees (Meliponula sp.), which may correspond to those previously found in the same site for fishing termites and to those found in Loango National Park, Gabon. However, we observed chimpanzees using a similar set of tools for hunting a medium-sized mammal (possibly mongoose) that hid inside a log. This is the first report of hunting with tools by a chimpanzee population in Central Africa. Chimpanzees may recognize the multiple functions and applicability of tools (extracting honey and driving prey), although it is still a preliminary speculation. Our findings may provide us a new insight on the chimpanzee’s flexibility of tool use and cognitive abilities of complex food gathering.     

Cognitive capacities for cooking in chimpanzees

The transition to a cooked diet represents an important shift in human ecology and evolution. Cooking requires a set of sophisticated cognitive abilities, including causal reasoning, self-control and anticipatory planning. Do humans uniquely possess the cognitive capacities needed to cook food? We address whether one of humans'' closest relatives, chimpanzees (Pan troglodytes), possess the domain-general cognitive skills needed to cook. Across nine studies, we show that chimpanzees: (i) prefer cooked foods; (ii) comprehend the transformation of raw food that occurs when cooking, and generalize this causal understanding to new contexts; (iii) will pay temporal costs to acquire cooked foods; (iv) are willing to actively give up possession of raw foods in order to transform them; and (v) can transport raw food as well as save their raw food in anticipation of future opportunities to cook. Together, our results indicate that several of the fundamental psychological abilities necessary to engage in cooking may have been shared with the last common ancestor of apes and humans, predating the control of fire.     

Tolerance allows bonobos to outperform chimpanzees on a cooperative task

To understand constraints on the evolution of cooperation, we compared the ability of bonobos and chimpanzees to cooperatively solve a food-retrieval problem. We addressed two hypotheses. The "emotional-reactivity hypothesis" predicts that bonobos will cooperate more successfully because tolerance levels are higher in bonobos. This prediction is inspired by studies of domesticated animals; such studies suggest that selection on emotional reactivity can influence the ability to solve social problems [1, 2]. In contrast, the "hunting hypothesis" predicts that chimpanzees will cooperate more successfully because only chimpanzees have been reported to cooperatively hunt in the wild [3-5]. We indexed emotional reactivity by measuring social tolerance while the animals were cofeeding and found that bonobos were more tolerant of cofeeding than chimpanzees. In addition, during cofeeding tests only bonobos exhibited socio-sexual behavior, and they played more. When presented with a task of retrieving food that was difficult to monopolize, bonobos and chimpanzees were equally cooperative. However, when the food reward was highly monopolizable, bonobos were more successful than chimpanzees at cooperating to retrieve it. These results support the emotional-reactivity hypothesis. Selection on temperament may in part explain the variance in cooperative ability across species, including hominoids.     

Chimpanzees spontaneously alter the location of their gestures to match the attentional orientation of others

Chimpanzees, Pan troglodytes, interacted with a familiar human partner across a table that contained a piece of highly desirable food and an undesirable distracter object. We explored how the chimpanzees' deployed their gestures on trials in which the food and object were out of their reach, thus creating a context in which they needed to recruit their partner's assistance in order to obtain the food. The attentional focus of their partner was directed either at the food, at the distracter object, or at a neutral location on the ceiling. Although the location of the food exerted primary control over the location of the chimpanzees' gestures, the results also clearly show that when their partner's focus was on the distracter object, the subjects modified their pattern of gesturing to accommodate the discrepancy between the object they wanted and the object on which their partner was focused. We discuss the results in the context of the evolution of joint and shared attention in humans, chimpanzees and other primates. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Cooperative problem solving in rooks (Corvus frugilegus)

