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.

Abstract: Many studies have been carried out into both motor and sensory laterality of horses in agonistic and stressful situations. Here we examine sensory laterality in affiliative interactions within four groups of domestic horses and ponies (N = 31), living in stable social groups, housed at a single complex close to Vienna, Austria, and demonstrate for the first time a significant population preference for the left side in affiliative approaches and interactions. No effects were observed for gender, rank, sociability, phenotype, group, or age. Our results suggest that right hemisphere specialization in horses is not limited to the processing of stressful or agonistic situations, but rather appears to be the norm for processing in all social interactions, as has been demonstrated in other species including chicks and a range of vertebrates. In domestic horses, hemispheric specialization for sensory input appears not to be based on a designation of positive versus negative, but more on the perceived need to respond quickly and appropriately in any given situation.

Abstract: Prima facie, the acquisition of novel behaviors in animals through observation of conspecifics seems straightforward. There are, however, various mechanisms through which the behavior of animals can be altered from observing others. These mechanisms range from simple hard-wired contagious processes to genuine learning by observation, which differ fundamentally in cognitive complexity. They range from social facilitation and local enhancement to true social learning. The different learning mechanisms are the subject of this review, largely because research on learning by observation can be confounded by difficulties in interpretation owing to the looming possibility of associative learning infecting experimental results. While it is often assumed that horses are capable of acquiring new behavior through intra-species observation, research on social learning in horses includes a variety of studies some of which may overestimate the possession of higher mental abilities. Assuming such abilities in their absence can have welfare implications, e.g. isolating stereotypical horses on the assumption that these behaviors can be learned though observation by neighboring horses. This review summarizes the definitions and criteria for the various types of social transmission and social learning and reviews the current documentation for each type in horses with the aim of clarifying whether horses possess the ability to learn through true social learning. As social ungulates, horses evolved in open landscapes, exposed to predators and grazing most of the day. Being in close proximity to conspecifics may theoretically offer an opportunity to learn socially, however anti-predator vigilance and locating forage may not require the neural complexity of social learning. Given the significant energetic expense of brain tissue, it is likely that social facilitation and local enhancement may have been sufficient in the adaptation of equids to their niche. As a consequence, social learning abilities may be maladaptive in horses. Collectively, the review proposes a novel differentiation between social transmission (social facilitation, local and stimulus enhancement) and social learning (goal emulation, imitation). Horses are undoubtedly sensitive to intra-species transfer of information but this transfer does not appear to satisfy the criteria for social learning, and thus there is no solid evidence for true social learning in horses.

Abstract: Learning through the observation of others allows the transfer of information without the costs incurred during individual trial and error learning. Horses (Equus caballus) are a highly social species, which might be expected to be capable of learning from others, but experimental findings are inconsistent, and potentially confounded by social facilitation effects not related directly to the learning of the task. We refined the methods used in previous equine social learning studies, to examine and distinguish specific social influences on learning of a task: we used predefined group leaders rather than agonistically dominant individuals to demonstrate a detour task to familiar conspecific observers; in addition we had two control groups: a non-observer (true control) and a group with the demonstrator simply present at the goal (social facilitation control). 44 socially kept horses were allocated to one of the three test conditions and took part in five trials each. Success rate, latency and detour direction were recorded. There was no significant difference between the three groups in the likelihood of them succeeding in the task nor latency to succeed; however there was a significant difference in the route chosen by the groups, with the true control choosing the side with the entrance gate significantly more than either the observer group or social facilitation group. Both of the latter two groups chose to go in the same direction relative to themselves, regardless of which side the gate was. Seven out of nine horses in the observer group chose the same direction as their demonstrator every time. Our results show a significant role of social facilitation on detour behaviour and highlight the importance of including adequate controls for simpler cognitive influences on behaviour before claims can be made about the specific learning of motor actions or goal directed behaviour. Social cues may be important to horses if the task is sufficiently challenging and motivationally important, so future work should consider more demanding, but ecologically relevant situations, in order to maximise the potential revelation of social learning effects which do not depend on simple local or stimulus enhancement effects.

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.