Spread of arbitrary conventions among chimpanzees: a controlled experiment

Wild chimpanzees (Pan troglodytes) have a rich cultural repertoire--traditions common in some communities are not present in others. The majority of reports describe functional, material traditions, such as tool use. Arbitrary conventions have received far less attention. In the same way that observations of material culture in wild apes led to experiments to confirm social transmission and identify underlying learning mechanisms, experiments investigating how arbitrary habits or conventions arise and spread within a group are also required. The few relevant experimental studies reported thus far have relied on cross-species (i.e. human-ape) interaction offering limited ecological validity, and no study has successfully generated a tradition not involving tool use in an established group. We seeded one of two rewarded alternative endpoints to a complex sequence of behaviour in each of two chimpanzee groups. Each sequence spread in the group in which it was seeded, with many individuals unambiguously adopting the sequence demonstrated by a group member. In one group, the alternative sequence was discovered by a low ranking female, but was not learned by others. Since the action-sequences lacked meaning before the experiment and had no logical connection with reward, chimpanzees must have extracted both the form and benefits of these sequences through observation of others.     

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.     

Cultural panthropology

Culture, in the most basic sense of “tradition,” has been shown to exist in many species. There is more to the phenomenon of culture in humans, however, than the mere existence of traditions. Thus, rather than expecting that culture can be assigned to living or ancestral species in an all‐or‐none fashion, reconstruction of the evolution of this uniquely complex phenomenon is likely to depend on successfully teasing apart its components, which may have evolved in a somewhat mosaic fashion. In this paper, we dissect ten different aspects of human culture and offer evidence that most of them are manifested in chimpanzees, even if in limited ways, permitting inferences about the cultural profile of our common ancestor. The aspects of culture examined include large‐scale patterns of behavioral variation across populations, the mechanisms available for social transmission, and cultural contents. The contrasts thus drawn for humans and chimpanzees offer a framework for cultural comparisons between other taxa from the past and present.     

Innovation in chimpanzees

The study of innovation in non‐human animals (henceforth: animals) has recently gained momentum across fields including primatology, animal behaviour and cultural evolution. Examining the rate of innovations, and the cognitive mechanisms driving these innovations across species, can provide insights into the evolution of human culture. Especially relevant to the study of human culture is one of our closest living relatives, the chimpanzee (Pan troglodytes). Both wild and captive chimpanzees demonstrate an impressive ability to innovate solutions to novel problems, but also a striking level of conservatism in some contexts, creating a unique and at times puzzling, picture of animal innovation. Whilst the animal innovation field is rife with potential for expanding our knowledge of human and non‐human cognition and problem‐solving, it is undermined by a lack of consistency across studies. The field is yet to settle on a definition of the term ‘innovation’, leading to studies being incomparable across and even within the same species. Here, we fill two gaps in the literature. First, we discuss some of the most prevalent definitions of ‘innovation’ from different fields, highlighting similarities and differences between them. Secondly, we provide an up‐to‐date review of accounts of innovations in both wild and captive chimpanzees. We hope this review will provide a resource for researchers interested in the study of innovation in chimpanzees and other animals, as well as emphasising the need for consistency in the way in which innovations are reported.     

Chimpanzees fail to plan in an exchange task but succeed in a tool-using procedure

Planning has long been considered a uniquely human capacity. Lately, however, it has been shown that apes and a corvid species act now to derive a material future benefit. Since primates are highly social animals and their sociality is considered a strong selective force that resulted in complex cognitive capacities, planning is also expected in social situations. Unfortunately, prompting from social partners cannot be excluded in a social setting. Therefore, we controlled for this factor by testing the capacity to plan in chimpanzees using an exchange paradigm, that involves both a material and a social component, and a tool-use paradigm, similar to the one used on two other ape species. All chimpanzees failed to plan in the exchange task, but three individuals showed planning behavior in the tool-use task. Our methods controlled for the fact that chimpanzees were not prompted by the visibility of the reward at the moment of planning and also could not repeat a previously acquired routine. The best interpretation for our results is that chimpanzees can plan. However, planning was limited to the situation where the action to attain the future benefit only depended on a chimpanzee's own behavior.     