Recent work has shown that captive rooks, like chimpanzees and other primates, develop cooperative alliances with their conspecifics. Furthermore, the pressures hypothesized to have favoured social intelligence in primates also apply to corvids. We tested cooperative problem-solving in rooks to compare their performance and cognition with primates. Without training, eight rooks quickly solved a problem in which two individuals had to pull both ends of a string simultaneously in order to pull in a food platform. Similar to chimpanzees and capuchin monkeys, performance was better when within-dyad tolerance levels were higher. In contrast to chimpanzees, rooks did not delay acting on the apparatus while their partner gained access to the test room. Furthermore, given a choice between an apparatus that could be operated individually over one that required the action of two individuals, four out of six individuals showed no preference. These results may indicate that cooperation in chimpanzees is underpinned by more complex cognitive processes than that in rooks. Such a difference may arise from the fact that while both chimpanzees and rooks form cooperative alliances, chimpanzees, but not rooks, live in a variable social network made up of competitive and cooperative relationships.     

The Relationship between Event-Based Prospective Memory and Ongoing Task Performance in Chimpanzees (Pan troglodytes)

Prospective memory is remembering to do something at a future time. A growing body of research supports that prospective memory may exist in nonhuman animals, but the methods used to test nonhuman prospective memory differ from those used with humans. The current work tests prospective memory in chimpanzees using a method that closely approximates a typical human paradigm. In these experiments, the prospective memory cue was embedded within an ongoing task. Tokens representing food items could be used in one of two ways: in a matching task with pictures of items (the ongoing task) or to request a food item hidden in a different location at the beginning of the trial. Chimpanzees had to disengage from the ongoing task in order to use the appropriate token to obtain a higher preference food item. In Experiment 1, chimpanzees effectively matched tokens to pictures, when appropriate, and disengaged from the ongoing task when the token matched the hidden item. In Experiment 2, performance did not differ when the target item was either hidden or visible. This suggested no effect of cognitive load on either the prospective memory task or the ongoing task, but performance was near ceiling, which may have contributed to this outcome. In Experiment 3, we created a more challenging version of the task. More errors on the matching task occurred before the prospective memory had been carried out, and this difference seemed to be limited to the hidden condition. This finding parallels results from human studies and suggests that working memory load and prospective memory may have a similar relationship in nonhuman primates.     

Image scoring in great apes

'Image scoring' occurs when person A monitors the giving behaviour of person B towards person C. We tested for 'image scoring' in chimpanzees, bonobos, gorillas, and orangutans. Subjects passively observed two types of incident: (i) a 'nice' person gave grapes to a human beggar, and (ii) a 'nasty' person refused to give. The subject witnessed both incidents in succession (but was unable to obtain the grapes). Shortly after, the ape had an opportunity to approach one or both human actors (nice/nasty), both of whom were now sitting side-by-side holding grapes. However, neither human offered their grapes if approached. The subject's expectation of which human was more likely to offer food was measured by comparing the proportion of time that subjects spent near each person. Chimpanzees (n=17) spent significantly more time at the 'nice' window compared to 'nasty'. Also, preference for 'nasty' declined as trials progressed. Results for other apes were not significant.     

Savanna chimpanzees, Pan troglodytes verus, hunt with tools

Although tool use is known to occur in species ranging from naked mole rats [1] to owls [2], chimpanzees are the most accomplished tool users [3-5]. The modification and use of tools during hunting, however, is still considered to be a uniquely human trait among primates. Here, we report the first account of habitual tool use during vertebrate hunting by nonhumans. At the Fongoli site in Senegal, we observed ten different chimpanzees use tools to hunt prosimian prey in 22 bouts. This includes immature chimpanzees and females, members of age-sex classes not normally characterized by extensive hunting behavior. Chimpanzees made 26 different tools, and we were able to recover and analyze 12 of these. Tool construction entailed up to five steps, including trimming the tool tip to a point. Tools were used in the manner of a spear, rather than a probe or rousing tool. This new information on chimpanzee tool use has important implications for the evolution of tool use and construction for hunting in the earliest hominids, especially given our observations that females and immature chimpanzees exhibited this behavior more frequently than adult males.     

