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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	Plotnik, J.; Nelson, P.A.; de Waal, F.B.M.
	Title	Visual field information in the face perception of chimpanzees (Pan troglodytes)			Type	Journal Article
	Year	2003	Publication	Annals of the New York Academy of Sciences	Abbreviated Journal	Ann N Y Acad Sci
	Volume	1000	Issue		Pages	94-98
	Keywords	Animals; Facial Expression; Pan troglodytes; Recognition (Psychology); Visual Fields/physiology; Visual Perception/*physiology
	Abstract	Evidence for a visual field advantage (VFA) in the face perception of chimpanzees was investigated using a modification of a free-vision task. Four of six chimpanzee subjects previously trained on a computer joystick match-to-sample paradigm were able to distinguish between images of neutral face chimeras consisting of two left sides (LL) or right sides (RR) of the face. While an individual's ability to make this distinction would be unlikely to determine their suitability for the VFA tests, it was important to establish that distinctive information was available in test images. Data were then recorded on their choice of the LL vs. RR chimera as a match to the true, neutral image; a bias for one of these options would indicate an hemispatial visual field advantage. Results suggest that chimpanzees, unlike humans, do not exhibit a left visual field advantage. These results have important implications for studies on laterality and asymmetry in facial signals and their perception in primates.
	Address	Department of Animal Science, Cornell University, Ithaca, New York 14853, USA. jmp63@cornell.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	0077-8923	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:14766624			Approved	no
	Call Number	refbase @ user @			Serial	175
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	Author	Vokey, J.R.; Rendall, D.; Tangen, J.M.; Parr, L.A.; de Waal, F.B.M.
	Title	Visual kin recognition and family resemblance in 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	194-199
	Keywords	Animals; Female; Male; Pan troglodytes; Random Allocation; Recognition (Psychology); Visual Perception
	Abstract	The male-offspring biased visual kin recognition in chimpanzees (Pan troglodytes) reported by L. A. Parr and F. B. M. de Waal (1999) was replicated with human (Homo sapiens) participants and a principal components analysis (PCA) of pixel maps of the chimpanzee face photos. With the same original materials and methods, both humans and the PCA produced the same asymmetry in kin recognition as found with the chimpanzees. The PCA suggested that the asymmetry was a function of differences in the distribution of global characteristics associated with the framing of the faces in the son and daughter test sets. Eliminating potential framing biases, either by cropping the photos tightly to the faces or by rebalancing the recognition foils, eliminated the asymmetry but not human participants' ability to recognize chimpanzee kin.
	Address	Department of Psychology and Neuroscience, University of Lethbridge, Lethbridge, Alberta, Canada. vokey@uleth.ca
	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:15250806			Approved	no
	Call Number	refbase @ user @			Serial	171
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	Author	Bonnie, K.E.; de Waal, F.B.M.
	Title	Affiliation promotes the transmission of a social custom: handclasp grooming among captive chimpanzees			Type	Journal Article
	Year	2006	Publication	Primates	Abbreviated Journal	Primates
	Volume	47	Issue	1	Pages	27-34
	Keywords	Animals; Animals, Zoo/physiology; Behavior, Animal/physiology; Cultural Evolution; Grooming/physiology; Observation; Pan troglodytes/physiology; Social Behavior
	Abstract	Handclasp grooming is a unique social custom, known to occur regularly among some, but not all populations of chimpanzees (Pan troglodytes). As with other cultural behaviors, it is assumed that this distinctive grooming posture is learned socially by one individual from another. However, statistical comparisons among factors thought to influence how a behavior spreads within a group have never, to our knowledge, been conducted. In the present study, the origination and spread of handclasp grooming in a group of captive chimpanzees was followed throughout more than 1,500 h of observation over a period of 12 years. We report on the frequency, bout duration, and number and demography of performers throughout the study period, and compare these findings to those reported for wild populations. We predicted that dyads with strong affiliative ties, measured by time spent in proximity to and grooming one another, were likely to develop a handclasp grooming partnership during the study period. A quadratic assignment procedure was used to compare correlations among observed frequencies of grooming and proximity with handclasp grooming in all possible dyads within the group. As predicted, the formation of new handclasp grooming dyads was positively correlated with the rate of overall grooming and proximity within a dyad. In addition, in nearly all dyads formed, at least one individual had been previously observed to handclasp groom. We concluded that affiliation and individual experience determines the transmission of handclasp grooming among captive chimpanzees.
