Abstract: 1. The social relationships in a group of Icelandic horses without a mature stallion were studied. The horses were all familiar to each other. Mutual grooming and play relationships, spatial associations, dominance-subordinate relations and the effect of kinship on these relationships were analysed.TAGSTARTBRTAGEND 2. The social structure was clearly dominated by the behaviour of the adult mares. The horses preferred to form bonds within their social class (sex/age) and they kept close proximity with their friends. The group was effectively divided into two social subgroups, adult mares as one group and adult geldings and sub-adults as another group. The sub-adults and adult geldings formed associations, which were based on mutual grooming and play, while the adult mares did not play. Differences between the sexes were evident. Males played more than the females, had more playing partners and were more popular as playmates.TAGSTARTBRTAGEND 3. Aggression rates were low. The dominance hierarchy was linear. Adult mares ranked higher than adult geldings, sub-adults and the foals. Rank was significantly correlated with age. The closer the adult mares were in rank, the more they groomed with each other. Such relationships were not found amongst the other social group.TAGSTARTBRTAGEND 4. Kinship was calculated between all pairs of animals for up to 4 or 5 generations. Allogrooming and play frequencies and proximity were all positively correlated with kinship. Adult mares, which were close in the dominance hierarchy, were on average more related than those further apart.TAGSTARTBRTAGEND 5. The social relationships in the Icelandic herd were, to some extent, different from relationships reported from unmanaged and feral horse-herds with mature stallions and bachelors. Our results suggest that adult mares groom more in groups without a stallion. Furthermore, they have more preferred partners than in natural harems and their partners are other adult mares, not their weaned offspring as seems to be the case in feral herds. The sub-adults also seem to be more socially active in the absence of stallions. Interestingly, in the Icelandic group, the adult mares showed stallion like behaviours, like mounting and protecting foals. Only by studying the behaviour and the nature of the relationships of horses in groups of different compositions, can we expect to gain a comprehensive understanding about individual social strategies and cognitive capabilities of the species. Such knowledge is valuable for management and welfare of the horse.

Abstract: During 1992 and 1993, 14 reintroduced Przewalski Horses or Takhi (Equus ferus przewalskii) were studied in the Hustain Nuruu Mountain Steppe reserve in Mongolia. Most of the individuals did not know each other before reintroduction. These Takhi were the first of five groups due to be released in the reserve after an acclimatisation period of at least 1 year. During acclimatisation the Takhi, lived visually and acoustically separately, in fenced enclosures of approximately 45 ha each. The observations, mostly scan-sampling, were carried out in each season. The observation bouts were divided over six periods and over two harem herds. Two of the periods were in the same consecutive seasons, so comparison over the years was possible. Social integration within the Takhi herds was very high from the beginning, as described by the spatial relation and synchronisation data. Between 50 and 89% of the observation time, the behaviour of all herd members was synchronised. The amount of time spent grazing by the Takhi (30-68% of the daylight period) was similar to that of feral horses and Takhi in captivity and semi-reserves. The Takhi tended to rest in the morning and have a bimodal period of grazing at dawn and in the afternoon. The Takhi displayed clear habitat preferences for certain activities. They had a strong preference to rest at the highest point in their enclosure. They fed preferably on two or three different vegetation types (with five types available in each enclosure). The amount of time spent grazing during the non-growing seasons (49 +/- 15%) indicates that the feeding value and availability of food were sufficient. Health changes were detected adequately using condition scoring sheets. No supplementary food or water was supplied during the harsh winters. Moreover, low mortality rates and high reproductive success show that the mountain steppe is a habitat which is potentially suitable for establishing a healthy Takhi population. Takhi is the first species to return to its native habitat after living only in zoos for so many generations.