Chimpanzees coordinate in a negotiation game

A crucially important aspect of human cooperation is the ability to negotiate to cooperative outcomes when interests over resources conflict. Although chimpanzees and other social species may negotiate conflicting interests regarding travel direction or activity timing, very little is known about their ability to negotiate conflicting preferences over food. In the current study, we presented pairs of chimpanzees with a choice between two cooperative tasks—one with equal payoffs (e.g., 5-5) and one with unequal payoffs (higher and lower than in the equal option, e.g., 10-1). This created a conflict of interests between partners with failure to work together on the same cooperative task resulting in no payoff for either partner. The chimpanzee pairs cooperated successfully in as many as 78–94% of the trials across experiments. Even though dominant chimpanzees preferred the unequal option (as they would obtain the largest payoff), subordinate chimpanzees were able to get their way (the equal option) in 22–56% of trials across conditions. Various analyses showed that subjects were both strategic and also cognizant of the strategies used by their partners. These results demonstrate that one of our two closest primate relatives, the chimpanzee, can settle conflicts of interest over resources in mutually satisfying ways—even without the social norms of equity, planned strategies of reciprocity, and the complex communication characteristic of human negotiation.     

Hunter-Gatherer Inter-Band Interaction Rates: Implications for Cumulative Culture

Our species exhibits spectacular success due to cumulative culture. While cognitive evolution of social learning mechanisms may be partially responsible for adaptive human culture, features of early human social structure may also play a role by increasing the number potential models from which to learn innovations. We present interview data on interactions between same-sex adult dyads of Ache and Hadza hunter-gatherers living in multiple distinct residential bands (20 Ache bands; 42 Hadza bands; 1201 dyads) throughout a tribal home range. Results show high probabilities (5%–29% per year) of cultural and cooperative interactions between randomly chosen adults. Multiple regression suggests that ritual relationships increase interaction rates more than kinship, and that affinal kin interact more often than dyads with no relationship. These may be important features of human sociality. Finally, yearly interaction rates along with survival data allow us to estimate expected lifetime partners for a variety of social activities, and compare those to chimpanzees. Hadza and Ache men are estimated to observe over 300 men making tools in a lifetime, whereas male chimpanzees interact with only about 20 other males in a lifetime. High intergroup interaction rates in ancestral humans may have promoted the evolution of cumulative culture.     

Behavioural diversity among the wild chimpanzee populations of Bossou and neighbouring areas, Guinea and Côte d'Ivoire, West Africa. A preliminary report

We present a preliminary report on the differences and similarities in material culture among four neighbouring chimpanzee (Pan troglodytes verus) communities. One of these communities includes Bossou, a long-term field site of wild chimpanzees, in Guinea, West Africa. We also conducted surveys of three new sites. Two of those surveyed areas, Seringbara in Guinea and Yealé in C?te d'Ivoire, are located less than 12 km away from Bossou in the Nimba Mountains region, which forms a natural boundary between Guinea and C?te d'Ivoire. The third, Diécké, is situated further south-west, closer to the border with Liberia. During the surveys, we gathered behavioural information about these neighbouring populations of chimpanzees. The differences, as well as similarities, in material culture were tabulated based on our findings. The three behavioural variants found so far involve differences in nut cracking behaviour with regard to the species of nut cracked. Some variation in materials used for nut cracking has also been recorded. However, we still need to establish whether these local variations can be explained by the demands of the physical and biotic environments in which the populations of chimpanzees live. If these alternative hypotheses can be excluded with continuing research at the study sites, these differences are likely to be cultural behaviours that are influenced by the social context and mode, i.e. horizontal, vertical or oblique, of transmission, by the social structure and organisation of each community and/or perhaps by some form of social norms prevalent within these communities.     

Is culture a golden barrier between human and chimpanzee?

