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	Author	Newton-Fisher, N.E.; Lee, P.C.
	Title	Grooming reciprocity in wild male chimpanzees			Type	Journal Article
	Year	2011	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	81	Issue	2	Pages	439-446
	Keywords	altruism; biological market theory; Budongo; chimpanzee; grooming; Pan troglodytes
	Abstract	Understanding cooperation between unrelated individuals remains a central problem in animal behaviour; evolutionary mechanisms are debated, and the importance of reciprocity has been questioned. Biological market theory makes specific predictions about the occurrence of reciprocity in social groups; applied to the social grooming of mammals, it predicts reciprocity in the absence of other benefits for which grooming can be exchanged. Considerable effort has been made to test this grooming trade model in nonhuman primates; such studies show mixed results, but may be confounded by kin effects. We examined patterns of reciprocity within and across bouts, and tested predictions of the grooming trade model, among wild male chimpanzees, Pan troglodytes: a system with negligible kin effects. In accord with the model's expectations, we found that some grooming was directed by lower- to higher-ranked individuals, and that, on average, higher-ranked individuals groomed more reciprocally. We found no support, however, for a prediction that more reciprocity should occur between individuals close in rank. For most dyads, reciprocity of effort occurred through unbalanced participation in grooming bouts, but reciprocity varied considerably between dyads and only a small proportion showed strongly reciprocal grooming. Despite this, each male had at least one reciprocal grooming relationship. In bouts where both individuals groomed, effort was matched through mutual grooming, not alternating roles. Our results provide mixed support for the current grooming trade, biological market model, and suggest that it needs to incorporate risks of currency inflation and cheating for species where reciprocity can be achieved through repeated dyadic interactions.
	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	0003-3472	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5329
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	Author	Brosnan, S.F.; de Waal, F.B.M.
	Title	Responses to a simple barter task in chimpanzees, Pan troglodytes			Type	Journal Article
	Year	2005	Publication	Primates	Abbreviated Journal	Primates
	Volume	46	Issue	3	Pages	173-182
	Keywords	Analysis of Variance; Animals; Choice Behavior/physiology; Conditioning (Psychology)/physiology; Learning/physiology; Pan troglodytes/physiology; Reward; Sex Factors; Social Behavior; *Token Economy
	Abstract	Chimpanzees (Pan troglodytes) frequently participate in social exchange involving multiple goods and services of variable value, yet they have not been tested in a formalized situation to see whether they can barter using multiple tokens and rewards. We set up a simple barter economy with two tokens and two associated rewards and tested chimpanzees on their ability to obtain rewards by returning the matching token in situations in which their access to tokens was unlimited or limited. Chimpanzees easily learned to associate value with the tokens, as expected, and did barter, but followed a simple strategy of favoring the higher-value token, regardless of the reward proffered, instead of a more complex but more effective strategy of returning the token that matched the reward. This response is similar to that shown by capuchin monkeys in our previous study. We speculate that this response, while not ideal, may be sufficient to allow for stability of the social exchange system in these primates, and that the importance of social barter to both species may have led to this convergence of strategies.
	Address	Living Links, Yerkes National Primate Research Center, Atlanta, GA 30322, USA. sbrosna@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	0032-8332	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15824938			Approved	no
	Call Number	refbase @ user @			Serial	167
Permanent link to this record



	Author	Baker, K.C.; Seres, E.; Aureli, F.; De Waal, F.B.
	Title	Injury risks among chimpanzees in three housing conditions			Type	Journal Article
	Year	2000	Publication	American journal of primatology	Abbreviated Journal	Am. J. Primatol.
	Volume	51	Issue	3	Pages	161-175
	Keywords	Aggression; Animal Welfare; Animals; Female; Housing, Animal; Male; Pan troglodytes; Risk Assessment; Wounds and Injuries/veterinary
	Abstract	Meeting the psychological needs of chimpanzees (Pan troglodytes) can be a challenge given their aggressiveness on the one hand and the complexity of their social lives on the other. It is unclear how to balance the need to provide opportunities for species-appropriate behavior against potential risks of injury chimpanzees may inflict on each other. This study evaluates the suggestion that simpler social environments protect chimpanzees from wounding. Over a two-year period all visible injuries to 46 adult males, 64 adult females, and 25 immature chimpanzees were recorded at the Yerkes Regional Primate Research Center. Approximately half of the subjects were mother-reared, and the rest were nursery-reared. Housing included compounds containing about 20 chimpanzees, interconnected indoor-outdoor runs for groups of up to 12 individuals, and smaller indoor-outdoor runs for pairs and trios. Annual wounding rates were calculated for serious wounds (extensive injuries and all those requiring veterinary intervention) as well as for minor wounds. Compound-housed chimpanzees incurred the highest level of minor wounding, but serious wounding levels were not affected by housing condition. Even with a period of dominance instability and elevated levels of wounding in one compound, compound chimpanzees were not injured more than those in smaller social groups over the long term. Nursery-reared females in moderate-sized groups were wounded more than mother-reared females. Also, nursery-reared males and females were wounded less often when paired with mother-reared companions. Overall, this study indicates that maintaining chimpanzees in pairs and trios would not be an effective means for reducing injuries. The management of wounding in chimpanzee colonies is influenced more by the sex and rearing composition of a colony.
	Address	Living Links Center, Yerkes Regional Primate Research Center, Emory University, Atlanta, Georgia 30322, USA. kateb@rmy.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	0275-2565	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:10902665			Approved	no
	Call Number	refbase @ user @			Serial	188
Permanent link to this record



