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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
Permanent link to this record



	Author	Sinha, A.
	Title	Knowledge acquired and decisions made: triadic interactions during allogrooming in wild bonnet macaques, Macaca radiata			Type	Journal Article
	Year	1998	Publication	Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences	Abbreviated Journal	Philos Trans R Soc Lond B Biol Sci
	Volume	353	Issue	1368	Pages	619-631
	Keywords	Aggression; Animals; Cognition; Computer Simulation; Decision Making; Evolution; Female; Grooming; Logistic Models; Macaca radiata/psychology; Social Behavior; Social Dominance
	Abstract	The pressures of developing and maintaining intricate social relationships may have led to the evolution of enhanced cognitive abilities in many nonhuman primates. Knowledge of the dominance ranks and social relationships of other individuals, in particular, is important in evaluating one's position in the rank hierarchy and affiliative networks. Triadic interactions offer an excellent opportunity to examine whether decisions are taken by individuals on the basis of such knowledge. Allogrooming supplants among wild female bonnet macaques (macaca radiata) usually involved the subordinate female of a grooming dyad retreating at the approach of a female dominant to both members of the dyad. In a few exceptional cases, however, the dominant member of the dyad retreated; simple non-cognitive hypotheses involving dyadic rank differences and agonistic relationships failed to explain this phenomenon. Instead, retreat by the dominant individual was positively correlated with the social attractiveness of her subordinate companion (as measured by the duration of grooming received by the latter from other females in the troop). This suggests that not only does an individual evaluate relationships among other females, but does so on the basis of the amount of grooming received by them. Similarly, the frequency of approaches received by any female was correlated with her social attractiveness when she was the dominant member of the dyad, but not when she was the subordinate. This indicated that approaching females might be aware of the relative dominance ranks of the two allogrooming individuals. In logistic regression analyses, the probability of any individual retreating was found to be influenced more by her knowledge of her rank difference with both the other interactants, rather than by their absolute ranks. Moreover, information about social attractiveness appeared to be used in terms of correlated dominance ranks. The nature of knowledge acquired by bonnet macaque females may thus be egotistical in that other individuals are evaluated relative to oneself, integrative in that information about all other interactants is used simultaneously, and hierarchical in the ability to preferentially use certain categories of knowledge for the storage of related information from other domains.
	Address	National Centre for Biological Sciences, TIFR Centre, Bangalore, India
	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	0962-8436	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:9602536			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4362
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	Author	Aureli, F.; Preston, S.D.; de Waal, F.B.
	Title	Heart rate responses to social interactions in free-moving rhesus macaques (Macaca mulatta): a pilot study			Type	Journal Article
	Year	1999	Publication	Journal of comparative psychology (Washington, D.C. : 1983)	Abbreviated Journal	J Comp Psychol
	Volume	113	Issue	1	Pages	59-65
	Keywords	Animals; Behavior, Animal/physiology; Female; Grooming/physiology; Heart Rate/physiology; Macaca mulatta/physiology; Male; Movement/physiology; Pilot Projects; Social Behavior
	Abstract	Heart rate telemetry was explored as a means to access animal emotion during social interactions under naturalistic conditions. Heart rates of 2 middle-ranking adult females living in a large group of rhesus macaques (Macaca mulatta) were recorded along with their behavior. Heart rate changes during 2 types of interactions were investigated, while controlling for the effects of posture and activity. The risk of aggression associated with the approach of a dominant individual was expected to provoke anxiety in the approachee. This prediction was supported by the heart rate increase after such an approach. No increase was found when the approacher was a kin or a subordinate individual. The tension-reduction function of allogrooming was also supported. Heart rate decelerated faster during the receipt of grooming than in matched control periods.
	Address	Yerkes Regional Primate Research Center, Emory University, Atlanta, Georgia 30322, USA. aureli@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	0735-7036	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:10098269			Approved	no
	Call Number	refbase @ user @			Serial	197
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	Author	de Waal, F.B.; Aureli, F.; Judge, P.G.
	Title	Coping with crowding			Type	Journal Article
	Year	2000	Publication	Scientific American	Abbreviated Journal	Sci Am
	Volume	282	Issue	5	Pages	76-81
	Keywords	Adaptation, Psychological; Animals; Behavior, Animal; Emotions; Female; Grooming; Homicide; Humans; Macaca mulatta; Male; Pan troglodytes; Population Density; Primates; Rodentia; Rural Population; Territoriality; Urban Population; Violence
	Abstract
	Address	Living Links Center, Yerkes Regional Primate Research Center, Atlanta, 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:11056991			Approved	no
	Call Number	refbase @ user @			Serial	184
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	Author	Whiten, A.; Boesch, C.
	Title	The cultures of chimpanzees			Type	Journal Article
	Year	2001	Publication	Scientific American	Abbreviated Journal	Sci Am
	Volume	284	Issue	1	Pages	60-67
	Keywords	Animals; Behavior, Animal; Culture; Feeding Behavior; Grooming; Hominidae; Humans; Pan troglodytes/physiology
	Abstract
	Address	University of St. Andrews
	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:11132425			Approved	no
	Call Number	refbase @ user @			Serial	740
Permanent link to this record



