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	Author	Wolter, R.; Krueger, K.
	Title	The analysis of social bonds in feral horses			Type	Conference Article
	Year	2015	Publication	Proceedings of the 3. International Equine Science Meeting	Abbreviated Journal	Proc. 3. Int. Equine. Sci. Mtg
	Volume		Issue		Pages
	Keywords	social bonds, grooming, nearest neighborhood, rank
	Abstract	In many social mammals, individuals preferentially affiliate with a small subset of available partners instead of distributing their social behaviors equally among all group members. The resulting social bonds have been investigated in several mammalian taxa, especially in primate societies, but also in other taxa such as birds, dolphins, rodents and ungulates. In feral horses, a great number of studies on social bonds can be found, but with a huge variety between methods for the analysis. There seems to be a lack of a clear and common definition of social bonds in horses and of comparable analyses. For example, there are irregularities between the studies regarding the research designs, the selection of recording methods and the interpretation of the measurements. Mutual grooming is used most often for the analysis of social relationships in many species. As mutual grooming is rare in horses, especially measurements of spatial proximity are commonly used for the analysis of social bonds in addition to other behavioral patterns. However, the combination of mutual grooming and nearest neighborhood analyses for the analysis of social bond is debatable, as in contrast to mutual grooming, which must occur deliberately by both grooming partners, the spatial distribution can be influenced by one partner alone, which may even force the other horse to keep a certain distance or to stay in close proximity. In this study, we investigated the comparability of mutual grooming and nearest neighborhood data for social bond analyses in feral horses. Therefore, we observed five groups of semi-wild living Przewalski’s horses and six groups of feral horses. We analysed the horses’ social ranks by applying an Average Dominance Index, we recorded the distances between the animals and observed the number of mutual grooming events as well as friendly approaches. Our results show that there was only a weak correlation between the frequency of staying in nearest neighborhood and mutual grooming in all observed horse groups. In contrast to this, the correlation between the number of friendly approaches and mutual grooming events was higher in most groups. Hierarchies did not affect social bonds, as mutual grooming was similarly induced by higher and lower ranking animals and the social rank did not affect the choice of the grooming partner. Similarly, likelihoods of staying in the neighborhood of particular animals were not affected by the animals’ social rank. The grooming frequencies differed between the different horse groups and between the individual horses living in the particular groups. They seem to be effected by individual predisposition. Altogether we suggest that the ratio of mutual grooming seems to be a better indicator for social bonds in feral horses than the frequency of staying in the nearest neighborhood. Mutual grooming occurs deliberately and is bidirectional, whereas nearest neighborhoods could be enforced and unidirectional. For the calculation of social bonds in horses, we consider it to be more reliable to combine the frequency of mutual grooming with the frequency of friendly approaches.
	Address
	Corporate Author	Wolter, R.			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	978-3-95625-000-2	Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5882
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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	Clutton-Brock, T.H.; Greenwood, P.J.; Powell, R.P.
	Title	Ranks and relationships in Highland ponies and Highland Cows			Type	Journal Article
	Year	1976	Publication	Zeitschrift für Tierpsychologie	Abbreviated Journal	Z. Tierpsychol.
	Volume	41	Issue	2	Pages	202-216
	Keywords	Agonistic Behavior; Animals; Cattle; Female; Grooming; Horses; Male; *Social Dominance; Spatial Behavior
	Abstract	Recent studies of primates have questioned the importance of dominance hierarchies in groups living under natural conditions. In a herd of Highland ponies and one of Highland cattle grazing under free-range conditions on the Isle of Rhum (Inner Hebrides) well defined hierarchies were present. The provision of food produced a marked increase in the frequency of agonistic interactions but had no effect on the rank systems of the two herds. While rank was clearly important in affecting the distribution of agonistic interactions, it was poorly related to behaviour in non-agonistic situations.
	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	0044-3573	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:961125			Approved	no
	Call Number	refbase @ user @			Serial	661
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	Author	Seyfarth, R.M.; Cheney, D.L.
	Title	Grooming, alliances and reciprocal altruism in vervet monkeys			Type	Journal Article
	Year	1984	Publication	Nature	Abbreviated Journal	Nature
	Volume	308	Issue	5959	Pages	541-543
	Keywords	Altruism; Animals; Cercopithecus/physiology; Cercopithecus aethiops/physiology; Grooming; *Social Behavior; Vocalization, Animal
	Abstract	Reciprocal altruism refers to the exchange of beneficial acts between individuals, in which the benefits to the recipient exceed the cost to the altruist. Theory predicts that cooperation among unrelated animals can occur whenever individuals encounter each other regularly and are capable of adjusting their cooperative behaviour according to experience. Although the potential for reciprocal altruism exists in many animal societies, most interactions occur between closely related individuals, and examples of reciprocity among non-kin are rare. The field experiments on vervet monkeys which we present here demonstrate that grooming between unrelated individuals increases the probability that they will subsequently attend to each others' solicitations for aid. Vervets appear to be more willing to aid unrelated individuals if those individuals have behaved affinitively toward them in the recent past. In contrast, recent grooming between close genetic relatives appears to have no effect on their willingness to respond to each other's solicitations for aid.
	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	0028-0836	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:6709060			Approved	no
	Call Number	refbase @ user @			Serial	704
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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	Sigurjónsdóttir, H.; Haraldsson, H.
