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	Author	Whiten, A.; Horner, V.; de Waal, F.B.M.
	Title	Conformity to cultural norms of tool use in chimpanzees			Type	Journal Article
	Year	2005	Publication	Nature	Abbreviated Journal	Nature
	Volume	437	Issue	7059	Pages	737-740
	Keywords	Aging/physiology; Animals; Culture; Feeding Behavior/physiology; Female; Pan troglodytes/physiology/psychology; *Social Conformity; Technology; Time Factors
	Abstract	Rich circumstantial evidence suggests that the extensive behavioural diversity recorded in wild great apes reflects a complexity of cultural variation unmatched by species other than our own. However, the capacity for cultural transmission assumed by this interpretation has remained difficult to test rigorously in the field, where the scope for controlled experimentation is limited. Here we show that experimentally introduced technologies will spread within different ape communities. Unobserved by group mates, we first trained a high-ranking female from each of two groups of captive chimpanzees to adopt one of two different tool-use techniques for obtaining food from the same 'Pan-pipe' apparatus, then re-introduced each female to her respective group. All but two of 32 chimpanzees mastered the new technique under the influence of their local expert, whereas none did so in a third population lacking an expert. Most chimpanzees adopted the method seeded in their group, and these traditions continued to diverge over time. A subset of chimpanzees that discovered the alternative method nevertheless went on to match the predominant approach of their companions, showing a conformity bias that is regarded as a hallmark of human culture.
	Address	Centre for Social Learning and Cognitive Evolution, School of Psychology, University of St Andrews, St Andrews, Fife, KY16 9JP, UK. 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	1476-4687	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16113685			Approved	no
	Call Number	refbase @ user @			Serial	163
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	Author	de Waal, F.B.M.
	Title	A century of getting to know the chimpanzee			Type	Journal Article
	Year	2005	Publication	Nature	Abbreviated Journal	Nature
	Volume	437	Issue	7055	Pages	56-59
	Keywords	Aggression; Animals; Behavior, Animal/physiology; Competitive Behavior; Cooperative Behavior; Female; Humans; Male; Pan troglodytes/genetics/physiology/psychology; Sexual Behavior, Animal; *Social Behavior
	Abstract	A century of research on chimpanzees, both in their natural habitat and in captivity, has brought these apes socially, emotionally and mentally much closer to us. Parallels and homologues between chimpanzee and human behaviour range from tool-technology and cultural learning to power politics and intercommunity warfare. Few behavioural domains have remained untouched by this increased knowledge, which has dramatically challenged the way we view ourselves. The sequencing of the chimpanzee genome will no doubt bring more surprises and insights. Humans do occupy a special place among the primates, but this place increasingly has to be defined against a backdrop of substantial similarity.
	Address	Living Links, Yerkes National Primate Research Center, Emory University, 954 North Gatewood Road, Atlanta, Georgia 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	1476-4687	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16136128			Approved	no
	Call Number	refbase @ user @			Serial	162
Permanent link to this record



	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	Flack, J.C.; Girvan, M.; de Waal, F.B.M.; Krakauer, D.C.
	Title	Policing stabilizes construction of social niches in primates			Type	Journal Article
	Year	2006	Publication	Nature	Abbreviated Journal	Nature
	Volume	439	Issue	7075	Pages	426-429
	Keywords	Animals; Conflict (Psychology); Female; Macaca nemestrina/physiology/psychology; Male; Models, Biological; *Social Behavior
	Abstract	All organisms interact with their environment, and in doing so shape it, modifying resource availability. Termed niche construction, this process has been studied primarily at the ecological level with an emphasis on the consequences of construction across generations. We focus on the behavioural process of construction within a single generation, identifying the role a robustness mechanism--conflict management--has in promoting interactions that build social resource networks or social niches. Using 'knockout' experiments on a large, captive group of pigtailed macaques (Macaca nemestrina), we show that a policing function, performed infrequently by a small subset of individuals, significantly contributes to maintaining stable resource networks in the face of chronic perturbations that arise through conflict. When policing is absent, social niches destabilize, with group members building smaller, less diverse, and less integrated grooming, play, proximity and contact-sitting networks. Instability is quantified in terms of reduced mean degree, increased clustering, reduced reach, and increased assortativity. Policing not only controls conflict, we find it significantly influences the structure of networks that constitute essential social resources in gregarious primate societies. The structure of such networks plays a critical role in infant survivorship, emergence and spread of cooperative behaviour, social learning and cultural traditions.
