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	Author	Assersohn, C.; Whiten, A.; Kiwede, Z.T.; Tinka, J.; Karamagi, J.
	Title	Use of leaves to inspect ectoparasites in wild chimpanzees: a third cultural variant?			Type	Journal Article
	Year	2004	Publication	Primates	Abbreviated Journal	Primates
	Volume	45	Issue	4	Pages	255-258
	Keywords	Animals; Animals, Wild/physiology; Ape Diseases/parasitology; Behavior, Animal/physiology; Ectoparasitic Infestations/parasitology/veterinary; Female; Grooming/physiology; Male; Pan troglodytes/physiology; Plant Leaves; Protozoa/*isolation & purification; Uganda
	Abstract	We report 26 cases of using leaves as tools with which wild chimpanzees (Pan troglodytes schweinfurthii) in the Sonso community, Budongo Forest, Uganda, appeared to inspect objects removed during grooming. Careful removal of potential ectoparasites and delicate lip or manual placement on leaves followed by intense visual examination characterised this behaviour. It appears to be done to judge whether either ingestion or discarding is most appropriate, the former occurring in most cases. This behaviour may represent a third variant of ectoparasite handling, different from those described at Tai and Gombe, yet sharing features with the latter. These two East African techniques may thus have evolved from leaf grooming.
	Address	Scottish Primate Research Group, School of Psychology, University of St Andrews, St Andrews, KY16 9JU, Fife, 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	0032-8332	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15179558			Approved	no
	Call Number	refbase @ user @			Serial	733
Permanent link to this record



	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
Permanent link to this record



	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
Permanent link to this record



	Author	Horowitz, A.C.
	Title	Do humans ape? Or do apes human? Imitation and intention in humans (Homo sapiens) and other animals			Type	Journal Article
	Year	2003	Publication	Journal of comparative psychology	Abbreviated Journal	J Comp Psychol
	Volume	117	Issue	3	Pages	325-336
	Keywords	Adolescent; Adult; Animals; Appetitive Behavior; Attention; Child, Preschool; Concept Formation; Female; Humans; Imitative Behavior; Male; Motivation; Pan troglodytes/psychology; Problem Solving; *Psychomotor Performance; Reaction Time; Species Specificity
	Abstract	A. Whiten, D. M. Custance, J.-C. Gomez, P. Teixidor, and K. A. Bard (1996) tested chimpanzees' (Pan troglodytes) and human children's (Homo sapiens) skills at imitation with a 2-action test on an “artificial fruit.” Chimpanzees imitated to a restricted degree; children were more thoroughly imitative. Such results prompted some to assert that the difference in imitation indicates a difference in the subjects' understanding of the intentions of the demonstrator (M. Tomasello, 1996). In this experiment, 37 adult human subjects were tested with the artificial fruit. Far from being perfect imitators, the adults were less imitative than the children. These results cast doubt on the inference from imitative performance to an ability to understand others' intentions. The results also demonstrate how any test of imitation requires a control group and attention to the level of behavioral analysis.
	Address	Department of Cognitive Science, University of California, San Diego, CA, USA. ahorowitz@crl.ucsd.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:14498809			Approved	yes
	Call Number	refbase @ user @			Serial	736
Permanent link to this record



	Author	Stoinski, T.S.; Whiten, A.
	Title	Social learning by orangutans (Pongo abelii and Pongo pygmaeus) in a simulated food-processing task			Type	Journal Article
	Year	2003	Publication	Journal of comparative psychology (Washington, D.C. : 1983)	Abbreviated Journal	J Comp Psychol
	Volume	117	Issue	3	Pages	272-282
	Keywords	Animal Communication; Animals; Feeding Behavior; Female; Imitative Behavior; Male; Pongo pygmaeus/psychology; Problem Solving; Psychomotor Performance; Social Environment; Species Specificity; *Transfer (Psychology)
	Abstract	Increasing evidence for behavioral differences between populations of primates has created a resurgence of interest in examining mechanisms of information transfer between individuals. The authors examined the social transmission of information in 15 captive orangutans (Pongo abelii and Pongo pygmaeus) using a simulated food-processing task. Experimental subjects were shown 1 of 2 methods for removing a suite of defenses on an “artificial fruit.” Control subjects were given no prior exposure before interacting with the fruit. Observing a model provided a functional advantage in the task, as significantly more experimental than control subjects opened the fruit. Within the experimental groups, the authors found a trend toward differences in the actual behaviors used to remove 1 of the defenses. Results support observations from the wild implying horizontal transfer of information in orangutans and show that a number of social learning processes are likely to be involved in the transfer of knowledge in this species.
	Address	Department of Primate Research, Zoo Atlanta, Atlanta, GA 30315, USA. tstoinski@zooatlanta.org
	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:14498803			Approved	no
	Call Number	refbase @ user @			Serial	737
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	Author	Stoinski, T.S.; Wrate, J.L.; Ure, N.; Whiten, A.
	Title	Imitative learning by captive western lowland gorillas (Gorilla gorilla gorilla) in a simulated food-processing task			Type	Journal Article
	Year	2001	Publication	Journal of comparative psychology (Washington, D.C. : 1983)	Abbreviated Journal	J Comp Psychol
	Volume	115	Issue	3	Pages	272-281
	Keywords	Animals; Behavior, Animal/physiology; Discrimination Learning; Feeding Behavior/physiology; Gorilla gorilla; *Imitative Behavior; Social Behavior
	Abstract	Although field studies have suggested the existence of cultural transmission of foraging techniques in primates, identification of transmission mechanisms has remained elusive. To test experimentally for evidence of imitation in the current study, we exposed gorillas (Gorilla gorilla gorilla) to an artificial fruit foraging task designed by A. Whiten and D. M. Custance (1996). Gorillas (n=6) watched a human model remove a series of 3 defenses around a fruit. Each of the defenses was removed using 1 of 2 alternative techniques. Subsequent video analysis of gorillas' behavior showed a significant tendency to copy the observed technique on 1 of the individual defenses and the direction of removal on another defense. This is the first statistically reliable evidence of imitation in gorillas. Sequence of defense removal was not replicated. The gorillas' responses were most similar to those of chimpanzees.
	Address	TECHlab, Zoo Atlanta, Georgia 30315, USA. stoinskit@mindspring.com
	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:11594496			Approved	no
	Call Number	refbase @ user @			Serial	738
Permanent link to this record



	Author	Whiten, A.; McGrew, W.C.
	Title	Is this the first portrayal of tool use by a chimp?			Type
	Year	2001	Publication	Nature	Abbreviated Journal	Nature
	Volume	409	Issue	6816	Pages	12
	Keywords	Animals; Behavior, Animal; Pan troglodytes/physiology; Philately
	Abstract
	Address
	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:11343083			Approved	no
	Call Number	refbase @ user @			Serial	739
Permanent link to this record



	Author	Whiten, A.; Boesch, C.
	Title	The cultures of chimpanzees			Type	Journal Article
	Year	2001	Publication	Scientific American	Abbreviated Journal	Sci Am
	Volume	284	Issue	1	Pages	60-67
	Keywords	Animals; Behavior, Animal; Culture; Feeding Behavior; Grooming; Hominidae; Humans; Pan troglodytes/physiology
	Abstract
	Address	University of St. Andrews
	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	0036-8733	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:11132425			Approved	no
	Call Number	refbase @ user @			Serial	740
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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
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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