Experimental investigation of orangutans' lithic percussive and sharp stone tool behaviours.

Early stone tools, and in particular sharp stone tools, arguably represent one of the most important technological milestones in human evolution. The production and use of sharp stone tools significantly widened the ecological niche of our ancestors, allowing them to exploit novel food resources. However, despite their importance, it is still unclear how these early lithic technologies emerged and which behaviours served as stepping-stones for the development of systematic lithic production in our lineage. One approach to answer this question is to collect comparative data on the stone tool making and using abilities of our closest living relatives, the great apes, to reconstruct the potential stone-related behaviours of early hominins. To this end, we tested both the individual and the social learning abilities of five orangutans to make and use stone tools. Although the orangutans did not make sharp stone tools initially, three individuals spontaneously engaged in lithic percussion, and sharp stone pieces were produced under later experimental conditions. Furthermore, when provided with a human-made sharp stone, one orangutan spontaneously used it as a cutting tool. Contrary to previous experiments, social demonstrations did not considerably improve the stone tool making and using abilities of orangutans. Our study is the first to systematically investigate the stone tool making and using abilities of untrained, unenculturated orangutans showing that two proposed pre-requisites for the emergence of early lithic technologies-lithic percussion and the recognition of sharp-edged stones as cutting tools-are present in this species. We discuss the implications that ours and previous great ape stone tool experiments have for understanding the initial stages of lithic technologies in our lineage.

Immature wild orangutans acquire relevant ecological knowledge through sex-specific attentional biases during social learning.

As a part of growing up, immature orangutans must acquire vast repertoires of skills and knowledge, a process that takes several years of observational social learning and subsequent practice. Adult female and male orangutans show behavioral differences including sex-specific foraging patterns and male-biased dispersal. We investigated how these differing life trajectories affect social interest and emerging ecological knowledge in immatures. We analyzed 15 years of detailed observational data on social learning, associations, and diet repertoires of 50 immatures (16 females and 34 males), from 2 orangutan populations. Specific to the feeding context, we found sex differences in the development of social interest: Throughout the dependency period, immature females direct most of their social attention at their mothers, whereas immature males show an increasing attentional preference for individuals other than their mothers. When attending to non-mother individuals, males show a significant bias toward immigrant individuals and a trend for a bias toward adult males. In contrast, females preferentially attend to neighboring residents. Accordingly, by the end of the dependency period, immature females show a larger dietary overlap with their mothers than do immature males. These results suggest that immature orangutans show attentional biases through which they learn from individuals with the most relevant ecological knowledge. Diversifying their skills and knowledge likely helps males when they move to a new area. In sum, our findings underline the importance of fine-grained social inputs for the acquisition of ecological knowledge and skills in orangutans and likely in other apes as well.

Comparative performance of orangutans (Pongo spp.), gorillas (Gorilla gorilla gorilla), and drills (Mandrillus leucophaeus), in an ephemeral foraging task.

A goal of the comparative approach is to test a variety of species on the same task. Here, we examined whether the factors that helped capuchin monkeys improve their performance in a dichotomous choice task would generalize to three other primate species: orangutans, gorillas, and drill monkeys. In this task, subjects have access to two options, each resulting in an identical food, but one (the ephemeral option) is only available if it is chosen first, whereas the other one (the permanent option) is always available. Therefore, the food-maximizing solution is to choose the ephemeral option first, followed by the permanent option for an additional reward. On the original version (plate task), the options were discriminated by the color and pattern of the plates holding the food, while on two subsequent versions we used altered cues that we predicted would improve performance: (1) the color of the foods themselves (color task), which we hypothesized was relevant to primates, who choose foods rather than substrates on which foods are found when foraging, and (2) patterned cups covering the foods (cup task), which we hypothesized would help primates avoid the prepotent response associated with visible food. Like capuchins, all three species initially failed to solve the plate task. However, while orangutans improved their performance from the plate to the color task, they did not for the cup task, and only a few gorillas and no drills succeeded in either task. Unfortunately, our ability to interpret these data was obscured by differences in the subjects' level of experience with cognitive testing and practical constraints that precluded the use of completely identical procedures across species. Nonetheless, we consider what these results can tell us, and discuss the value of conducting studies across multiple sites despite unavoidable differences.

Spontaneous (minimal) ritual in non-human great apes?

