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	Author	Horner, V.; Whiten, A.
	Title	Causal knowledge and imitation/emulation switching in chimpanzees (Pan troglodytes) and children (Homo sapiens)			Type	Journal Article
	Year	2005	Publication	Animal cognition	Abbreviated Journal	Anim. Cogn.
	Volume	8	Issue	3	Pages	164-181
	Keywords	Animals; Child Behavior; Child, Preschool; Concept Formation; Female; Humans; Imitative Behavior; Learning; Male; Pan troglodytes; Problem Solving; Psychomotor Performance; *Social Environment; Species Specificity
	Abstract	This study explored whether the tendency of chimpanzees and children to use emulation or imitation to solve a tool-using task was a response to the availability of causal information. Young wild-born chimpanzees from an African sanctuary and 3- to 4-year-old children observed a human demonstrator use a tool to retrieve a reward from a puzzle-box. The demonstration involved both causally relevant and irrelevant actions, and the box was presented in each of two conditions: opaque and clear. In the opaque condition, causal information about the effect of the tool inside the box was not available, and hence it was impossible to differentiate between the relevant and irrelevant parts of the demonstration. However, in the clear condition causal information was available, and subjects could potentially determine which actions were necessary. When chimpanzees were presented with the opaque box, they reproduced both the relevant and irrelevant actions, thus imitating the overall structure of the task. When the box was presented in the clear condition they instead ignored the irrelevant actions in favour of a more efficient, emulative technique. These results suggest that emulation is the favoured strategy of chimpanzees when sufficient causal information is available. However, if such information is not available, chimpanzees are prone to employ a more comprehensive copy of an observed action. In contrast to the chimpanzees, children employed imitation to solve the task in both conditions, at the expense of efficiency. We suggest that the difference in performance of chimpanzees and children may be due to a greater susceptibility of children to cultural conventions, perhaps combined with a differential focus on the results, actions and goals of the demonstrator.
	Address	Centre for Social Learning and Cognitive Evolution, School of Psychology, University of St Andrews, St Andrews, KY16 9JU, UK. vkh1@st-andrews.ac.uk
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15549502			Approved	no
	Call Number	refbase @ user @			Serial	732
Permanent link to this record



	Author	Whiten, A.; Horner, V.; de Waal, F.B.M.
	Title	Conformity to cultural norms of tool use in chimpanzees			Type	Journal Article
	Year	2005	Publication	Nature	Abbreviated Journal	Nature
	Volume	437	Issue	7059	Pages	737-740
	Keywords	Aging/physiology; Animals; Culture; Feeding Behavior/physiology; Female; Pan troglodytes/physiology/psychology; *Social Conformity; Technology; Time Factors
	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.
	Address	Centre for Social Learning and Cognitive Evolution, School of Psychology, University of St Andrews, St Andrews, Fife, KY16 9JP, UK. a.whiten@st-and.ac.uk
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1476-4687	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16113685			Approved	no
	Call Number	refbase @ user @			Serial	163
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	Author	Whiten, A.; Goodall, J.; McGrew, W.C.; Nishida, T.; Reynolds, V.; Sugiyama, Y.; Tutin, C.E.; Wrangham, R.W.; Boesch, C.
	Title	Cultures in chimpanzees			Type	Journal Article
	Year	1999	Publication	Nature	Abbreviated Journal	Nature
	Volume	399	Issue	6737	Pages	682-685
	Keywords	Animals; Behavior, Animal; Cognition; Culture; Humans; Pan troglodytes/*physiology; Species Specificity
	Abstract	As an increasing number of field studies of chimpanzees (Pan troglodytes) have achieved long-term status across Africa, differences in the behavioural repertoires described have become apparent that suggest there is significant cultural variation. Here we present a systematic synthesis of this information from the seven most long-term studies, which together have accumulated 151 years of chimpanzee observation. This comprehensive analysis reveals patterns of variation that are far more extensive than have previously been documented for any animal species except humans. We find that 39 different behaviour patterns, including tool usage, grooming and courtship behaviours, are customary or habitual in some communities but are absent in others where ecological explanations have been discounted. Among mammalian and avian species, cultural variation has previously been identified only for single behaviour patterns, such as the local dialects of song-birds. The extensive, multiple variations now documented for chimpanzees are thus without parallel. Moreover, the combined repertoire of these behaviour patterns in each chimpanzee community is itself highly distinctive, a phenomenon characteristic of human cultures but previously unrecognised in non-human species.
	Address	Scottish Primate Research Group, School of Psychology, University of St Andrews, UK
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0028-0836	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:10385119			Approved	no
	Call Number	refbase @ user @			Serial	742
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	Author	Barton, R.A.; Byrne, R.W.; Whiten, A.
	Title	Ecology, feeding competition and social structure in baboons			Type	Journal Article
	Year	1996	Publication	Behavioral Ecology and Sociobiology	Abbreviated Journal	Behav. Ecol. Sociobiol.
	Volume	38	Issue	5	Pages	321-329
	Keywords	Key words Ecology – Competition – Group size – Baboons
	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.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	refbase @ user @			Serial	807
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	Author	Caldwell, C.A.; Whiten, A.
	Title	Evolutionary perspectives on imitation: is a comparative psychology of social learning possible?			Type	Journal Article
	Year	2002	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	5	Issue	4	Pages	193-208
	Keywords	Animals; Behavior, Animal; Evolution; Humans; Imitative Behavior; Learning; Models, Animal
	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.
