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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	Hewitt, S.E.; Macdonald, D.W.; Dugdale, H.L.
	Title	Context-dependent linear dominance hierarchies in social groups of European badgers, Meles meles			Type	Journal Article
	Year	2009	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	77	Issue	1	Pages	161-169
	Keywords	David's score; directed aggression; dominance rank; European badger; grooming behaviour; linear dominance hierarchy; Meles meles; reproductive success; relatedness; social status
	Abstract	A social hierarchy is generally assumed to exist in those mammalian societies in which the costs and benefits of group living are distributed unevenly among group members. We analysed infrared closed-circuit television footage, collected over 3 years in Wytham Woods, Oxfordshire, to test whether social groups of European badgers have dominance hierarchies. Analysis of directed aggression between dyads revealed linear dominance hierarchies in three social-group-years, but patterns within social groups were not consistent across years. Dominance hierarchies were significantly steeper than random in five out of six social-group-years. In those social-group-years where a linear hierarchy was determined, there was an effect of sex on dominance rank, with females gaining significantly higher rank than males in two social-group-years. Overall, rank was not related to age, nor did it appear to affect the likelihood of an individual being wounded, or an individual's breeding status. The latter resulted from nonorthogonality between sex and breeding status, as there were only two breeding males. Overall, hierarchies were primarily dominated by breeding females, and may occur when breeding competition arises. Relatedness, unreciprocated allogrooming and sequential allomarking were not consistently related to levels of directed aggression across social-group-years. We suggest that dominance structures within European badger groups may be context dependent, with future study required to complete our understanding of where, and when, they arise.
	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	4695
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	Author	Seyfarth, R.M.
	Title	A model of social grooming among adult female monkeys			Type	Journal Article
	Year	1977	Publication	Journal of Theoretical Biology	Abbreviated Journal	J. Theor. Biol.
	Volume	65	Issue	4	Pages	671-698
	Keywords	Animals; Behavior, Animal; Female; Grooming; Haplorhini/physiology; *Models, Biological; Reproduction; Social Dominance; Time Factors
	Abstract	Grooming networks among adult female monkeys exhibit two similar features across a number of different species. High-ranking animals receive more grooming than others, and the majority of grooming occurs between females of adjacent rank. A theoretical model which duplicates these features is presented, and the properties of the model are used to explain the possible causation and function of female grooming behaviour. The model illustrates how relatively simple principles governing the behaviour of individuals may be used to explain more complex aspects of the social structure of non-human primate groups.
	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	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:406485			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5259
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	Author	de Waal, F.B.; Johanowicz, D.L.
	Title	Modification of reconciliation behavior through social experience: an experiment with two macaque species			Type	Journal Article
	Year	1993	Publication	Child development	Abbreviated Journal	Child Dev
	Volume	64	Issue	3	Pages	897-908
	Keywords	Aggression; Animals; Behavior, Animal; Female; Grooming; Macaca; Macaca mulatta; Male; Play and Playthings; Socialization; Vocalization, Animal
	Abstract	Reconciliation, defined as a friendly reunion between former opponents shortly after an aggressive encounter, is common in the stumptail macaque (Macaca arctoides) but rare in the rhesus macaque (M. mulatta). Juveniles of the two species were cohoused for 5 months, after which they were observed with conspecifics only. Control rhesus monkeys, matched in age and sex to the experimental subjects, went through the same procedure without exposure to the other species. A threefold increase in the proportion of reconciled fights was measured in the rhesus subjects. The difference emerged gradually during cohousing with the tutor species and was sustained following removal of this species. Other behavior, such as grooming and aggression, decreased over time. It is suggested that the social attitude of the subjects was affected through contact with a species characterized by a more relaxed dominance style.
	Address	Wisconsin Regional Primate Research Center, University of Wisconsin-Madison
	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	0009-3920	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:8339702			Approved	no
	Call Number	refbase @ user @			Serial	209
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	Author	Judge, P.G.; de Waal, F.B.
	Title	Intergroup grooming relations between alpha females in a population of free-ranging rhesus macaques			Type	Journal Article
	Year	1994	Publication	Folia primatologica; international journal of primatology	Abbreviated Journal	Folia Primatol (Basel)
	Volume	63	Issue	2	Pages	63-70
	Keywords	Animals; Animals, Wild; Female; Grooming; Macaca mulatta/psychology; Male; Population Density; *Social Behavior
	Abstract	Intergroup affiliation among female rhesus macaques, Macaca mulatta, was examined in the captive free-ranging colony of Morgan Island, S.C., USA. The provisioned colony has many social groups (35) and is maintained at a relatively high population density (21 animals/ha) with a relatively low adult male to female ratio (1:8.8). Focal and ad libitum samples were collected on 32 adults (3 males and 29 females) from two groups. Although infrequent, grooming was observed between adult females from different groups, and alpha females were the main participants in these interactions. Colony records indicated that none of the intergroup grooms was between females formerly from a common group. Relations between familiar neighboring groups may be maintained by a combination of both affiliative and aggressive behavior.
