Abstract: In most studies of social learning in animals, no attempt has been made to examine the nature of the strategy adopted by animals when they copy others. Researchers have expended considerable effort in exploring the psychological processes that underlie social learning and amassed extensive data banks recording purported social learning in the field, but the contexts under which animals copy others remain unexplored. Yet, theoretical models used to investigate the adaptive advantages of social learning lead to the conclusion that social learning cannot be indiscriminate and that individuals should adopt strategies that dictate the circumstances under which they copy others and from whom they learn. In this article, I discuss a number of possible strategies that are predicted by theoretical analyses, including copy when uncertain, copy the majority, and copy if better, and consider the empirical evidence in support of each, drawing from both the animal and human social learning literature. Reliance on social learning strategies may be organized hierarchically, their being employed by animals when unlearned and asocially learned strategies prove ineffective but before animals take recourse in innovation.

Abstract: In humans, speech perception is lateralized, with the left hemisphere of the brain dominant in processing the communicative content and the right hemisphere dominant in processing the emotional content. However, still little is known about such a division of tasks in other species. We therefore investigated lateralized processing of communicative and emotionally relevant calls in a social mammal, the pig (Sus scrofa). Based on the contralateral connection between ears and hemispheres, we compared the behavioural and cardiac responses of 36 young male pigs during binaural and monaural (left or right) playback to the same sounds. The playback stimuli were calls of social isolation and physical restraint, whose communicative and emotional relevance, respectively, were validated prior to the test by acoustic analyses and during binaural playbacks. There were indications of lateralized processing mainly in the initial detection (left head-turn bias, indicating right hemispheric dominance) of the more emotionally relevant restraint calls. Conversely, there were indications of lateralized processing only in the appraisal (increased attention during playback to the right ear) of the more communicative relevant isolation calls. This implies differential involvement of the hemispheres in the auditory processing of vocalizations in pigs and thereby hints at similarities in the auditory processing of vocal communication in non-human animals and speech in humans. Therefore, these findings provide interesting new insight in the evolution of human language and auditory lateralization.

Abstract: In Experiment 1, rats choosing in an E maze preferred to release a rat standing in a pool of water to dry ground over a rat already standing on dry ground. Five additional experiments showed that the choosing rat's preference for releasing the wet rat was maintained by two separable outcomes: (1) the social contact offered by the released rat and (2) the reinforcing value of proximity to a pool of water. These results call into question Sato et al.'s (Anim Cogn 18:1039-1047, 2015) claim to have demonstrated that a rat's releasing of a wet rat to dry ground is empathically motivated.

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: Identifying valid indicators to assess animals' emotional states is a critical objective of animal welfare science. In horses, eye wrinkles above the eyeball have been shown to be affected by pain and other emotional states. From other species we know that individual characteristics, e.g. age in humans, affect facial wrinkles, but it has not yet been investigated whether eye wrinkle expression in horses is systematically affected by such characteristics. Therefore, the aim of this study was to assess how age, sex, breed type, body condition and coat colour affect the expression and/or the assessment of eye wrinkles in horses. To this end, we adapted the eye wrinkle assessment scale from Hintze et al. (2016) and assessed eye wrinkle expression in pictures taken from the left and the right eye of 181 horses in a presumably neutral situation, using five outcome measures: a qualitative first impression reflecting how worried the horse is perceived by humans, the extent to which the brow is raised, the number of wrinkles, their markedness and the angle between a line through both corners of the eye and the topmost wrinkle. All measures could be assessed highly reliable with respect to intra- and inter-observer agreement. Breed type affected the width of the angle (F2, 114 = 8.20, p < 0.001), with thoroughbreds having the narrowest angle (M = 23.80, SD = 1.60), followed by warmbloods (M = 28.00, SD = 0.60), and coldbloods (M = 31.00, SD = 0.90). None of the other characteristics affected any of the outcome measures, and eye wrinkle expression did not differ between the left and the right eye area (all p-values > 0.05). In conclusion, horses' eye wrinkle expression and its assessment in neutral situations was not systematically affected by the investigated characteristics, except for 'breed type', which accounted for some variation in 'angle'; how much eye wrinkle expression is affected by emotion or perhaps mood needs further investigation and validation.