	Address	Santa Fe Institute, Santa Fe, New Mexico 87501, 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	1476-4687	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16437106			Approved	no
	Call Number	refbase @ user @			Serial	298
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	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	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.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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	Author	Dindo, M.; De Waal, F.B.M.
	Title	Partner effects on food consumption in brown capuchin monkeys			Type	Journal Article
	Year	2007	Publication	American journal of primatology	Abbreviated Journal	Am. J. Primatol.
	Volume	69	Issue	4	Pages	448-456
	Keywords
	Abstract	It has been claimed that capuchin monkeys (Cebus apella) show inequity aversion in relation to food rewards for a simple exchange task. However, other factors may affect the willingness of a monkey to consume foods of high or low value in the presence of a conspecific. In this study, pairs of monkeys were presented with unequally valued foods, but without any task-performance: they simply received the food under four experimental conditions. By looking at the rate of collection and consumption of low-valued cucumber slices we expected to see variation dependent on whether the partner either had 1) cucumber (equity), 2) grape (inequity), 3) inaccessible cucumber or 4) inaccessible grape. Testing 12 adult capuchin monkeys, our findings differed from those of other authors in that the monkeys failed to show negative reactions to inequity, but rather responded with scramble competition (i.e., fast food collection) in the presence of a conspecific without access to food. They also showed facilitated consumption in the presence of a conspecific consuming high-valued food. Possibly, (in)equity plays a different role if food serves as a reward for a task rather than if it is simply made available for consumption. Am. J. Primatol. 69:1-9, 2007. (c) 2006 Wiley-Liss, Inc.
	Address	Living Links, Yerkes National Primate Research Center, Emory University, Atlanta, Georgia
	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:17146793			Approved	no
	Call Number	refbase @ user @			Serial	158
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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	de Waal, F.B.
	Title	The integration of dominance and social bonding in primates			Type	Journal Article
	Year	1986	Publication	The Quarterly review of biology	Abbreviated Journal	Q Rev Biol
	Volume	61	Issue	4	Pages	459-479
	Keywords	Animals; Female; Humans; Male; Object Attachment; Primates; *Social Dominance
	Abstract	Social dominance is usually viewed from the perspective of intragroup competition over access to limited resources. The present paper, while not denying the importance of such competition, discusses the dominance concept among monkeys and apes in the context of affiliative bonding, social tolerance, and the reconciliation of aggressive conflicts. Two basic proximate mechanisms are supposed to provide a link between dominance and interindividual affiliation, namely, formalization of the dominance relationship (i.e., unequivocal communication of status), and conditional reassurance (i.e., the linkage of friendly coexistence to formalization of the relationship). Ritualized submission is imposed upon losers of dominance struggles by winners; losers are offered a “choice” between continued hostility or a tolerant relationship with a clearly signalled difference in status. If these two social mechanisms are lacking, aggression is bound to have dispersive effects. In their presence, aggression becomes a well-integrated, even constructive component of social life. In some higher primates this process of integration has reached the stage where status differences are strongly attenuated. In these species, sharing and trading can take the place of overt competition. The views underlying this “reconciled hierarchy” model are only partly new, as is evident from a review of the ethological literature. Many points are illustrated with data on a large semi-captive colony of chimpanzees (Pan troglodytes), particularly data related to striving for status, reconciliation behavior, and general association patterns. These observations demonstrate that relationships among adult male chimpanzees cannot be described in terms of a dichotomy between affiliative and antagonistic tendencies. Male bonding in this species has not been achieved by an elimination of aggression, but by a set of powerful buffering mechanisms that mitigate its effects. Although female chimpanzees do exhibit a potential for bonding under noncompetitive conditions, they appear to lack the buffering mechanisms of the males.
	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	0033-5770	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:3543991			Approved	no
	Call Number	refbase @ user @			Serial	210
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