	Author	de Waal, F.B.M.
	Title	Darwin's legacy and the study of primate visual communication			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	7-31
	Keywords	Affect; Aggression/psychology; Animals; Culture; Evolution; Facial Expression; Gestures; Grooming; Humans; Laughter; Nonverbal Communication; Primates/physiology; Smiling; *Visual Perception
	Abstract	After Charles Darwin's The Expression of the Emotions in Man and Animals, published in 1872, we had to wait 60 years before the theme of animal expressions was picked up by another astute observer. In 1935, Nadezhda Ladygina-Kohts published a detailed comparison of the expressive behavior of a juvenile chimpanzee and of her own child. After Kohts, we had to wait until the 1960s for modern ethological analyses of primate facial and gestural communication. Again, the focus was on the chimpanzee, but ethograms on other primates appeared as well. Our understanding of the range of expressions in other primates is at present far more advanced than that in Darwin's time. A strong social component has been added: instead of focusing on the expressions per se, they are now often classified according to the social situations in which they typically occur. Initially, quantitative analyses were sequential (i.e., concerned with temporal associations between behavior patterns), and they avoided the language of emotions. I will discuss some of this early work, including my own on the communicative repertoire of the bonobo, a close relative of the chimpanzee (and ourselves). I will provide concrete examples to make the point that there is a much richer matrix of contexts possible than the common behavioral categories of aggression, sex, fear, play, and so on. Primate signaling is a form of negotiation, and previous classifications have ignored the specifics of what animals try to achieve with their exchanges. There is also increasing evidence for signal conventionalization in primates, especially the apes, in both captivity and the field. This process results in group-specific or “cultural” communication patterns.
	Address	Yerkes Primate Center, and Psychology Department, Emory University, 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	0077-8923	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:14766618			Approved	no
	Call Number	refbase @ user @			Serial	177
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	Author	Dunbar, R.
	Title	Evolution of the social brain			Type	Journal Article
	Year	2003	Publication	Science	Abbreviated Journal	Science
	Volume	302	Issue	5648	Pages	1160-1161
	Keywords	Animals; Animals, Wild; Cognition; Endorphins/physiology; Evolution; Female; Grooming; Hierarchy, Social; Language; Neocortex/anatomy & histology/physiology; Papio/physiology/psychology; Reproduction; *Social Behavior; Social Dominance; Social Support; Vocalization, Animal
	Abstract
	Address	School of Biological Sciences, University of Liverpool, 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	1095-9203	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:14615522			Approved	no
	Call Number	refbase @ user @			Serial	548
Permanent link to this record



	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	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
Permanent link to this record



	Author	VanDierendonck, M.C.
	Title	Social relationships in a group of horses without a mature stallion			Type	Manuscript
	Year	2006	Publication	The Importance of Social Relationships in Horses	Abbreviated Journal
	Volume	Chapter 4	Issue		Pages
	Keywords	Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
	Abstract	Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication	Universiteit Utrecht	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	Equine Behaviour @ team @			Serial	2369
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