Culture pervades much of human existence. Its significance to human social interaction and cognitive development has convinced some researchers that the phenomenon and its underlying mechanisms represent a defining criterion for humankind. However, care should be taken not to make hasty conclusions in light of the growing number of observations on the cultural abilities of different species, ranging from chimpanzees and orangutans to whales and dolphins. The present review concentrates on wild chimpanzees and shows that they all possess an extensive cultural repertoire. In the light of what we know from humans, I evaluate the importance of social learning leading to acquisition of cultural traits, as well as of collective meaning of communicative traits. Taking into account cross‐cultural variations in humans, I argue that the cultural abilities we observe in wild chimpanzees present a broad level of similarity between the two species.     

Innovation in wild chimpanzees (Pan troglodytes)

Innovations of behavior have major implications for the concept of culture in animals. Innovation has rarefy been documented in wild animal populations. The chimpanzees of the Taï National Park, Côte d’Ivoire, spontaneously included new patterns of behavior in their repertoire during our study. Some innovations were incorporated into a minority of the group members’ repertoire, whereas others, such as building day nests in trees and on the ground more frequently, became general behaviors. Similarly, new contexts for use of leaf- clipping behavior and novel ways of feeding on some leaves appeared and rapidly became part of the behavior of most group members. The environmental parameters were too stable to explain these new forms of behavior, as most of them took place within 1 month during the same dry season. In a similar way, leaf- grooming acquired a new function in Gombe chimpanzees. A process similar to social conventions could explain the emergence of these new functions for an existing behavior in a way rather similar to human “fashion. ” Three major characteristics of human culture — the absence of individual variations in the performance of the behavior, the general use of the behavior by group members, and the ability to modify the function of a behavior — characterize leaf-clipping in Taï chimpanzees and leaf- grooming in Gombe chimpanzees.     

Human children rely more on social information than chimpanzees do

Human societies are characterized by more cultural diversity than chimpanzee communities. However, it is currently unclear what mechanism might be driving this difference. Because reliance on social information is a pivotal characteristic of culture, we investigated individual and social information reliance in children and chimpanzees. We repeatedly presented subjects with a reward-retrieval task on which they had collected conflicting individual and social information of equal accuracy in counterbalanced order. While both species relied mostly on their individual information, children but not chimpanzees searched for the reward at the socially demonstrated location more than at a random location. Moreover, only children used social information adaptively when individual knowledge on the location of the reward had not yet been obtained. Social information usage determines information transmission and in conjunction with mechanisms that create cultural variants, such as innovation, it facilitates diversity. Our results may help explain why humans are more culturally diversified than chimpanzees.     

Human cumulative culture: a comparative perspective

Many animals exhibit social learning and behavioural traditions, but human culture exhibits unparalleled complexity and diversity, and is unambiguously cumulative in character. These similarities and differences have spawned a debate over whether animal traditions and human culture are reliant on homologous or analogous psychological processes. Human cumulative culture combines high‐fidelity transmission of cultural knowledge with beneficial modifications to generate a ‘ratcheting’ in technological complexity, leading to the development of traits far more complex than one individual could invent alone. Claims have been made for cumulative culture in several species of animals, including chimpanzees, orangutans and New Caledonian crows, but these remain contentious. Whilst initial work on the topic of cumulative culture was largely theoretical, employing mathematical methods developed by population biologists, in recent years researchers from a wide range of disciplines, including psychology, biology, economics, biological anthropology, linguistics and archaeology, have turned their attention to the experimental investigation of cumulative culture. We review this literature, highlighting advances made in understanding the underlying processes of cumulative culture and emphasising areas of agreement and disagreement amongst investigators in separate fields.     

The scope of culture in chimpanzees, humans and ancestral apes

More studies have focused on aspects of chimpanzee behaviour and cognition relevant to the evolution of culture than on any other species except our own. Accordingly, analysis of the features shared by chimpanzees and humans is here used to infer the scope of cultural phenomena in our last common ancestor, at the same time clarifying the nature of the special characteristics that advanced further in the hominin line. To do this, culture is broken down into three major aspects: the large scale, population-level patterning of traditions; social learning mechanisms; and the behavioural and cognitive contents of culture. Each of these is further dissected into subcomponents. Shared features, as well as differences, are identified in as many as a dozen of these, offering a case study for the comparative analysis of culture across animal taxa and a deeper understanding of the roots of our own cultural capacities.     

