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Custance, D., Whiten, A., Sambrook, T., & Galdikas, B. (2001). Testing for social learning in the “artificial fruit” processing of wildborn orangutans (Pongo pygmaeus), Tanjung Puting, Indonesia. Anim. Cogn., 4(3), 305–313.
Abstract: Social learning about actions, objects and sequencing was investigated in a group of 14 wildborn orangutans (four adult females and ten 3- to 5-year-old juveniles). Human models showed alternative methods and sequences for dismantling an artificial fruit to groups of participants matched by gender and age. Each participant received three to six 2-min trials in which they were given access to the artificial fruit for manipulation. Independent coders, who were unaware of which method each participant had seen, gave confidence ratings and collected action frequencies from watching video recordings of the experimental trials. No significant differences were found between groups in terms of the coders' confidence ratings, the action frequencies or the sequence of manipulations. These negative results may at least partly reflect the immaturity of a large proportion of the participants. A positive correlation was found between age and the degree of matching to the method shown. Although none of the juveniles succeeded in opening the “fruit”, two out of the four adults did so and they also seemed to match more closely the sequence of elements touched over successive trials. The results are compared with similar data previously collected from human children, chimpanzees, gorillas, capuchin monkeys and common marmosets.
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Caldwell, C. A., & Whiten, A. (2004). Testing for social learning and imitation in common marmosets, Callithrix jacchus, using an artificial fruit. Anim. Cogn., 7(2), 77–85.
Abstract: We tested for social learning and imitation in common marmosets using an artificial foraging task and trained conspecific demonstrators. We trained a demonstrator marmoset to open an artificial fruit, providing a full demonstration of the task to be learned. Another marmoset provided a partial demonstration, controlling for stimulus enhancement effects, by eating food from the outside of the apparatus. We thus compared three observer groups, each consisting of four animals: those that received the full demonstration, those that received the partial demonstration, and a control group that saw no demonstration prior to testing. Although none of the observer marmosets succeeded in opening the artificial fruit during the test periods, there were clear effects of demonstration type. Those that saw the full demonstration manipulated the apparatus more overall, whereas those from the control group manipulated it the least of the three groups. Those from the full-demonstration group also contacted the particular parts of the artificial fruit that they had seen touched (localised stimulus enhancement) to a greater extent than the other two groups. There was also an interaction between the number of hand and mouth touches made to the artificial fruit for the full- and partial-demonstration groups. Whether or not these data represent evidence for imitation is discussed. We also propose that the clear differences between the groups suggest that social learning mechanisms provide real benefits to these animals in terms of developing novel food-processing skills analogous to the one presented here.
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Caldwell, C. A., & Whiten, A. (2002). Evolutionary perspectives on imitation: is a comparative psychology of social learning possible? Anim. Cogn., 5(4), 193–208.
Abstract: Studies of imitation in animals have become numerous in recent times, but do they contribute to a comparative psychology of social learning? We review this burgeoning field to identify the problems and prospects for such a goal. Difficulties of two main kinds are identified. First, researchers have tackled questions about social learning from at least three very different theoretical perspectives, the “phylogenetic”, “animal model”, and “adaptational”. We examine the conflicts between them and consider the scope for integration. A second difficulty arises in the methodological approaches used in the discipline. In relation to one of these – survey reviews of published studies – we tabulate and compare the contrasting conclusions of nine articles that together review 36 studies. The basis for authors' disagreements, including the matters of perceptual opacity, novelty, sequential structure, and goal representation, are examined. In relation to the other key method, comparative experimentation, we identify 12 studies that have explicitly compared species' imitative ability on similar tasks. We examine the principal problems of comparing like with like in these studies and consider solutions, the most powerful of which we propose to be the use of a systematic range of task designs, rather than any single “gold standard” task.
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Barton, R. A., Byrne, R. W., & Whiten, A. (1996). Ecology, feeding competition and social structure in baboons. Behav. Ecol. Sociobiol., 38(5), 321–329.
Abstract: Predictions of the model of van Schaik (1989) of female-bonding in primates are tested by systematically comparing the ecology, level of within-group contest competition for food (WGC), and patterns of social behaviour found in two contrasting baboon populations. Significant differences were found in food distribution (percentage of the diet from clumped sources), feeding supplant rates and grooming patterns. In accord with the model, the tendencies of females to affiliate and form coalitions with one another, and to be philopatric, were strongest where ecological conditions promoted WGC. Group fission in the population with strong WGC was “horizontal” with respect to female dominance rank, and associated with female-female aggression during a period of elevated feeding competition. In contrast, where WGC was low, females' grooming was focused on adult males rather than other females. Recent evidence suggests that group fission here is initiated by males, tends to result in the formation of one-male groups, and is not related to feeding competition but to male-male competition for mates. An ecological model of baboon social structure is presented which incorporates the effects of female-female competition, male-male competition, and predation pressure. The model potentially accounts for wide variability in group size, group structure and social relationships within the genus Papio. Socio-ecological convergence between common baboons and hamadryas baboons, however, may be limited in some respects by phylogenetic inertia.
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Whiten, A. (2000). Social complexity and social intelligence. In Novartis Foundation Symposium (Vol. 233, pp. 185–96; discussion pp. 196–201).