Helping behaviour and regard for others in capuchin monkeys (Cebus apella)

Altruism is an evolutionary puzzle. To date, much debate has focused on whether helping others without regard to oneself is a uniquely human behaviour, with a variety of empirical studies demonstrating a lack of altruistic behaviour in chimpanzees even when the demands of behaving altruistically seem minimal. By contrast, a recent experiment has demonstrated that chimpanzees will help a human experimenter to obtain an out-of-reach object, irrespective of whether or not they are offered a reward for doing so, suggesting that the cognitions underlying altruistic behaviour may be highly sensitive to situational demands. Here, we examine the cognitive demands of other-regarding behaviour by testing the conditions under which primates more distantly related to humans--capuchin monkeys--help an experimenter to obtain an out-of-reach object. Like chimpanzees, capuchin monkeys helped human experimenters even in the absence of a reward, but capuchins systematically failed to take into account the perspective of others when they stood to obtain food for themselves. These results suggest an important role for perspective taking and inhibition in altruistic behaviour and seem to reflect a significant evolutionary development in the roots of altruism, and specifically in other-regarding behaviour, between the divergence of New World monkeys and apes.     

The evolutionary origins of human patience: temporal preferences in chimpanzees, bonobos, and human adults

To make adaptive choices, individuals must sometimes exhibit patience, forgoing immediate benefits to acquire more valuable future rewards [1-3]. Although humans account for future consequences when making temporal decisions [4], many animal species wait only a few seconds for delayed benefits [5-10]. Current research thus suggests a phylogenetic gap between patient humans and impulsive, present-oriented animals [9, 11], a distinction with implications for our understanding of economic decision making [12] and the origins of human cooperation [13]. On the basis of a series of experimental results, we reject this conclusion. First, bonobos (Pan paniscus) and chimpanzees (Pan troglodytes) exhibit a degree of patience not seen in other animals tested thus far. Second, humans are less willing to wait for food rewards than are chimpanzees. Third, humans are more willing to wait for monetary rewards than for food, and show the highest degree of patience only in response to decisions about money involving low opportunity costs. These findings suggest that core components of the capacity for future-oriented decisions evolved before the human lineage diverged from apes. Moreover, the different levels of patience that humans exhibit might be driven by fundamental differences in the mechanisms representing biological versus abstract rewards.     

Children,but not chimpanzees,have facial correlates of determination

Facial expressions have long been proposed to be important agents in forming and maintaining cooperative interactions in social groups. Human beings are inordinately cooperative when compared with their closest-living relatives, the great apes, and hence one might expect species differences in facial expressivity in contexts in which cooperation could be advantageous. Here, human children and chimpanzees were given an identical task designed to induce an element of frustration (it was impossible to solve). In children, but not chimpanzees, facial expressions associated with effort and determination positively correlated with persistence at the task. By contrast, bodily indicators of stress (self-directed behaviour) negatively correlated with task persistence in chimpanzees. Thus, children exhibited more behaviour as they persisted, and chimpanzees exhibited less. The facial expressions produced by children, could, therefore, function to solicit prosocial assistance from others.     

The ontogeny of human prosociality: behavioral experiments with children aged 3 to 8

Humans regularly engage in prosocial behavior that differs strikingly from that of even our closest living relatives, the chimpanzees (Pan troglodytes). In laboratory settings, chimpanzees are indifferent when given the opportunity to deliver valued rewards to conspecifics, while even very young human children have repeatedly been shown to behave prosocially. Although this broadly suggests that prosocial behavior in chimpanzees differs from that of young human children, the methods used in prior work with children have also differed from the methods used in studies of chimpanzees in potentially crucial ways. Here we test 92 pairs of 3–8-year-old children from urban American (Los Angeles, CA, USA) schools in a face-to-face task that closely parallels tasks used previously with chimpanzees. We found that children were more prosocial than chimpanzees have previously been in similar tasks, and our results suggest that this was driven more by a desire to provide benefits to others than a preference for egalitarian outcomes. We did not find consistent evidence that older children were more prosocial than younger children, implying that younger children behaved more prosocially in the current study than in previous studies in which participants were fully anonymous. These findings strongly suggest that humans are more prosocial than chimpanzees from an early age and that anonymity influences children's prosocial behavior, particularly at the youngest ages.     