	Address	Department of Psychology, Emory University, and Living Links Center, Yerkes National Primate Research Center, 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	0032-8332	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16142425			Approved	no
	Call Number	refbase @ user @			Serial	161
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	Author	de Waal, F.B.M.
	Title	How animals do business			Type	Journal Article
	Year	2005	Publication	Scientific American	Abbreviated Journal	Sci Am
	Volume	292	Issue	4	Pages	54-61
	Keywords	Animals; Attitude; Behavior, Animal; Cebus; Cooperative Behavior; Economics; Emotions; Fishes; Food; Humans; Pan troglodytes; Papio; Social Behavior
	Abstract
	Address	Emory University, 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	0036-8733	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15915815			Approved	no
	Call Number	refbase @ user @			Serial	166
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	Author	de Waal, F.B.M.; Dindo, M.; Freeman, C.A.; Hall, M.J.
	Title	The monkey in the mirror: hardly a stranger			Type	Journal Article
	Year	2005	Publication	Proceedings of the National Academy of Sciences of the United States of America	Abbreviated Journal	Proc. Natl. Acad. Sci. U.S.A.
	Volume	102	Issue	32	Pages	11140-11147
	Keywords	Analysis of Variance; Animals; Cebus/physiology; Discrimination (Psychology); Empathy; Female; Male; Observation; Recognition (Psychology); Self Concept; Sex Factors
	Abstract	It is widely assumed that monkeys see a stranger in the mirror, whereas apes and humans recognize themselves. In this study, we question the former assumption by using a detailed comparison of how monkeys respond to mirrors versus live individuals. Eight adult female and six adult male brown capuchin monkeys (Cebus apella) were exposed twice to three conditions: (i) a familiar same-sex partner, (ii) an unfamiliar same-sex partner, and (iii) a mirror. Females showed more eye contact and friendly behavior and fewer signs of anxiety in front of a mirror than they did when exposed to an unfamiliar partner. Males showed greater ambiguity, but they too reacted differently to mirrors and strangers. Discrimination between conditions was immediate, and blind coders were able to tell the difference between monkeys under the three conditions. Capuchins thus seem to recognize their reflection in the mirror as special, and they may not confuse it with an actual conspecific. Possibly, they reach a level of self-other distinction intermediate between seeing their mirror image as other and recognizing it as self.
	Address	Living Links Center, Emory University, Atlanta, GA 30322, USA. dewaal@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	0027-8424	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16055557			Approved	no
	Call Number	refbase @ user @			Serial	164
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	Author	Brosnan, S.F.; de Waal, F.B.M.
	Title	A concept of value during experimental exchange in brown capuchin monkeys, Cebus apella			Type	Journal Article
	Year	2004	Publication	Folia primatologica; international journal of primatology	Abbreviated Journal	Folia Primatol (Basel)
	Volume	75	Issue	5	Pages	317-330
	Keywords	Animals; Cebus/psychology; Choice Behavior; Female; Food Preferences; Learning; Male; Sex Factors; Statistics, Nonparametric; Token Economy; Video Recording
	Abstract	We evaluated the response of brown capuchin monkeys to two differentially valued tokens in an experimental exchange situation akin to a simple barter. Monkeys were given a series of three tests to evaluate their ability to associate tokens with food, then their responses were examined in a barter situation in which tokens were either limited or unlimited. Capuchins did not perform barter in the typical sense, returning the tokens which were associated with the reward. However, females, but not males, showed a different response, preferring the higher-value token. This may indicate that they learned to prefer one token over the other rather than to associate the tokens with their specific rewards. This sex difference parallels previous findings of greater reciprocity in female brown capuchins than in males.
	Address	Living Links Center, Emory University, Yerkes National Primate Research Center, 954 N. Gatewood Drive, Atlanta, GA 30329, 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	0015-5713	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15486443			Approved	no
	Call Number	refbase @ user @			Serial	170
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	Author	Brosnan, S.F.; de Waal, F.B.M.