The potential for rituals in non-human great apes (apes) is an understudied topic. We derive a minimal definition of ritual and then examine the currently available evidence for it in untrained and non-enculturated apes. First, we examine whether such apes show evidence for the two main components of our minimal definition of ritual: symbolism and copying. Second, we examine if there are actual cases already identifiable today that may fit all aspects of our minimal definition of ritual-or whether there are at least cases that fit some aspects (proto-ritual). We find that apes are not likely to spontaneously practise minimal ritual, but we claim that the highest expected likelihood of occurrence is in the results-copying domain. Yet, we did not find actual cases of minimal ritual in apes-including those involving environmental results. We did, however, find some cases that may match at least part of our minimal ritual definition-which we termed proto-ritual. At least two out of three potential cases of such proto-rituals that we identified (rain dance, object-in-ear and surplus nest-making procedures) do revolve around results. Overall, apes do not show much, or very clear, evidence for even minimal ritual, but may sometimes show proto-ritual. However, dedicated ape ritual studies are currently lacking, and future work may identify ape ritual (or clearer cases of proto-ritual). We discuss the implications of our preliminary finding for inferences of ritual in the last common ancestor of humans and apes. This article is part of the theme issue 'Ritual renaissance: new insights into the most human of behaviours'.

Specialization in the vicarious learning of novel arbitrary sequences in humans but not orangutans.

Sequence learning underlies many uniquely human behaviours, from complex tool use to language and ritual. To understand whether this fundamental cognitive feature is uniquely derived in humans requires a comparative approach. We propose that the vicarious (but not individual) learning of novel arbitrary sequences represents a human cognitive specialization. To test this hypothesis, we compared the abilities of human children aged 3-5 years and orangutans to learn different types of arbitrary sequences (item-based and spatial-based). Sequences could be learned individually (by trial and error) or vicariously from a human (social) demonstrator or a computer (ghost control). We found that both children and orangutans recalled both types of sequence following trial-and-error learning; older children also learned both types of sequence following social and ghost demonstrations. Orangutans' success individually learning arbitrary sequences shows that their failure to do so in some vicarious learning conditions is not owing to general representational problems. These results provide new insights into some of the most persistent discontinuities observed between humans and other great apes in terms of complex tool use, language and ritual, all of which involve the cultural learning of novel arbitrary sequences. This article is part of the theme issue 'Ritual renaissance: new insights into the most human of behaviours'.

Social inhibition and behavioural flexibility when the context changes: a comparison across six primate species.

The ability to inhibit previously employed strategies and flexibly adjust behavioural responses to external conditions may be critical for individual survival. However, it is unclear which factors predict their distribution across species. Here, we investigated social inhibition and behavioural flexibility in six primate species (chimpanzees, bonobos, orangutans, gorillas, capuchin monkeys and spider monkeys) differing in terms of phylogenetic relatedness, foraging ecology and social organization. Depending on the social context, individuals could maximize their food intake by inhibiting the selection of a larger food reward in one condition (i.e. inhibition), but not in others, which required them to flexibly switching strategies across conditions (i.e. behavioural flexibility). Overall, our study revealed inter-specific differences in social inhibition and behavioural flexibility, which partially reflected differences in fission-fusion dynamics. In particular, orangutans and chimpanzees showed the highest level of inhibitory skills, while gorillas and capuchin monkeys showed the lowest one. In terms of behavioural flexibility, orangutans and spider monkeys were the best performers, while bonobos and capuchin monkeys were the worst ones. These results contribute to our understanding that inhibition and behavioural flexibility may be linked in more complex ways than usually thought, although both abilities play a crucial role in efficient problem solving.

An orangutan in Paris: pondering Proximity at the Muséum d'histoire naturelle in 1836.

When the Muséum d'histoire naturelle in Paris learned in 1836 that it had the chance to buy a live, young orangutan, it was excited by the prospect. Specimens were the focus of the Museum's activities, and this particular specimen seemed especially promising, not only because the Museum had very few orangutan specimens in its collection, but also because of what was perceived to be the orangutan's unique place in the natural order of things, namely, at the very boundary between the animal kingdom and humans. Frédéric Cuvier, the superintendent of the Museum's menagerie, urged that studying the orangutan's mental faculties would help resolve fundamental questions regarding the similarities and differences between animals and humans. Archival and printed sources allow one to reconstruct the orangutan's capture, acquisition, and subsequent career at the menagerie in greater detail than has generally been possible for animals of nineteenth-century zoos. Scientists, artists, the public, the press, and even musicians (Franz Liszt included) sought to engage with the orangutan, seeing in it not just another ape or monkey but a special creature unto itself at the animal/human boundary. Key to their fascination with the orangutan was the question of proximity-just how close was the orangutan to humans? The orangutan's story illuminates not only how the animal-human boundary was conceived at the time but also the problematic status of the zoo as a site for scientific research and the roles of scientific and non-scientific actors alike in constructing how the orangutan was understood.