Abstract: We observed the activity and movements of Konik horses (Equus caballus) in order to determine the initiators of movements and the individuals taking the lead in movements of the main groups. We conducted our observations between March-June 2010 along the shores of the Rhine river, in the Blauwe Kamer reserve, in the Netherlands. The horses were introduced to the reserve alongside with cattle to prevent the growth of the forest and maintain the grass habitat. We videotaped all observations on two digital video cameras, one providing the general view of the group and the other scanning and focusing on the individuals, to aid with identification. Horses were recognized based on individual profiles that were created for each one, consisting of photos of both sides and notes of the main characters, such as orientation and coloration of the mane, prominent scars and markings, etc. Twenty three horses (not counting foals) were organized in two harem groups with 11 individuals (two of which were dominant stallions) in the large group and six individuals (one of which was a stallion) in the second group. These two main groups were always within sight of each other, and two bachelor males moved usually in their vicinity. An additional group of three young bachelor males roamed elsewhere in the reserve. We divided the movements of the horses to local movements while grazing and to long-distance movements, in which the horses moved to a different grazing location, to a pond of water, resting area or groups of trees that were used by the horses for scratching themselves. During the local movements, any two of the three oldest females in the large harem group were enough to cause the whole group to follow them and gradually change position within the pasture. The smaller harem always followed the large harem’s movements. The long-distance movements of the large harem were sometimes initiated by one of the harem stallions that herded their group and at other times – by the oldest females. Soon after the movement was clearly initiated, the dominant stallions positioned themselves in the back of the group or in the center and had no influence on the direction of move that was only determined by the leading females. In the long-distance movements too, the small harem followed the large harem group, and the two bachelor males usually followed behind them. Social interactions included aggressive interactions between the two dominant males within the large harem or between dominant stallions and the two bachelor males accompanying the two harems. In addition, dominant males courting or attempting to mate with mares sometimes caused a turmoil that eventually initiated movement of the harems. KW -

Abstract: For some time now, social network analysis provides tools to describe concisely the social structure of animals. Nevertheless, the factors that shape the social network and determine the frequency of different types of affiliations are often unclear. For example, the effects of relatedness on social networks have been only studied on relatively few species. Both among social or sub-social species the effects ranged from no effect to a substantial effect. Here we test the hypothesis that relatedness has an effect on the social network of horses (Equus caballus) that live freely in semi-natural conditions, and specifically, that individuals with high values of relatedness to their neighbors in the network will have fewer links Wolf et al, 2011). This hypothesis is based on the notion that related individuals have common interests and are likely to assist each other, and may need less links to other, unrelated individuals. In addition, individuals of similar age and sex are likely to have common needs, and thus are more likely to be associated. We thus tested a second hypothesis: homophily to individuals with similar age and sex will have a role in determining the associations within the social network. The field study was conducted on 27 horses in the Blauwe Kamer reserve in the Netherlands (1.1 sq km). We videotaped horses and their groups and used the information from 22 horses, after excluding the five foals from the analysis. Relatedness was calculated from the pedigree, which was based on parentage, determined by DNA analysis. The social network was constructed based on spatial proximity data. We assessed the influence of relatedness, age-homophily and sex-homophily on the network structure with Multiple Regression Quadratic Assignment Procedure (MRQAP) (Krackhardt 1988), with the R package sna. The results show that there was no significant effect of relatedness on the network, nor an effect of age-homophily. Nevertheless, we found a significant effect of sex-homophily, the tendency of individuals to associate with individuals of the same sex. We argue that the lack of a relatedness effect is not likely to have been caused due to the inability to detect who is kin. The structured social system in horses includes strong associations between often unrelated individuals, alongside with young individuals leaving their natal harem as part of the inherent inbreeding avoidance (Linklater & Cameron 2009; Boyd et al. in press). The significant effect of sex-homophily could stem from the protection females get from associations with other females, e.g., in the reduction of harassment from males. Previous studies on feral horses showed that mares that were better connected with other females in their harem benefited from higher survival rates to their foals (Cameron et al. 2009). The associations among bachelor males could also contribute to the strength of the sex-homophily effect.
In order to generalize from our results, one needs to examine additional populations of horses, because the conditions in the Blauwe Kamer reserve may not be representative, mainly due to the limited opportunity for dispersal in a restricted area.
Keywords: Long-term affiliation; spatial proximity, kin detection
Boyd, L., Scorolli, A. L., Nowzari, H., & Bouskila, A. (2016). Chapter 2: Social organization. In J. I. Ransom, & P. Kaczensky (Eds.), Wild equids. Baltimore, Maryland (in press): The Johns Hopkins University Press.
Cameron, E. Z., Setsaas, T. H., & Linklater, W. L. (2009). Social bonds between unrelated females increase reproductive success in feral horses. Proceedings of the National Academy of Sciences, 106(33), 13850-13853
Krackhardt, D. (1988). Predicting with networks: Nonparametric multiple regression analysis of dyadic data. Social Networks, 10(4), 359-381.
Linklater, W. L., & Cameron, E. Z. (2009). Social dispersal but with philopatry reveals incest avoidance in a polygynous ungulate. Animal Behaviour, 77(5), 1085-1093.
Wolf, J. B., Traulsen, A., & James, R. (2011). Exploring the link between genetic relatedness r and social contact structure k in animal social networks. The American Naturalist, 177(1), 135-142.