	Address	School of Psychology, Washington Singer Laboratories, University of Exeter, Exeter EX4 4QG, UK. C.A.Caldwell@exeter.ac.uk
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:12461597			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2593
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	Author	Horner, V.; Whiten, A.; Flynn, E.; de Waal, F.B.M.
	Title	Faithful replication of foraging techniques along cultural transmission chains by chimpanzees and children			Type	Journal Article
	Year	2006	Publication	Proceedings of the National Academy of Sciences of the United States of America	Abbreviated Journal	Proc. Natl. Acad. Sci. U.S.A.
	Volume	103	Issue	37	Pages	13878-13883
	Keywords	Animals; Child, Preschool; Humans; Imitative Behavior; Pan troglodytes/psychology
	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.
	Address	Centre for Social Learning and Cognitive Evolution, School of Psychology, University of St. Andrews, Fife KY16 9JP, United Kingdom
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0027-8424	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16938863			Approved	no
	Call Number	refbase @ user @			Serial	159
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	Author	Barton, R.A.; Whiten, A.
	Title	Feeding competition among female olive baboons, Papio anubis			Type	Journal Article
	Year	1993	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	46	Issue	4	Pages	777-789
	Keywords
	Abstract	Abstract. Competition for food is thought to play a key role in the social organization of group-living female primates, leading to the prediction that individual foraging success will be partly regulated by dominance relationships. Among adult females in a group of free-ranging olive baboons, dominance rank was significantly correlated with nutrient acquisition rates (feeding rates and daily intakes), but not with dietary diversity or quality, nor with activity budgets. The mean daily food intake of the three highest-ranking females was 30% greater than that of the three lowest-ranking females, providing an explanation for relationships between female rank and fertility found in a number of other studies of group-living primates. The intensity of feeding competition, as measured by supplant rates and spatial clustering of individuals, increased during the dry season, a period of low food availability, seemingly because foods eaten then were more clumped in distribution than those eaten in the wet season. Implications for models of female social structure and maximum group size are discussed.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4258
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	Author	Whiten, A.; Horner, V.; Litchfield, C.A.; Marshall-Pescini, S.
	Title	How do apes ape?			Type	Journal Article
	Year	2004	Publication	Learning & Behavior	Abbreviated Journal	Learn. Behav.
	Volume	32	Issue	1	Pages	36-52
	Keywords	Adaptation, Psychological; Animals; Behavior, Animal; Hominidae/psychology; Imitative Behavior; Imprinting (Psychology); Learning; Psychological Theory; Social Environment; *Social Facilitation
	Abstract	In the wake of telling critiques of the foundations on which earlier conclusions were based, the last 15 years have witnessed a renaissance in the study of social learning in apes. As a result, we are able to review 31 experimental studies from this period in which social learning in chimpanzees, gorillas, and orangutans has been investigated. The principal question framed at the beginning of this era, Do apes ape? has been answered in the affirmative, at least in certain conditions. The more interesting question now is, thus, How do apes ape? Answering this question has engendered richer taxonomies of the range of social-learning processes at work and new methodologies to uncover them. Together, these studies suggest that apes ape by employing a portfolio of alternative social-learning processes in flexibly adaptive ways, in conjunction with nonsocial learning. We conclude by sketching the kind of decision tree that appears to underlie the deployment of these alternatives.
	Address	Centre for Social Learning and Cognitive Evolution, Scottish Primate Research Group, School of Psychology, University of St. Andrews, St. Andrews, Fife, Scotland. a.whiten@st-and.ac.uk
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1543-4494	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15161139			Approved	no
	Call Number	refbase @ user @			Serial	734
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	Author	Whiten, A.
	Title	Imitation of the sequential structure of actions by chimpanzees			Type	Journal Article
	Year	1998	Publication	J Comp Psychol	Abbreviated Journal
	Volume	11	Issue		Pages
	Keywords
	Abstract
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @ Whiten1998			Serial	6291
Permanent link to this record



	Author	Whiten, A.
	Title	Imitation of the sequential structure of actions by chimpanzees (Pan troglodytes)			Type	Journal Article
	Year	1998	Publication	Journal of comparative psychology (Washington, D.C. : 1983)	Abbreviated Journal	J Comp Psychol
	Volume	112	Issue	3	Pages	270-281
	Keywords	Animals; Appetitive Behavior/physiology; Ceremonial Behavior; Exploratory Behavior/physiology; Female; Fruit; Imitative Behavior/physiology; Learning/physiology; Male; Pan troglodytes/psychology; Practice (Psychology); Problem Solving/*physiology
	Abstract	Imitation was studied experimentally by allowing chimpanzees (Pan troglodytes) to observe alternative patterns of actions for opening a specially designed “artificial fruit.” Like problematic foods primates deal with naturally, with the test fruit several defenses had to be removed to gain access to an edible core, but the sequential order and method of defense removal could be systematically varied. Each subject repeatedly observed 1 of 2 alternative techniques for removing each defense and 1 of 2 alternative sequential patterns of defense removal. Imitation of sequential organization emerged after repeated cycles of demonstration and attempts at opening the fruit. Imitation in chimpanzees may thus have some power to produce cultural convergence, counter to the supposition that individual learning processes corrupt copied actions. Imitation of sequential organization was accompanied by imitation of some aspects of the techniques that made up the sequence.
	Address	Scottish Primate Research Group, School of Psychology, University of St. Andrews, Fife, Scotland. a.whiten@st-andrews.ac.uk
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0735-7036	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:9770315			Approved	no
	Call Number	refbase @ user @			Serial	743
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