	Author	Aureli, F.; de Waal, F.B.
	Title	Inhibition of social behavior in chimpanzees under high-density conditions			Type	Journal Article
	Year	1997	Publication	American journal of primatology	Abbreviated Journal	Am. J. Primatol.
	Volume	41	Issue	3	Pages	213-228
	Keywords	Aggression/psychology; Animals; Behavior, Animal; Crowding; Female; Male; Pan troglodytes/psychology; Population Density; *Social Environment; Stress, Psychological
	Abstract	This is the first study to investigate the short-term effects of high population density on captive chimpanzees (Pan troglodytes). Subjects of the study were 45 chimpanzees living in five different groups at the Yerkes Regional Primate Research Center. The groups were observed under two conditions: 1) when they had access to both the indoor and outdoor sections of their enclosures; 2) during cold days when they were locked into the indoor runs, which reduced the available space by more than half. Under the high-density condition, allogrooming and submissive greetings decreased, but juvenile play increased. Remarkably, the rate of various forms of agonistic behavior, such as aggression, bluff charge, bluff display, and hooting, occurred less frequently under the high-density condition. This general decrease in adult social activity, including agonistic behavior, can be interpreted as an inhibition strategy to reduce opportunities for conflict when interindividual distances are reduced. This strategy is probably effective only in the short run, however. Behavioral indicators of anxiety, such as rough scratching and yawning, showed elevated rates, suggesting increased social tension under the high-density condition.
	Address	Yerkes Regional Primate Research Center, Emory University, Atlanta, GA 30322, USA
	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:9057966			Approved	no
	Call Number	refbase @ user @			Serial	203
Permanent link to this record



	Author	de Waal, F.B.
	Title	Bonobo sex and society			Type	Journal Article
	Year	1995	Publication	Scientific American	Abbreviated Journal	Sci Am
	Volume	272	Issue	3	Pages	82-88
	Keywords	Animals; Evolution; Female; Hominidae; Humans; Male; Pan troglodytes/physiology; Sexual Behavior, Animal/physiology; Social Behavior
	Abstract
	Address	Yerkes Regional Primate Research Center, Atlanta
	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:7871411			Approved	no
	Call Number	refbase @ user @			Serial	206
Permanent link to this record



	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
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	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
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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	Dunbar, R.I.M.; McAdam, M.R.; O'connell, S.
	Title	Mental rehearsal in great apes (Pan troglodytes and Pongo pygmaeus) and children			Type	Journal Article
	Year	2005	Publication	Behavioural Processes	Abbreviated Journal	Behav. Process.
	Volume	69	Issue	3	Pages	323-330
	Keywords	Algorithms; Animals; Child; Child, Preschool; Food; Frontal Lobe/anatomy & histology/physiology; Humans; Imagination; Pan troglodytes; Pongo pygmaeus; Problem Solving/physiology; Psychomotor Performance/physiology; Reward
	Abstract	The ability to rehearse possible future courses of action in the mind is an important feature of advanced social cognition in humans, and the “social brain” hypothesis implies that it might also be a feature of primate social cognition. We tested two chimpanzees, six orangutans and 63 children aged 3-7 years on a set of four puzzle boxes, half of which were presented with an opportunity to observe the box before being allowed to open it (“prior view”), the others being given without an opportunity to examine the boxes before handling them (“no prior view”). When learning effects are partialled out, puzzle boxes in the “prior view” condition were opened significantly faster than boxes given in the “no prior view” condition by the children, but not by either of the great apes. The three species differ significantly in the speed with which they opened boxes in the “no prior view” condition. The three species' performance on this task was a function of relative frontal lobe volume, suggesting that it may be possible to identify quantitative neuropsychological differences between species.
	Address	Evolutionary Psychology Research Group, School of Biological Sciences, University of Liverpool, Biosciences Building, Crown Street, Liverpool L69 7ZB, UK. rimd@liv.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	0376-6357	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15896530			Approved	no
	Call Number				Serial	2097
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	Author	Mulcahy, N.J.; Call, J.
	Title	How great apes perform on a modified trap-tube task			Type	Journal Article
	Year	2006	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	9	Issue	3	Pages	193-199
	Keywords	Animals; Discrimination Learning; Female; Gorilla gorilla; Hominidae/psychology; Intelligence; Male; Motor Skills; Pan paniscus; Pan troglodytes; Pongo pygmaeus; *Problem Solving; Species Specificity
	Abstract	To date, neither primates nor birds have shown clear evidence of causal knowledge when attempting to solve the trap tube task. One factor that may have contributed to mask the knowledge that subjects may have about the task is that subjects were only allowed to push the reward away from them, which is a particularly difficult action for primates in certain problem solving situations. We presented five orangutans (Pongo pygmaeus), two chimpanzees (Pan troglodytes), two bonobos (Pan paniscus), and one gorilla (Gorilla gorilla) with a modified trap tube that allowed subjects to push or rake the reward with the tool. In two additional follow-up tests, we inverted the tube 180 degrees rendering the trap nonfunctional and also presented subjects with the original task in which they were required to push the reward out of the tube. Results showed that all but one of the subjects preferred to rake the reward. Two orangutans and one chimpanzee (all of whom preferred to rake the reward), consistently avoided the trap only when it was functional but failed the original task. These findings suggest that some great apes may have some causal knowledge about the trap-tube task. Their success, however, depended on whether they were allowed to choose certain tool-using actions.
	Address	Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103, Leipzig, Germany. mulcahy@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	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16612632			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2469
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