	Address	Yerkes Regional Primate Research Center, Lawrenceville, Ga 30243
	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:7813977			Approved	no
	Call Number	refbase @ user @			Serial	208
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	Author	Kimura, R.
	Title	Mutual grooming and preferred associate relationships in a band of free-ranging horses			Type	Journal Article
	Year	1998	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	59	Issue	4	Pages	265-276
	Keywords	Horses; Grooming; Dominance entropy
	Abstract	Preferred associate (nearest neighbour) and mutual grooming relationships among mares, in an isolated family band of free-ranging horses (Equus caballus), were studied, and the structural and functional differences between these two relationships were examined. The frequent partners accompanying the mare were not the same in both these relationships (p<0.05) and mares changed their partners during the study period between 1988-1990. Individual horses of similar rank tended to remain closer together in both winter and summer. Aggressive-submissive behaviour was so infrequent during spring that rank determinations could not be made; however, in fall, although rank could be determined, rank was not correlated with nearest neighbour. Three subgroups, based on preferred associate relationships in summer, fall and winter, directly reflected the age and social rank of the mares in the group. Individual horses of higher rank tended to have many partners in winter, while individuals of lower rank had fewer. There was no significant correlation between the frequency of mutual grooming and individual rank. The mutual grooming relationship was strongly influenced by seasonal changes as the relative amount of grazing time per day increased. Thus, the frequency of mutual grooming was lowest in winter and highest in summer. The mutual grooming relationship was based on the bonds between individual horses, which were little influenced by social rank. Lower ranking individuals tended to have a greater variety of grooming partners in summer.
	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		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number				Serial	2022
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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.; Luttrell, L.M.
	Title	The similarity principle underlying social bonding among female rhesus monkeys			Type	Journal Article
	Year	1986	Publication	Folia primatologica; international journal of primatology	Abbreviated Journal	Folia Primatol (Basel)
	Volume	46	Issue	4	Pages	215-234
	Keywords	Aggression; Animals; Dominance-Subordination; Female; Grooming; Group Processes; Macaca/physiology; Macaca mulatta/physiology; Male; Object Attachment
	Abstract	Twenty adult female rhesus monkeys (Macaca mulatta) were observed over a three-year period. They lived in a mixed captive group with kinship relations known for three generations. The study's aim was to test Seyfarth's [J. theor. Biol. 65: 671-698, 1977] model of rank-related grooming and to investigate two other possible determinants of social bonding, i.e. relative age and the group's stratification into two social classes. Data on affiliation, coalitions, and social competition were collected by means of both focal observation and instantaneous time sampling. Whereas certain elements of the existing model were confirmed, its explanatory principles were not. Social competition did not result in more contact among close-ranking females (the opposite effect was found), and the relation between affiliative behavior and coalitions was more complex than predicted. Based on multivariate analyses and a comparison of theoretical models, we propose a simpler, more encompassing principle underlying interfemale attraction. According to this 'similarity principle', rhesus females establish bonds with females whom they most resemble. The similarity may concern genetical and social background, age, hierarchical position and social class. Effects of these four factors were independently demonstrated. The most successful model assumed that similarity factors influence female bonding in a cumulative fashion.
	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	0015-5713	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:3557225			Approved	no
	Call Number	refbase @ user @			Serial	211
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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
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	Author	de Waal, F.B.; Seres, M.
	Title	Propagation of handclasp grooming among captive chimpanzees			Type	Journal Article
	Year	1997	Publication	American journal of primatology	Abbreviated Journal	Am. J. Primatol.
	Volume	43	Issue	4	Pages	339-346
	Keywords	Animals; Family Relations; Female; Grooming; Learning; Male; Pan troglodytes/psychology; *Social Behavior
	Abstract	A grooming posture previously reported for two wild chimpanzee (Pan troglodytes) communities developed spontaneously in a captive group of the same species. This offered a unique opportunity to follow the propagation of a new social custom. The posture consists of two partners grasping hands--either both right hands or both left hands--and raising the arms in an A-frame above their heads while mutually grooming with their free hands. The propagation of this pattern was followed over a 5 year period. In the beginning, handclasps were always initiated by the same adult female. This female initiated the posture mainly with her adult female kin. In subsequent years, these relatives became frequent participants in the posture with each other as well as with nonrelatives. Over the years the posture increased in frequency and duration and spread to the majority of adults and also to a few adolescents and older juveniles. The pattern persisted after removal of the apparent originator.
	Address	Yerkes Primate Center, Emory University, Atlanta, GA 30329, 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	0275-2565	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:9403098			Approved	no
	Call Number	refbase @ user @			Serial	202
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