	Title	Significance of Group Composition for the Welfare of Pastured Horses			Type	Journal Article
	Year	2018	Publication	Animals	Abbreviated Journal	Animals
	Volume	9	Issue	1	Pages	14
	Keywords	horse welfare; aggression; allogrooming; pastured horses; Icelandic horse
	Abstract	Simple Summary Because of their social nature, horses need to have plenty of opportunities to interact with others to establish bonds and learn from their elders. Comparison of social behaviour of 426 horses in 20 groups of Icelandic horses in pastures, showed that aggression was lowest where the group composition was like the natural system, i.e., with a stallion, mares and their young foals. In groups without a stallion, the presence of foals is also associated with low aggression. Stability of the group with respect to group composition is of great importance; the horses are less aggressive in the more stable groups. The highest aggression was found in groups of unfamiliar yearlings. The horses allogroomed more in groups with relatively more young horses, which suggests they are forming bonds. Later, they groom less but prefer certain individuals. Horse owners should all be aware of the importance of planning the composition of horse groups and to keep the membership as stable as possible in order to ensure good welfare. Abstract We explore how herd composition and management factors correlate with frequencies of social interactions in horse groups. Since the welfare of horses correlates with low aggression levels and social contact opportunities, information of this kind is important. The data are a collection of records of social interactions of 426 Icelandic horses in 20 groups of at least eight horses. The complexities and limitations of the data prohibit useful statistical modelling so the results are presented descriptively. Interesting and informative patterns emerge which can be of use both in management and in future studies. Of special interest are the low levels of agonistic behaviours in breeding groups where one stallion was present. The horses were less agonistic when in groups with young foals and where group membership was stable. Unfamiliar yearlings in peer groups were especially aggressive. Allogrooming was most frequent in groups with relatively more young horses and in unstable and small groups. Interestingly, the horses allogroomed more if they had few preferred allogrooming partners. The findings show that composition (age/sex) and stability of groups are of great importance with respect to aggression levels and opportunities for establishing bonds.
	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	Equine Behaviour @ team @			Serial	6585
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	Author	Wolter, R.; Stefanski, V.; Krueger, K.
	Title	Parameters for the Analysis of Social Bonds in Horses			Type	Journal Article
	Year	2018	Publication	Animals	Abbreviated Journal	Animals
	Volume	8	Issue	11	Pages	191
	Keywords	feral horses; mutual grooming; social bonds; social bond analysis; spatial proximity
	Abstract	Social bond analysis is of major importance for the evaluation of social relationships in group housed horses. However, in equine behaviour literature, studies on social bond analysis are inconsistent. Mutual grooming (horses standing side by side and gently nipping, nuzzling, or rubbing each other), affiliative approaches (horses approaching each other and staying within one body length), and measurements of spatial proximity (horses standing with body contact or within two horse-lengths) are commonly used. In the present study, we assessed which of the three parameters is most suitable for social bond analysis in horses, and whether social bonds are affected by individual and group factors. We observed social behaviour and spatial proximity in 145 feral horses, five groups of Przewalskiâ��s horses (N = 36), and six groups of feral horses (N = 109) for 15 h per group, on three days within one week. We found grooming, friendly approaches, and spatial proximity to be robust parameters, as their correlation was affected only by the animalsâ�� sex (GLMM: N = 145, SE = 0.001, t = â��2.7, p = 0.008) and the group size (GLMM: N = 145, SE < 0.001, t = 4.255, p < 0.001), but not by the horse breed, the aggression ratio, the social rank, the group, the group composition, and the individuals themselves. Our results show a trend for a correspondence between all three parameters (GLMM: N = 145, SE = 0.004, t = 1.95, p = 0.053), a strong correspondence between mutual grooming and friendly approaches (GLMM: N = 145, SE = 0.021, t = 3.922, p < 0.001), and a weak correspondence between mutual grooming and spatial proximity (GLMM: N = 145, SE = 0.04, t = 1.15, p = 0.25). We therefore suggest either using a combination of the proactive behaviour counts mutual grooming and friendly approaches, or using measurements of close spatial proximity, for the analysis of social bonds in horses within a limited time frame.
	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	2076-2615	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6428
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	Author	VanDierendonck, M.C.; de Vries, H.; Schilder, M.B.H.; Colenbrander, B.; Þorhallsdóttir, A.G. and Sigurjónsdóttir, H.
	Title	Interventions in social behaviour in a herd of mares and geldings			Type	Journal Article
	Year	2009	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	116	Issue	1	Pages	67-73
	Keywords	Horses; Social relationships; Interventions; Mares; Geldings; Play; Allogrooming; Social network
	Abstract	Social dynamics and maintenance of social cohesion were studied by analysing social interventions in two groups of horses consisting of adult mares, their offspring, adult geldings and sub-adults. The animals were observed for a total of 1316Â h. All relevant dyadic and triadic social interactions, including initial behaviour, possible intervention and outcome, were recorded. The main question was: do horses use interventions in affiliative interactions to safeguard their social network? Horses were significantly more likely to intervene in allogrooming or play interactions that involved a preferred partner. The stronger the preferred association in allogrooming, the higher the likelihood the intervener took over allogrooming with an initial dyad member. Interveners originating from two newly introduced groups (nÂ =Â 3 and 5), intervened significantly more often when a member of their own group allogroomed with an unfamiliar horse. In play, no correlation with unfamiliarity was found. Overall, the intervening horses stopped more than half of the initial allogrooming interactions, and a third of all interactions. Therefore, social facilitation cannot sufficiently explain interference behaviour. This study shows that maintaining relationships with preferred partners is important to horses and has implications for equine husbandry and management.
	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	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4766
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	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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