Differential sensitivity to conspecific and allospecific cues in chimpanzees and humans: a comparative eye-tracking study

Previous studies have shown that a variety of animals including humans are sensitive to social cues from others and shift their attention to the same objects attended to by others. However, little is known about how animals process conspecifics'' and another species'' actions, although primates recognize conspecific faces better than those of another species. In this study, using unrestrained eye-tracking techniques, we first demonstrated that conspecific social cues modulated looking behaviours of chimpanzees more than human cues, whereas human observers were equally sensitive to both species. Additionally, first pass gaze duration at the face indicates that chimpanzees looked at the chimpanzees'' face longer than the human face, suggesting that chimpanzees might extract more referential information from a conspecific face. These results also imply that a unique ability for extracting referential information from a variety of social objects has emerged during human evolution.     

Subsistence Technology of Nigerian Chimpanzees

A trademark of Homo sapiens is the enormous variation in behavioral patterns across populations. Insight into the development of human cultures can be aided by studies on communities of Pan across Africa that display unique combinations of social behavior and elementary technology. Only cross-population comparisons can reveal whether the diversity reflects differential genetics, environmental constraints, or is a cultural variant. However, the recently recognized and most endangered subspecies, Pan troglodytes vellerosus, remains completely unstudied in this respect. We report first evidence from a new long-term study of Nigerian chimpanzees at Gashaka. Their dietary composition is highly varied and they have to cope with high concentrations of antifeedant defenses of plants against consumption. Gashaka chimpanzees use a varied tool kit for extractive foraging. For example, they harvest insects throughout the year, via digging sticks and probes, to obtain honey from stingless-bee and honeybee nests, dipping wands to prey on army ants, and fishing rods to eat arboreal ants. Tools appeared to be custom-made with a considerable degree of standardization in length, diameter, and preferential use of distal ends. Moreover, compared to the rainy season, tools were longer during the dry season when insects retreat further into their nests. Many of the expressions of subsistence technology seem to be environmentally constrained. Most notably, the absence of termite-eating could reflect a low abundance of mounds. Other traits may represent cultural variation. For example, the chimpanzees did not hammer open 2 types of hard-shelled nuts with tools, unlike what occurs elsewhere in West Africa. The prevalence of elementary technology may indicate that the material culture of Gashaka chimpanzees is most related to core cultural tendencies of Central African populations.     

Transmission of multiple traditions within and between chimpanzee groups

Field reports provide increasing evidence for local behavioral traditions among fish, birds, and mammals. These findings are significant for evolutionary biology because social learning affords faster adaptation than genetic change and has generated new (cultural) forms of evolution. Orangutan and chimpanzee field studies suggest that like humans, these apes are distinctive among animals in each exhibiting over 30 local traditions. However, direct evidence is lacking in apes and, with the exception of vocal dialects, in animals generally for the intergroup transmission that would allow innovations to spread widely and become evolutionarily significant phenomena. Here, we provide robust experimental evidence that alternative foraging techniques seeded in different groups of chimpanzees spread differentially not only within groups but serially across two further groups with substantial fidelity. Combining these results with those from recent social-diffusion studies in two larger groups offers the first experimental evidence that a nonhuman species can sustain unique local cultures, each constituted by multiple traditions. The convergence of these results with those from the wild implies a richness in chimpanzees' capacity for culture, a richness that parsimony suggests was shared with our common ancestor.     

Population-level variability in the social climates of four chimpanzee societies

Recent debates have questioned the extent to which culturally-transmitted norms drive behavioral variation in resource sharing across human populations. We shed new light on this discussion by examining the group-level variation in the social dynamics and resource sharing of chimpanzees, a species that is highly social and forms long-term community associations but differs from humans in the extent to which cultural norms are adopted and enforced. We rely on theory developed in primate socioecology to guide our investigation in four neighboring chimpanzee groups at a sanctuary in Zambia. We used a combination of experimental and observational approaches to assess the distribution of resource holding potential in each group. In the first assessment, we measured the proportion of the population that gathered in a resource-rich zone, in the second we assessed naturally occurring social spacing via social network analysis, and in the third we assessed the degree to which benefits were equally distributed within the group. We report significant, stable group-level variation across these multiple measures, indicating that group-level variation in resource sharing and social tolerance is not necessarily reliant upon human-like cultural norms.     

