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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	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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	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
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	Author	Bonnie, K.E.; Horner, V.; Whiten, A.; de Waal, F.B.M.
	Title	Spread of arbitrary conventions among chimpanzees: a controlled experiment			Type	Journal Article
	Year	2007	Publication	Proceedings. Biological sciences / The Royal Society	Abbreviated Journal	Proc Biol Sci
	Volume	274	Issue	1608	Pages	367-372
	Keywords
	Abstract	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.
	Address	Living Links, Yerkes National Primate Research Center, Atlanta, GA 30329, USA. kebonni@emory.edu
	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	0962-8452	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:17164200			Approved	no
	Call Number	refbase @ user @			Serial	157
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	Author	Horner, V.; Whiten, A.
	Title	*Learning from others' mistakes limits on understanding a trap-tube task by young chimpanzees (Pan troglodytes) and children (Homo sapiens)*			Type	Journal Article
	Year	2007	Publication	Journal of comparative psychology	Abbreviated Journal	J Comp Psychol
	Volume	121	Issue	1	Pages	12-21
	Keywords
	Abstract	A trap-tube task was used to determine whether chimpanzees (Pan troglodytes) and children (Homo sapiens) who observed a model's errors and successes could master the task in fewer trials than those who saw only successes. Two- to 7-year-old chimpanzees and 3- to 4-year-old children did not benefit from observing errors and found the task difficult. Two of the 6 chimpanzees developed a successful anticipatory strategy but showed no evidence of representing the core causal relations involved in trapping. Three- to 4-year-old children showed a similar limitation and tended to copy the actions of the demonstrator, irrespective of their causal relevance. Five- to 6-year-old children were able to master the task but did not appear to be influenced by social learning or benefit from observing errors.
	Address	Centre for Social Learning and Cognitive Evolution, School of Psychology, University of St Andrews, Fife, Scotland, UK. vhorner@rmy.emory.edu
	Corporate Author				Thesis
	Publisher		Place of Publication	Washington, D.C. : 1983	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:17324071			Approved	yes
	Call Number	refbase @ user @			Serial	728
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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	Caldwell, C.A.; Whiten, A.
	Title	Testing for social learning and imitation in common marmosets, Callithrix jacchus, using an artificial fruit			Type	Journal Article
	Year	2004	Publication	Animal cognition	Abbreviated Journal	Anim. Cogn.
	Volume	7	Issue	2	Pages	77-85
	Keywords	Animals; Association Learning; Callithrix/psychology; Discrimination Learning; Feeding Behavior; Female; Food Preferences; Fruit; Imitative Behavior; Male; *Social Behavior; Social Environment
	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.
	Address	Centre for Social Learning and Cognitive Evolution and Scottish Primate Research Group, University of St Andrews, KY16 9JU, St Andrews, Fife, Scotland. 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:15069606			Approved	no
	Call Number	refbase @ user @			Serial	735
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	Author	Castles, D.L.; Whiten, A.; Aureli, F.
	Title	Social anxiety, relationships and self-directed behaviour among wild female olive baboons			Type	Journal Article
	Year	1999	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	58	Issue	6	Pages	1207-1215
	Keywords
	Abstract	Self-directed behaviour (SDB) can be used as a behavioural indicator of stress and anxiety in nonhuman primates (Maestripieri et al. 1992, Animal Behaviour, 44, 967-979). We investigated the effect of nearest neighbours' relative dominance status on the SDB of sexually mature female olive baboons, Papio anubis. When the animal nearest to (within 5 m of) a female was a dominant individual, SDB rates (a combined measure of self-scratching, self-grooming, self-touching, body shaking and yawning) increased by ca. 40% over those observed when the nearest neighbour was a subordinate. The results indicate that (1) SDB can be used as a measure of uncertainty during the social interactions of cercopithecine primates and (2) as there was considerable variation in SDB response according to the nature of the dominant individual, SDB can be used to assess relationship security (i.e. the perceived predictability of a relationship for one partner). Finally, in combination with measures of affiliation rate, SDB may provide insight into relationship value.
	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	745
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	Author	Whiten, A.; Custance, D.M.; Gomez, J.C.; Teixidor, P.; Bard, K.A.
	Title	Imitative learning of artificial fruit processing in children (Homo sapiens) and chimpanzees (Pan troglodytes)			Type	Journal Article
	Year	1996	Publication	Journal of comparative psychology (Washington, D.C. : 1983)	Abbreviated Journal	J Comp Psychol
	Volume	110	Issue	1	Pages	3-14
	Keywords	Animals; Child, Preschool; Discrimination Learning; Female; Food Preferences/psychology; Fruit; Humans; Imitative Behavior; Male; Mental Recall; Pan troglodytes/psychology; Social Environment
	Abstract	Observational learning in chimpanzees and young children was investigated using an artificial fruit designed as an analog of natural foraging problems faced by primates. Each of 3 principal components could be removed in 2 alternative ways, demonstration of only one of which was watched by each subject. This permitted subsequent imitation by subjects to be distinguished from stimulus enhancement. Children aged 2-4 years evidenced imitation for 2 components, but also achieved demonstrated outcomes through their own techniques. Chimpanzees relied even more on their own techniques, but they did imitate elements of 1 component of the task. To our knowledge, this is the first experimental evidence of chimpanzee imitation in a functional task designed to simulate foraging behavior hypothesized to be transmitted culturally in the wild.
	Address	Scottish Primate Research Group, University of St. Andrews, Fife, Scotland. aw2@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:8851548			Approved	no
	Call Number	refbase @ user @			Serial	744
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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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