Social manipulation in nonhuman primates: Cognitive and motivational determinants.

Social interactions are the result of individuals' cooperative and competitive tendencies expressed over an extended period of time. Although social manipulation, i.e., using another individual to achieve one's own goals, is a crucial aspect of social interactions, there has been no comprehensive attempt to differentiate its various types and to map its cognitive and motivational determinants. For this purpose, we survey in this article the experimental literature on social interactions in nonhuman primates. We take social manipulation, illustrated by a case study with orangutans (Pongo abelii), as our starting point and move in two directions. First, we will focus on a flexibility/sociality axis that includes technical problem solving, social tool-use and communication. Second, we will focus on a motivational/prosociality axis that includes exploitation, cooperation, and helping. Combined, the two axes offer a way to capture a broad range of social interactions performed by human and nonhuman primates.

Differences in between-reinforcer value modulate the selective-value effect in great apes (Pan troglodytes, P. Paniscus, Gorilla gorilla, Pongo abelii).

We investigated how apes allocated their choices between 2 food options that varied in terms of their quantity and quality. Experiment 1 tested whether subjects preferred an AB option over an A option, where the A item is preferred to the B item (e.g., apple + carrot vs. apple). Additionally, we tested whether the length of the intertrial interval (ITI) affected subjects' choices. Five orangutans, 4 gorillas, 7 bonobos, and 10 chimpanzees received 3 types of trials: preference (A vs. B), quantity (AA vs. A), and mixed (AB vs. A where A is the preferred food). We used 3 food items that substantially differed in terms of preference (carrots, apples, and pellets). Subjects showed no overall preference for the mixed option (AB) compared with the single option (A), even though they showed clear preferences during both the preference and quantity trials. The intertrial length had no effect on choice behavior. Experiment 2 further explored apes' choices by using 3 highly preferred food items (bananas, grapes, and pellets) in 6 orangutans, 4 gorillas, 8 bonobos, and 18 chimpanzees. Unlike the results of Experiment 1, apes generally chose the mixed option. Our results indicated that apes did not show a general "selective-value" effect but chose depending on the relative value of the food items involved. Subjects were more likely to select the mixed over the single option when (a) the mixed option was composed of items that were closer in value and (b) they were compared against the less valuable item forming the mixed option.

Apes produce tools for future use.

There is now growing evidence that some animal species are able to plan for the future. For example great apes save and exchange tools for future use. Here we raise the question whether chimpanzees, orangutans, and bonobos would produce tools for future use. Subjects only had access to a baited apparatus for a limited duration and therefore should use the time preceding this access to create the appropriate tools in order to get the rewards. The apes were tested in three conditions depending on the need for pre-prepared tools. Either eight tools, one tool or no tools were needed to retrieve the reward. The apes prepared tools in advance for future use and they produced them mainly in conditions when they were really needed. The fact that apes were able to solve this new task indicates that their planning skills are flexible. However, for the condition in which eight tools were needed, apes produced less than two tools per trial in advance. However, they used their chance to produce additional tools in the tool use phase-thus often obtaining most of the reward from the apparatus. Increased pressure to prepare more tools in advance did not have an effect on their performance.

The ecology of primate material culture.

Tool use in extant primates may inform our understanding of the conditions that favoured the expansion of hominin technology and material culture. The 'method of exclusion' has, arguably, confirmed the presence of culture in wild animal populations by excluding ecological and genetic explanations for geographical variation in behaviour. However, this method neglects ecological influences on culture, which, ironically, may be critical for understanding technology and thus material culture. We review all the current evidence for the role of ecology in shaping material culture in three habitual tool-using non-human primates: chimpanzees, orangutans and capuchin monkeys. We show that environmental opportunity, rather than necessity, is the main driver. We argue that a better understanding of primate technology requires explicit investigation of the role of ecological conditions. We propose a model in which three sets of factors, namely environment, sociality and cognition, influence invention, transmission and retention of material culture.

Children conform to the behavior of peers; other great apes stick with what they know.

All primates learn things from conspecifics socially, but it is not clear whether they conform to the behavior of these conspecifics--if conformity is defined as overriding individually acquired behavioral tendencies in order to copy peers' behavior. In the current study, chimpanzees, orangutans, and 2-year-old human children individually acquired a problem-solving strategy. They then watched several conspecific peers demonstrate an alternative strategy. The children switched to this new, socially demonstrated strategy in roughly half of all instances, whereas the other two great-ape species almost never adjusted their behavior to the majority's. In a follow-up study, children switched much more when the peer demonstrators were still present than when they were absent, which suggests that their conformity arose at least in part from social motivations. These results demonstrate an important difference between the social learning of humans and great apes, a difference that might help to account for differences in human and nonhuman cultures.