Evolutionary psychology of spatial representations in the hominidae

Comparatively little is known about the inherited primate background underlying human cognition, the human cognitive "wild-type." Yet it is possible to trace the evolution of human cognitive abilities and tendencies by contrasting the skills of our nearest cousins, not just chimpanzees, but all the extant great apes, thus showing what we are likely to have inherited from the common ancestor. By looking at human infants early in cognitive development, we can also obtain insights into native cognitive biases in our species. Here, we focus on spatial memory, a central cognitive domain. We show, first, that all nonhuman great apes and 1-year-old human infants exhibit a preference for place over feature strategies for spatial memory. This suggests the common ancestor of all great apes had the same preference. We then examine 3-year-old human children and find that this preference reverses. Thus, the continuity between our species and the other great apes is masked early in human ontogeny. These findings, based on both phylogenetic and ontogenetic contrasts, open up the prospect of a systematic evolutionary psychology resting upon the cladistics of cognitive preferences.     

Scavenging by chimpanzees at Ngogo and the relevance of chimpanzee scavenging to early hominin behavioral ecology

Chimpanzees regularly hunt a variety of prey species. However, they rarely scavenge, which distinguishes chimpanzee carnivory from that of some modern hunter-gatherers and, presumably, at least some Plio-Pleistocene hominins. I use observations made over an 11-year period to document all known opportunities for scavenging encountered by chimpanzees at Ngogo, Kibale National Park, Uganda, and describe all cases of scavenging. I also review data on scavenging from other chimpanzee research sites. Chimpanzees at Ngogo encountered scavenging opportunities only about once per 100 days and ate meat from scavenged carcasses only four times. Scavenging opportunities are also rare at other sites, even where leopards are present (Mahale, Ta?, Gombe), and scavenging of leopard kills is known only from Mahale. Feeding on prey that chimpanzees had hunted but then abandoned is the most common form of scavenging reported across study sites. For example, several individuals at Ngogo ate meat from a partially consumed red colobus carcass abandoned after a hunt the previous day. Such behavior probably was not common among Oldowan hominins. Ngogo data and those from other sites also show that chimpanzees sometimes eat meat from carcasses of prey that they did not see killed and that were not killed by chimpanzees, and that scavenging allows access to carcasses larger than those of any prey items. However, chimpanzees ignore relatively many opportunities to obtain meat from such carcasses. Scavenging may be rare because fresh carcasses are rare, because the risk of bacterial infections and zoonoses is high, and because chimpanzees may not recognize certain species as potential prey or certain size classes of prey species as food sources. Its minimal nutritional importance, along with the absence of technology to facilitate confrontational scavenging and rapid carcass processing, apparently distinguishes chimpanzee foraging strategies from those of at least some Oldowan hominins.     

Complex tool sets for honey extraction among chimpanzees in Loango National Park, Gabon

Homo faber was once proposed as a label for humans specifically to highlight their unique propensity for tool use. However, new observations on complex tool use by the chimpanzees of Loango National Park, Gabon, expand our knowledge about tool-using abilities in Pan troglodytes. Chimpanzees in Loango, when using tools to extract honey from three types of bee nests, were observed to regularly use three- to five-element tool sets. In other words, different types of tools were used sequentially to access a single food source. Such tool sets included multi-function tools that present typical wear for two distinct uses. In addition, chimpanzees exploited underground bee nests and used ground-perforating tools to locate nest chambers that were not visible from the ground surface. These new observations concur with others from Central African chimpanzees to highlight the importance of honey extraction in arguments favoring the emergence of complex tool use in hominoids, including different tool types, expanded tool sets, multifunction tools, and the exploitation of underground resources. This last technique requires sophisticated cognitive abilities concerning unseen objects. A sequential analysis reveals a higher level of complexity in honey extraction than previously proposed for nut cracking or hunting tools, and compares with some technologies attributed to early hominins from the Early and Middle Stone Age. A better understanding of similarities in human and chimpanzee tool use will allow for a greater understanding of tool-using skills that are uniquely human.     

