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.

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.

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.

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.

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 1316 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 (n = 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.