Vocal culture in New Caledonian crows Corvus moneduloides

While recent research suggests that some animal species may possess ‘cultural’ traditions, much of the current evidence for wild populations remains contentious. This is largely due to the difficulty of demonstrating a fundamental prerequisite for the existence of culture: social learning. As the only case where social learning has been demonstrated conclusively, and subsequently linked to spatial or temporal trait variation, avian vocal dialects are the best studied, and most widely accepted, form of animal culture. Here, we investigate the potential for vocal culture in one of the few animals for which material culture has been suggested: the New Caledonian crow Corvus moneduloides. We show that this species: (1) possesses the capacity for social learning of vocalizations (experimental evidence in the form of a captive subject that reproduces human speech and other anthropogenic noises); and (2) exhibits significant large‐scale, population‐level variation in its vocalizations (cross‐island playback experiments, with analyses controlling for a substantial set of potentially confounding variables). In combination, this provides strong evidence for the existence of ‘culture’ in these birds. More specifically, our findings reveal that the species exhibits sufficient social learning mechanisms, and within‐population structuring, to generate and perpetuate cultural variation in at least one behavioural domain. This is a critical first step towards demonstrating cultural transmission in other behaviours, including tool manufacture and tool use, opening the door for the simultaneous investigation of vocal and material culture in a nonhuman species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 767–776.     

Sex Differences in Object Manipulation in Wild Immature Chimpanzees (Pan troglodytes schweinfurthii) and Bonobos (Pan paniscus): Preparation for Tool Use?

Sex differences in immatures predict behavioural differences in adulthood in many mammal species. Because most studies have focused on sex differences in social interactions, little is known about possible sex differences in ‘preparation’ for adult life with regards to tool use skills. We investigated sex and age differences in object manipulation in immature apes. Chimpanzees use a variety of tools across numerous contexts, whereas bonobos use few tools and none in foraging. In both species, a female bias in adult tool use has been reported. We studied object manipulation in immature chimpanzees at Kalinzu (Uganda) and bonobos at Wamba (Democratic Republic of Congo). We tested predictions of the ‘preparation for tool use’ hypothesis. We confirmed that chimpanzees showed higher rates and more diverse types of object manipulation than bonobos. Against expectation, male chimpanzees showed higher object manipulation rates than females, whereas in bonobos no sex difference was found. However, object manipulation by male chimpanzees was play-dominated, whereas manipulation types of female chimpanzees were more diverse (e.g., bite, break, carry). Manipulation by young immatures of both species was similarly dominated by play, but only in chimpanzees did it become more diverse with age. Moreover, in chimpanzees, object types became more tool-like (i.e., sticks) with age, further suggesting preparation for tool use in adulthood. The male bias in object manipulation in immature chimpanzees, along with the late onset of tool-like object manipulation, indicates that not all (early) object manipulation (i.e., object play) in immatures prepares for subsistence tool use. Instead, given the similarity with gender differences in human children, object play may also function in motor skill practice for male-specific behaviours (e.g., dominance displays). In conclusion, even though immature behaviours almost certainly reflect preparation for adult roles, more detailed future work is needed to disentangle possible functions of object manipulation during development.     

Behavioral diversity in populations of free-ranging chimpanzees in Africa: is it culture?

Behavioral differences exist among populations of wild chimpanzees (Pan troglodytes, Hominoidea) across Africa. Deciding if these are cross-cultural differences demands careful interpretation and rigorous scrutiny. Even standard patterns like social grooming may show variation in detail between neighboring populations. Some patterns of tool use, such as using hammers to crack nuts, also vary across far western Africa. At least 37 populations of free-ranging chimpanzees show tool use, and 14 show at least one habitual pattern of tool use. Regional differences exist among the three subspecies or geographical races of chimpanzees. Convincing empirical demonstration of social customs and traditions in non-human species is problematical, and even something so simple in principle as innovation is hard to show in practice. However, culture need not depend on imitation, teaching, or language, either in humans or in other species.