Orangutan (Pongo spp.) whistling and implications for the emergence of an open-ended call repertoire: a replication and extension.

One of the most apparent discontinuities between non-human primate (primate) call communication and human speech concerns repertoire size. The former is essentially fixed to a limited number of innate calls, while the latter essentially consists of numerous learned components. Consequently, primates are thought to lack laryngeal control required to produce learned voiced calls. However, whether they may produce learned voiceless calls awaits investigation. Here, a case of voiceless call learning in primates is investigated--orangutan (Pongo spp.) whistling. In this study, all known whistling orangutans are inventoried, whistling-matching tests (previously conducted with one individual) are replicated with another individual using original test paradigms, and articulatory and acoustic whistle characteristics are compared between three orangutans. Results show that whistling has been reported for ten captive orangutans. The test orangutan correctly matched human whistles with significantly high levels of performance. Whistle variation between individuals indicated voluntary control over the upper lip, lower lip, and respiratory musculature, allowing individuals to produce learned voiceless calls. Results are consistent with inter- and intra-specific social transmission in whistling orangutans. Voiceless call learning in orangutans implies that some important components of human speech learning and control were in place before the homininae-ponginae evolutionary split.

Memory for distant past events in chimpanzees and orangutans.

Determining the memory systems that support nonhuman animals' capacity to remember distant past events is currently the focus an intense research effort and a lively debate [1-3]. Comparative psychology has largely adopted Tulving's framework by focusing on whether animals remember what-where-when something happened (i.e., episodic-like memory) [4-6]. However, apes have also been reported to recall other episodic components [7] after single-trial exposures [8, 9]. Using a new experimental paradigm we show that chimpanzees and orangutans recalled a tool-finding event that happened four times 3 years earlier (experiment 1) and a tool-finding unique event that happened once 2 weeks earlier (experiment 2). Subjects were able to distinguish these events from other tool-finding events, which indicates binding of relevant temporal-spatial components. Like in human involuntary autobiographical memory, a cued, associative retrieval process triggered apes' memories: when presented with a particular setup, subjects instantaneously remembered not only where to search for the tools (experiment 1), but also the location of the tool seen only once (experiment 2). The complex nature of the events retrieved, the unexpected and fast retrieval, the long retention intervals involved, and the detection of binding strongly suggest that chimpanzees and orangutans' memories for past events mirror some of the features of human autobiographical memory.

Personality structure in brown capuchin monkeys (Sapajus apella): comparisons with chimpanzees (Pan troglodytes), orangutans (Pongo spp.), and rhesus macaques (Macaca mulatta).

Species comparisons of personality structure (i.e., how many personality dimensions and the characteristics of those dimensions) can facilitate questions about the adaptive function of personality in nonhuman primates. Here we investigate personality structure in the brown capuchin monkey (Sapajus apella), a New World primate species, and compare this structure to those of chimpanzees (Pan troglodytes), orangutans (Pongo spp.), and rhesus macaques (Macaca mulatta). Brown capuchins evolved behavioral and cognitive traits that are qualitatively similar to those of great apes, and individual differences in behavior and cognition often reflect differences in personality. Thus, we hypothesized that brown capuchin personality structure would overlap more with great apes than with rhesus macaques. We obtained personality ratings from seven sites, including 127 brown capuchin monkeys. Principal-components analysis identified five personality dimensions (Assertiveness, Openness, Neuroticism, Sociability, and Attentiveness), which were reliable across raters and, in a subset of subjects, significantly correlated with relevant behaviors up to a year later. Comparisons between species revealed that brown capuchins and great apes overlapped in personality structure, particularly chimpanzees in the case of Neuroticism. However, in some respects (i.e., capuchin Sociability and Openness) the similarities between capuchins and great apes were not significantly greater than those between capuchins and rhesus macaques. We discuss the relevance of our results to brown capuchin behavior and the evolution of personality structure in primates.

Orangutans (Pongo pygmaeus and Pongo abelii) understand connectivity in the skewered grape tool task.

