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	Author	Sousa, C.; Okamoto, S.; Matsuzawa, T.
	Title	Behavioural development in a matching-to-sample task and token use by an infant chimpanzee reared by his mother			Type	Journal Article
	Year	2003	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	6	Issue	4	Pages	259-267
	Keywords	Animals; Cognition; Discrimination Learning; Female; Imitative Behavior; Male; Mothers/psychology; Pan troglodytes/growth & development/psychology; *Transfer (Psychology)
	Abstract	We investigated the behavioural and cognitive development of a captive male infant chimpanzee, Ayumu, raised by his mother, Ai. Here we report Ayumu's achievements up to the age of 2 years and 3 months, in the context of complex computer-controlled tasks. From soon after birth, Ayumu had been present during an experiment performed by his mother. The task consisted of two phases, a matching-to-sample task in which she received token rewards, and the insertion of these tokens into a vending machine to obtain food rewards. Ayumu himself received no reward or encouragement from humans for any of the actions he exhibited during the experiment. At the age of 9 months and 3 weeks, Ayumu performed his first matching-to-sample trial. At around 1 year and 3 months, he began to perform them consistently. Also during this period, he frequently stole food rewards from his mother. At 2 years and 3 months, Ayumu succeeded for the first time in inserting a token into the vending machine. Once he had succeeded in using a token, he performed both phases of the task in sequence 20 times consecutively. The infant's behaviour was not shaped by food rewards but by a strong motivation to copy his mother's behaviour. Our observations of Ayumu thus mirror the learning processes shown by wild chimpanzees.
	Address	Primate Research Institute, Kyoto University, Inuyama Aichi 484-8506, Japan. csousa@fcsh.unl.pt
	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:13680400			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2556
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	Author	Hirata, S.; Celli, M.L.
	Title	Role of mothers in the acquisition of tool-use behaviours by captive infant chimpanzees			Type	Journal Article
	Year	2003	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	6	Issue	4	Pages	235-244
	Keywords	Animals; Cooking and Eating Utensils; Feeding Behavior; Female; Imitative Behavior/physiology; Male; Mothers/psychology; Motor Skills/physiology; Pan troglodytes/growth & development/psychology; Problem Solving/physiology
	Abstract	This article explores the maternal role in the acquisition of tool-use behaviours by infant chimpanzees ( Pan troglodytes). A honey-fishing task, simulating ant/termite fishing found in the wild, was introduced to three dyads of experienced mother and naive infant chimpanzees. Four fishing sites and eight sets of 20 objects to be used as tools, not all appropriate, were available. Two of the mothers constantly performed the task, using primarily two kinds of tools; the three infants observed them. The infants, regardless of the amount of time spent observing, successfully performed the task around the age of 20-22 months, which is earlier than has been recorded in the wild. Two of the infants used the same types of tools that the adults predominantly used, suggesting that tool selectivity is transmitted. The results also show that adults are tolerant of infants, even if unrelated; infants were sometimes permitted to lick the tools, or were given the tools, usually without honey, as well as permitted to observe the adult performances closely.
	Address	Primate Research Institute, Kyoto University, Inuyama, Kanrin, 484-8506 Aichi, Japan. hirata@gari.be.to
	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:13680401			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2555
Permanent link to this record



	Author	Biro, D.; Inoue-Nakamura, N.; Tonooka, R.; Yamakoshi, G.; Sousa, C.; Matsuzawa, T.
	Title	Cultural innovation and transmission of tool use in wild chimpanzees: evidence from field experiments			Type	Journal Article
	Year	2003	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	6	Issue	4	Pages	213-223
	Keywords	Animals; Cooking and Eating Utensils; Culture; Diffusion of Innovation; Feeding Behavior/psychology; Female; Functional Laterality; Imitative Behavior; Male; Motor Skills; Pan troglodytes/psychology; Social Environment
	Abstract	Chimpanzees (Pan troglodytes) are the most proficient and versatile users of tools in the wild. How such skills become integrated into the behavioural repertoire of wild chimpanzee communities is investigated here by drawing together evidence from three complementary approaches in a group of oil-palm nut- ( Elaeis guineensis) cracking chimpanzees at Bossou, Guinea. First, extensive surveys of communities adjacent to Bossou have shown that population-specific details of tool use, such as the selection of species of nuts as targets for cracking, cannot be explained purely on the basis of ecological differences. Second, a 16-year longitudinal record tracing the development of nut-cracking in individual chimpanzees has highlighted the importance of a critical period for learning (3-5 years of age), while the similar learning contexts experienced by siblings have been found to result in near-perfect (13 out of 14 dyads) inter-sibling correspondence in laterality. Third, novel data from field experiments involving the introduction of unfamiliar species of nuts to the Bossou group illuminates key aspects of both cultural innovation and transmission. We show that responses of individuals toward the novel items differ markedly with age, with juveniles being the most likely to explore. Furthermore, subjects are highly specific in their selection of conspecifics as models for observation, attending to the nut-cracking activities of individuals in the same age group or older, but not younger than themselves. Together with the phenomenon of inter-community migration, these results demonstrate a mechanism for the emergence of culture in wild chimpanzees.