	Title	Socially learned preferences for differentially rewarded tokens in the brown capuchin monkey (Cebus apella)			Type	Journal Article
	Year	2004	Publication	Journal of comparative psychology (Washington, D.C. : 1983)	Abbreviated Journal	J Comp Psychol
	Volume	118	Issue	2	Pages	133-139
	Keywords	Animals; Behavior, Animal; Cebus; Choice Behavior; Female; Learning; Male; Reward; Social Behavior
	Abstract	Social learning is assumed to underlie traditions, yet evidence indicating social learning in capuchin monkeys (Cebus apella), which exhibit traditions, is sparse. The authors tested capuchins for their ability to learn the value of novel tokens using a previously familiar token-exchange economy. Capuchins change their preferences in favor of a token worth a high-value food reward after watching a conspecific model exchange 2 differentially rewarded tokens, yet they fail to develop a similar preference after watching tokens paired with foods in the absence of a conspecific model. They also fail to learn that the value of familiar tokens has changed. Information about token value is available in all situations, but capuchins seem to pay more attention in a social situation involving novel tokens.
	Address	Living Links Center, Yerkes National Primate Research Center, and Graduate Division of Biological and Biomedical Sciences, Emory University, Atlanta, GA 30329, 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	0735-7036	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:15250800			Approved	no
	Call Number	refbase @ user @			Serial	173
Permanent link to this record



	Author	Brosnan, S.F.; Freeman, C.; De Waal, F.B.M.
	Title	Partner's behavior, not reward distribution, determines success in an unequal cooperative task in capuchin monkeys			Type	Journal Article
	Year	2006	Publication	American journal of primatology	Abbreviated Journal	Am. J. Primatol.
	Volume	68	Issue	7	Pages	713-724
	Keywords	Animals; Behavior, Animal/physiology; Cebus/physiology; Cooperative Behavior; Female; Food Preferences/physiology; Male; Reward
	Abstract	It was recently demonstrated that capuchin monkeys notice and respond to distributional inequity, a trait that has been proposed to support the evolution of cooperation in the human species. However, it is unknown how capuchins react to inequitable rewards in an unrestricted cooperative paradigm in which they may freely choose both whether to participate and, within the bounds of their partner's behavior, which reward they will receive for their participation. We tested capuchin monkeys with such a design, using a cooperative barpull, which has been used with great success in the past. Contrary to our expectations, the equity of the reward distribution did not affect success or pulling behavior. However, the behavior of the partner in an unequal situation did affect overall success rates: pairs that had a tendency to alternate which individual received the higher-value food in unequal reward situations were more than twice as successful in obtaining rewards than pairs in which one individual dominated the higher-value food. This ability to equitably distribute rewards in inherently biased cooperative situations has profound implications for activities such as group hunts, in which multiple individuals work together for a single, monopolizable reward.
	Address	Living Links Center, Yerkes National Primate Research Center, Emory University, Atlanta, Georgia, 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	0275-2565	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16786518			Approved	no
	Call Number	refbase @ user @			Serial	160
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	Author	Plotnik, J.M.; de Waal, F.B.M.; Reiss, D.
	Title	Self-recognition in an Asian elephant			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	45	Pages	17053-17057
	Keywords	Animals; Asia; Behavior, Animal; Cognition; Elephants/psychology; Female; Photic Stimulation
	Abstract	Considered an indicator of self-awareness, mirror self-recognition (MSR) has long seemed limited to humans and apes. In both phylogeny and human ontogeny, MSR is thought to correlate with higher forms of empathy and altruistic behavior. Apart from humans and apes, dolphins and elephants are also known for such capacities. After the recent discovery of MSR in dolphins (Tursiops truncatus), elephants thus were the next logical candidate species. We exposed three Asian elephants (Elephas maximus) to a large mirror to investigate their responses. Animals that possess MSR typically progress through four stages of behavior when facing a mirror: (i) social responses, (ii) physical inspection (e.g., looking behind the mirror), (iii) repetitive mirror-testing behavior, and (iv) realization of seeing themselves. Visible marks and invisible sham-marks were applied to the elephants' heads to test whether they would pass the litmus “mark test” for MSR in which an individual spontaneously uses a mirror to touch an otherwise imperceptible mark on its own body. Here, we report a successful MSR elephant study and report striking parallels in the progression of responses to mirrors among apes, dolphins, and elephants. These parallels suggest convergent cognitive evolution most likely related to complex sociality and cooperation.
	Address	Living Links, Yerkes National Primate Research Center, and Department of Psychology, Emory University, 532 North Kligo Circle, 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	0027-8424	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:17075063			Approved	no
	Call Number	refbase @ user @			Serial	408
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