Great apes appear to have limited knowledge of tool functionality when they are presented with tasks that involve a physical connection between a tool and a reward. For instance, they fail to understand that pulling a rope with a reward tied to its end is more beneficial than pulling a rope that only touches a reward. Apes show more success when both ropes have rewards tied to their ends but one rope is nonfunctional because it is clearly separated into aligned sections. It is unclear, however, whether this success is based on perceptual features unrelated to connectivity, such as perceiving the tool's separate sections as independent tools rather than one discontinuous tool. Surprisingly, there appears to be no study that has tested any type of connectivity problem using natural tools made from branches with which wild and captive apes often have extensive experience. It is possible that such ecologically valid tools may better help subjects understand connectivity that involves physical attachment. In this study, we tested orangutans with natural tools and a range of connectivity problems that involved the physical attachment of a reward on continuous and broken tools. We found that the orangutans understood tool connectivity involving physical attachment that apes from other studies failed when tested with similar tasks using artificial as opposed to natural tools. We found no evidence that the orangutans' success in broken tool conditions was based on perceptual features unrelated to connectivity. Our results suggest that artificial tools may limit apes' knowledge of connectivity involving physical attachment, whereas ecologically valid tools may have the opposite effect.

Evidence for a midlife crisis in great apes consistent with the U-shape in human well-being.

Recently, economists and behavioral scientists have studied the pattern of human well-being over the lifespan. In dozens of countries, and for a large range of well-being measures, including happiness and mental health, well-being is high in youth, falls to a nadir in midlife, and rises again in old age. The reasons for this U-shape are still unclear. Present theories emphasize sociological and economic forces. In this study we show that a similar U-shape exists in 508 great apes (two samples of chimpanzees and one sample of orangutans) whose well-being was assessed by raters familiar with the individual apes. This U-shaped pattern or "midlife crisis" emerges with or without use of parametric methods. Our results imply that human well-being's curved shape is not uniquely human and that, although it may be partly explained by aspects of human life and society, its origins may lie partly in the biology we share with great apes. These findings have implications across scientific and social-scientific disciplines, and may help to identify ways of enhancing human and ape well-being.

Visual discrimination in an orangutan (Pongo pygmaeus): measuring visual preference.

Although previous studies have confirmed that trained orangutans visually discriminate between mammals and artificial objects, whether orangutans without operant conditioning can discriminate remains unknown. The visual discrimination ability in an orangutan (Pongo pygmaeus) with no experience in operant learning was examined using measures of visual preference. Sixteen color photographs of inanimate objects and of mammals with four legs were randomly presented to an orangutan. The results showed that the mean looking time at photographs of mammals with four legs was longer than that for inanimate objects, suggesting that the orangutan discriminated mammals with four legs from inanimate objects. The results implied that orangutans who have not experienced operant conditioning may possess the ability to discriminate visually.

Old world monkeys compare to apes in the primate cognition test battery.

Understanding the evolution of intelligence rests on comparative analyses of brain sizes as well as the assessment of cognitive skills of different species in relation to potential selective pressures such as environmental conditions and social organization. Because of the strong interest in human cognition, much previous work has focused on the comparison of the cognitive skills of human toddlers to those of our closest living relatives, i.e. apes. Such analyses revealed that apes and children have relatively similar competencies in the physical domain, while human children excel in the socio-cognitive domain; in particular in terms of attention sharing, cooperation, and mental state attribution. To develop a full understanding of the evolutionary dynamics of primate intelligence, however, comparative data for monkeys are needed. We tested 18 Old World monkeys (long-tailed macaques and olive baboons) in the so-called Primate Cognition Test Battery (PCTB) (Herrmann et al. 2007, Science). Surprisingly, our tests revealed largely comparable results between Old World monkeys and the Great apes. Single comparisons showed that chimpanzees performed only better than the macaques in experiments on spatial understanding and tool use, but in none of the socio-cognitive tasks. These results question the clear-cut relationship between cognitive performance and brain size and--prima facie--support the view of an accelerated evolution of social intelligence in humans. One limitation, however, is that the initial experiments were devised to tap into human specific skills in the first place, thus potentially underestimating both true nonhuman primate competencies as well as species differences.

Nest-building orangutans demonstrate engineering know-how to produce safe, comfortable beds.

Nest-building orangutans must daily build safe and comfortable nest structures in the forest canopy and do this quickly and effectively using the branches that surround them. This study aimed to investigate the mechanical design and architecture of orangutan nests and determine the degree of technical sophistication used in their construction. We measured the whole nest compliance and the thickness of the branches used and recorded the ways in which the branches were fractured. Branch samples were also collected from the nests and subjected to three-point bending tests to determine their mechanical properties. We demonstrated that the center of the nest is more compliant than the edges; this may add extra comfort and safety to the structure. During construction orangutans use the fact that branches only break half-way across in "greenstick" fracture to weave the main nest structure. They choose thicker branches with greater rigidity and strength to build the main structure in this way. They then detach thinner branches by following greenstick fracture with a twisting action to make the lining. These results suggest that orangutans exhibit a degree of technical knowledge and choice in the construction of nests.