Evolutionary precursors of social norms in chimpanzees: a new approach

Moral behaviour, based on social norms, is commonly regarded as a hallmark of humans. Hitherto, humans are perceived to be the only species possessing social norms and to engage in moral behaviour. There is anecdotal evidence suggesting their presence in chimpanzees, but systematic studies are lacking. Here, we examine the evolution of human social norms and their underlying psychological mechanisms. For this, we distinguish between conventions, cultural social norms and universal social norms. We aim at exploring whether chimpanzees possess evolutionary precursors of universal social norms seen in humans. Chimpanzees exhibit important preconditions for their presence and enforcement: tolerant societies, well-developed social-cognitive skills and empathetic competence. Here, we develop a theoretical framework for recognizing different functional levels of social norms and distinguish them from mere statistical behavioural regularities. Quasi social norms are found where animals behave functionally moral without having moral emotions. In proto social norms, moral emotions might be present but cannot be collectivized due to the absence of a uniquely human psychological trait, i.e. shared intentionality. Human social norms, whether they are universal or cultural, involve moral emotions and are collectivized. We will discuss behaviours in chimpanzees that represent potential evolutionary precursors of human universal social norms, with special focus on social interactions involving infants. We argue that chimpanzee infants occupy a special status within their communities and propose that tolerance towards them might represent a proto social norm. Finally, we discuss possible ways to test this theoretical framework.     

Diet of chimpanzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Park, Uganda, 1. Diet composition and diversity

Chimpanzees (Pan troglodytes) are ecologically flexible omnivores with broad diets comprising many plant and animal foods, although they mostly eat fruit (including figs). Like other ecologically flexible nonhuman primates (e.g., baboons, Papio spp.) with broad diets, their diets vary across habitats. Much data on diets come from short studies that may not capture the range of variation, however, and data are scant on variation within habitats and populations. We present data on diet composition and diversity for chimpanzees at Ngogo, in Kibale National Park, Uganda, collected over a 15-year period, with a focus on the plant components of the diet. We compare Ngogo data to those on chimpanzees at the nearby Kibale site of Kanyawara, on other chimpanzee populations, and on some other frugivorous-omnivorous primates. Results support the argument that chimpanzees are ripe fruit specialists: Ngogo chimpanzees ate a broad, mostly fruit-based diet, feeding time devoted to fruit varied positively with fruit availability, and diet diversity varied inversely with fruit availability. Comparison of Ngogo and Kanyawara shows much similarity, but also pronounced within-population dietary variation. Chimpanzees fed much more on leaves, and much less on pith and stems, at Ngogo. Figs accounted for somewhat less feeding time at Ngogo, but those of Ficus mucuso were quantitatively the most important food. This species is essentially absent at Kanayawara; its abundance and high productivity at Ngogo, along with much higher abundance of several other important food species, help explain why chimpanzee community size and population density are over three times higher at Ngogo. High inter-annual variation at Ngogo highlights the value of long-term data for documenting the extent of ecological variation among chimpanzee populations and understanding how such variation might affect population biology and social dynamics.     

What's in it for me? Self-regard precludes altruism and spite in chimpanzees

Sensitivity to fairness may influence whether individuals choose to engage in acts that are mutually beneficial, selfish, altruistic, or spiteful. In a series of three experiments, chimpanzees (Pan troglodytes) could pull a rope to access out-of-reach food while concomitantly pulling another piece of food further away. In the first study, they could make a choice that solely benefited themselves (selfishness), or both themselves and another chimpanzee (mutualism). In the next two experiments, they could choose between providing food solely for another chimpanzee (altruism), or for neither while preventing the other chimpanzee from receiving a benefit (spite). The main result across all studies was that chimpanzees made their choices based solely on personal gain, with no regard for the outcomes of a conspecific. These results raise questions about the origins of human cooperative behaviour.