Abstract: When we talk of the 'nature of intelligence', or any other attribute, we may be referring to its essential structure, or to its place in nature, particularly the function it has evolved to serve. Here I examine both, from the perspective of the evolution of intelligence in primates. Over the last 20 years, the Social (or 'Machiavellian') Intelligence Hypothesis has gained empirical support. Its core claim is that the intelligence of primates is primarily an adaptation to the special complexities of primate social life. In addition to this hypothesis about the function of intellect, a secondary claim is that the very structure of intelligence has been moulded to be 'social' in character, an idea that presents a challenge to orthodox views of intelligence as a general-purpose capacity. I shall outline the principal components of social intelligence and the environment of social complexity it engages with. This raises the question of whether domain specificity is an appropriate characterization of social intelligence and its subcomponents, like theory of mind. As a counter-argument to such specificity I consider the hypothesis that great apes exhibit a cluster of advanced cognitive abilities that rest on a shared capacity for second-order mental representation.
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Whiten, A., & Ham, R. (1992). On the nature and evolution of imitation in the animal kingdom: reappraisal of a century of research. Adv. Study Behav., 21, 239–283.
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Byrne, R. W., & Whiten, A. (1990). Tactical deception in primates: the 1990 database (Vol. 27). German Primate Center.
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Whiten, A., & van Schaik, C. P. (2007). The evolution of animal 'cultures' and social intelligence. Philos Trans R Soc Lond B Biol Sci, 362(1480), 603–620.
Abstract: Decades-long field research has flowered into integrative studies that, together with experimental evidence for the requisite social learning capacities, have indicated a reliance on multiple traditions ('cultures') in a small number of species. It is increasingly evident that there is great variation in manifestations of social learning, tradition and culture among species, offering much scope for evolutionary analysis. Social learning has been identified in a range of vertebrate and invertebrate species, yet sustained traditions appear rarer, and the multiple traditions we call cultures are rarer still. Here, we examine relationships between this variation and both social intelligence-sophisticated information processing adapted to the social domain-and encephalization. First, we consider whether culture offers one particular confirmation of the social ('Machiavellian') intelligence hypothesis that certain kinds of social life (here, culture) select for intelligence: 'you need to be smart to sustain culture'. Phylogenetic comparisons, particularly focusing on our own study animals, the great apes, support this, but we also highlight some paradoxes in a broader taxonomic survey. Second, we use intraspecific variation to address the converse hypothesis that 'culture makes you smart', concluding that recent evidence for both chimpanzees and orang-utans support this proposition.
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Horner, V., Whiten, A., Flynn, E., & de Waal, F. B. M. (2006). Faithful replication of foraging techniques along cultural transmission chains by chimpanzees and children. Proc. Natl. Acad. Sci. U.S.A., 103(37), 13878–13883.
Abstract: Observational studies of wild chimpanzees (Pan troglodytes) have revealed population-specific differences in behavior, thought to represent cultural variation. Field studies have also reported behaviors indicative of cultural learning, such as close observation of adult skills by infants, and the use of similar foraging techniques within a population over many generations. Although experimental studies have shown that chimpanzees are able to learn complex behaviors by observation, it is unclear how closely these studies simulate the learning environment found in the wild. In the present study we have used a diffusion chain paradigm, whereby a behavior is passed from one individual to the next in a linear sequence in an attempt to simulate intergenerational transmission of a foraging skill. Using a powerful three-group, two-action methodology, we found that alternative methods used to obtain food from a foraging device (“lift door” versus “slide door”) were accurately transmitted along two chains of six and five chimpanzees, respectively, such that the last chimpanzee in the chain used the same method as the original trained model. The fidelity of transmission within each chain is remarkable given that several individuals in the no-model control group were able to discover either method by individual exploration. A comparative study with human children revealed similar results. This study is the first to experimentally demonstrate the linear transmission of alternative foraging techniques by non-human primates. Our results show that chimpanzees have a capacity to sustain local traditions across multiple simulated generations.
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Whiten, A., Horner, V., & de Waal, F. B. M. (2005). Conformity to cultural norms of tool use in chimpanzees. Nature, 437(7059), 737–740.
Abstract: Rich circumstantial evidence suggests that the extensive behavioural diversity recorded in wild great apes reflects a complexity of cultural variation unmatched by species other than our own. However, the capacity for cultural transmission assumed by this interpretation has remained difficult to test rigorously in the field, where the scope for controlled experimentation is limited. Here we show that experimentally introduced technologies will spread within different ape communities. Unobserved by group mates, we first trained a high-ranking female from each of two groups of captive chimpanzees to adopt one of two different tool-use techniques for obtaining food from the same 'Pan-pipe' apparatus, then re-introduced each female to her respective group. All but two of 32 chimpanzees mastered the new technique under the influence of their local expert, whereas none did so in a third population lacking an expert. Most chimpanzees adopted the method seeded in their group, and these traditions continued to diverge over time. A subset of chimpanzees that discovered the alternative method nevertheless went on to match the predominant approach of their companions, showing a conformity bias that is regarded as a hallmark of human culture.
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