	Address	Primate Research Institute, Kyoto University, Japan. dora.biro@zoology.oxford.ac.uk
	Corporate Author				Thesis
	Publisher	Springer-Verlag	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:12898285			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2560
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	Author	Izumi, A.; Kojima, S.
	Title	Matching vocalizations to vocalizing faces in a chimpanzee (Pan troglodytes)			Type	Journal Article
	Year	2004	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	7	Issue	3	Pages	179-184
	Keywords	Acoustic Stimulation; Animal Communication; Animals; Discrimination Learning; Facial Expression; Female; Individuality; Pan troglodytes/psychology; Photic Stimulation; Recognition (Psychology); Vocalization, Animal
	Abstract	Auditory-visual processing of species-specific vocalizations was investigated in a female chimpanzee named Pan. The basic task was auditory-visual matching-to-sample, where Pan was required to choose the vocalizer from two test movies in response to a chimpanzee's vocalization. In experiment 1, movies of vocalizing and silent faces were paired as the test movies. The results revealed that Pan recognized the status of other chimpanzees whether they vocalized or not. In experiment 2, two different types of vocalizing faces of an identical individual were prepared as the test movies. Pan recognized the correspondence between vocalization types and faces. These results suggested that chimpanzees possess crossmodal representations of their vocalizations, as do humans. Together with the ability of vocal individual recognition, this ability might reflect chimpanzees' profound understanding of the status of other individuals.
	Address	Primate Research Institute, Kyoto University, Kanrin, Inuyama, 484-8506, Aichi, Japan. izumi@pri.kyoto-u.ac.jp
	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:15015035			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2541
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	Author	Langergraber, K.; Mitani, J.; Vigilant, L.
	Title	Kinship and social bonds in female chimpanzees (Pan troglodytes)			Type	Journal Article
	Year	2009	Publication	American journal of primatology	Abbreviated Journal	Am. J. Primatol.
	Volume	71	Issue	10	Pages	840-851
	Keywords	Animals; Family; Female; Male; Pan troglodytes/physiology/psychology; *Social Distance
	Abstract	A large body of theoretical and empirical research suggests that kinship influences the development and maintenance of social bonds among group-living female mammals, and that human females may be unusual in the extent to which individuals form differentiated social relationships with nonrelatives. Here we combine behavioral observations of party association, spatial proximity, grooming, and space use with extensive molecular genetic analyses to determine whether female chimpanzees form strong social bonds with unrelated individuals of the same sex. We compare our results with those obtained from male chimpanzees who live in the same community and have been shown to form strong social bonds with each other. We demonstrate that party association is as good a predictor of spatial proximity and grooming in females as it is in males, that the highest party association indices are consistently found between female dyads, that the sexes do not differ in the long-term stability of their party association patterns, and that these results cannot be explained as a by-product of the tendency of females to selectively range in particular areas of the territory. We also show that close kin (i.e. mother-daughter and sibling dyads) are very rare, indicating that the vast majority of female dyads that form strong social bonds are not closely related. Additional analyses reveal that “subgroups” of females, consisting of individuals who frequently associate with one another in similar areas of the territory, do not consist of relatives. This suggests that a passive form of kin-biased dispersal, involving the differential migration of females from neighboring communities into subgroups, was also unlikely to be occurring. These results show that, as in males, kinship plays a limited role in structuring the intrasexual social relationships of female chimpanzees.
	Address	Primatology Department, Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany. langergraber@eva.mpg.de
	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	0275-2565	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:19475543			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5166
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	Author	Flack, J.C.; Jeannotte, L.A.; de Waal, F.B.M.
	Title	Play signaling and the perception of social rules by juvenile chimpanzees (Pan troglodytes)			Type	Journal Article
	Year	2004	Publication	Journal of comparative psychology (Washington, D.C. : 1983)	Abbreviated Journal	J Comp Psychol
	Volume	118	Issue	2	Pages	149-159
	Keywords	Age Factors; Animals; Behavior, Animal; Female; Male; Pan troglodytes; Play and Playthings; Recognition (Psychology); Signal Detection (Psychology); *Social Perception
	Abstract	Prescriptive social rules are enforced statistical regularities. The authors investigated whether juvenile chimpanzees (Pan troglodytes) recognize and use enforced statistical regularities to guide dyadic play behavior. They hypothesized (a) that proximity of adults, especially mothers of younger play partners, to play bouts will increase the play signaling of older partners and (b) that when juvenile-juvenile play bouts occur in proximity to adults, older partners will play at a lower intensity than when no adults are present. They found that older and younger partners increase their play signaling in the presence of the mothers of younger partners, particularly as the intensity of play bouts increases. In contrast to their hypothesis, older partners played more roughly when the mothers of younger partners were in proximity. These results suggest that juvenile chimpanzees increase play signaling to prevent termination of the play bouts by mothers of younger partners.
	Address	Santa Fe Institute, NM 97501, USA. jflack@santafe.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	0735-7036	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15250802			Approved	no
	Call Number	refbase @ user @			Serial	172
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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.; 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
Permanent link to this record



	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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