Abstract: In recent years, Przewalski’s horses have been increasingly kept in semi-reserves. However,there areonly few studies ontheir behaviour and their ability to adaptto management interventions.In the main part of my dissertation, I focus on investigatingthe animals’ behaviour in different semi-reserves with varyinghabitats and living spaces. In addition, I investigate the horses’ behaviour during various management interventionsand analysetheensuing changes instress levels. Another aspect of my dissertation is the studyof social behaviour inPrzewalski’s horses. I investigate theparameters that should be used to demonstrate social bonds between individualsandassess whichdata provide the most meaningful results.In the commentary tochapter 1,several studies investigatingsocial bonds in horsesare discussed. Comparing the various studies, it is strikingthat no homogeneous analyses orevaluation criteria exist. While some authors only considersocial grooming, others include data onthe spatial proximity of the individuals in their evaluations, and various definitionsof proximity can also be found in the literature. Additionally, someauthors use friendly approaches between individuals asa furtherparameter wheninvestigating the social bonds.Continuing with this theme, in chapter 2I investigate the social behaviour of the horses and compare various analysis methods. I show that proactive behaviour, such as friendly approaches, is a good alternative to spatial proximity when investigating social bonds between group members, andis also useful for expanding the often very small data sets of mutual grooming in horses. Comparing Przewalski’s horses with wild living horses, I found no significant differences in the social behavior and the frequency of social interactions, regardlessof group size, group composition, habitat, and individual parameters such as age and gender.Inchapter 3,I investigate the behaviour of a Przewalski’s horse group when exploring a new area of their enclosure. Their behaviour changed, showing less resting and more feeding. Furthermore, the animals maintained greater distances from each other, and the alpha male, instead of herding the group from behind, led the group around the new area and walked in front of the other group members. Moreover, he showed a substantial increase in stress level during the first day.A general comparison of the behaviour of the Przewalski’s horses in different semi-reserves is provided in chapter 4. In it, the habitat choice of the animals and their reactions to various management interventions are investigated. It is shown that Przewalski’s horses prefer open grassland to dense woods, although keeping Przewalski’s horses in a pine forest does not influence the animals’ stress level. In contrast to habitat, food range, and changes in the group composition, which do not appear to change stress levels, individual factors, such as the hierarchy, influence the glucocorticoid level of the animals significantly. The largest increases in stress hormones were demonstrated when the horses were temporarily confined in smaller areas.The importance of the available space is also discussed in chapter 5, where it is shown that horses show less aggressive behaviour when more space is provided. In contrast, the husbandry system does not influence the animals’ aggression, but the way of feeding can additionally reduce agonistic behaviour, for example if food is offered ad libitum.In summary, the results of this study provide indications for the optimization of keeping Przewalski’s horses in semi-reserves. The animals can adapt themselves to the environment and thrive in habitats which do not correspond to their original steppe-like home. Nevertheless, the semi-reserves should provide sufficient grassland, as the horses prefer this type of habitat. General speaking, any types of habitat can only offer a suitable living space if the food range is sufficient for the number of horses. Otherwise, and especially during could winter months, supplementary feeding is necessary according to the body condition of the animals. This is particularly important for older, weakened, or very young animals, which are still adapting to life in the semi-reserve. Without sufficient food, stress hormones can increase and negatively influence the well-being of the horses. The same is true for management interventions: restricting the animals to small enclosures, for example, can adversely affect the horses’ well-being and should be only done if absolutely necessary. Targetedbehaviour observations allow the animals that have a special meaning for the group to be identified, and these should not be taken out of the group unless it is unavoidable, as young and unexperienced horses orientate themselves on those animals. This is especially true for the alpha male in a bachelor group, as these groups are often composed of young horses and the alpha-male provides the necessary stability and experience. Social bonds between individuals can be investigated by observing friendly and proactive behaviour, and social grooming and friendly approaches yield suitable data for such analysis.

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: 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&#65533;&#65533;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&#65533;&#65533; sex (GLMM: N = 145, SE = 0.001, t = â&#65533;&#65533;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.