Abstract: Growing evidence shows a variety of sensorial and motor asymmetries in social and non-social interactions in various species, indicating a lateralized processing of information by the brain. Using digital video cameras on tripods and drones, this study investigated lateralization in frequency and duration of social behavior patterns, in affiliative, agonistic, and resting contexts, in a feral population of horses (Equus ferus caballus) in Northern Portugal, consisting of 37 individuals organized in eight harem groups. Affiliative interactions (including grooming) were more often performed, and lasted longer, when recipients were positioned to the right side. In recumbent resting (animals lying down) episodes on the left side lasted longer. Our results of an affiliative behavior having a right side tendency, provide partial support to the valence-specific hypothesis of Ahern and Schwartz (1979) – left hemisphere dominance for positive affect, affiliative behaviors. Longer recumbent resting episodes on the left side may be due to synchronization. However, in both instances it is discussed how lateralization may be context dependent. Investigating the position asymmetries of social behaviors in feral equids will contribute to a better understanding of differential lateralization and hemispheric specialization from the ecological and evolutionary perspectives.

Abstract: Horses are a highly social species living in complex social systems which should require them to memorise and generalise social experiences and distinguish between familiar and unfamiliar conspecifics. In the main part of my thesis I concentrated on the specific conflict situation of a horse being introduced into a new social group, and investigated its behaviour and stress level. Horses were either introduced (1) immediately, (2) after an observation period, or (3) together with an integration horse after an observation period. Additionally, in the second part of my thesis I arranged several experiments to elaborate additional aspects which could affect the behaviour of horses during introductions. In this study I could describe a simplified method for measuring stress through the analysis of faecal GCMs in horses. An enzyme immunoassay (EIA) for 11-oxoaetiocholanolone using 11-oxoaetiocholanolone-17-CMO: BSA (3?,11-oxo-A EIA) as antigen showed high amounts of immunoreactive substances. The new assay increases the accuracy of the test and lowers the expenses per sample; also storing of samples at room temperature after collection is less critical. This is a big advantage both in the field of wildlife management of equids and in the field of equestrian sports (chapter 1). Comparing the different introduction techniques, the introduction with an integration horse led to significantly less total interactions and lower levels of aggression than the introduction of single horses, both immediately and after several days of observing the new group. Additionally, by observing the behaviour of the horses during everyday sociality I could develop a formula describing the interrelationship between expected aggression level and enclosure size per horse. The curve takes an exponential shape. Starting from a space allowance of 300 m2 and more per horse, the amount of aggressions per hour approaches zero. For the reduction of aggression levels and injury risks in socially kept horses I recommend an enclosure size of at least 300 m2 per horse (chapter 2). I further investigated the stress level of the introduced animals. Horses which were immediately introduced did not show elevated faecal GCMs. In contrast, horses which were introduced after an observation period had slightly elevated values 2 and 3 days after the introduction. For horses introduced together with an integration horse faecal GCMs were significantly above the baseline value on the day of introduction and 1 day after it. These differences between introduction techniques indicate that the introduction event itself is not as stressful as previously assumed. Rather standing together with an integration horse and not being able to integrate immediately into the complete group elicits stress in horses (chapter 3). In the commentary of chapter 4 several studies are discussed which failed to demonstrate social learning in horses. It is argued that they did not consider important aspects which could have an influence, such as the dominance status or the social background of the horses (chapter 4). In chapter 5 a social feeding situation was investigated. The social rank as well as the position of conspecifics affected the feeding strategy of horses. Domestic horses used social cognition and strategic decision making in order to decide where to feed. When possible they tended to return to the same, continuously supplied feeding site and switched to an ?avoidance tendency? in the presence of dominant horses or when another horse was already feeding there (chapter 5). One possibility to recognize group members is through olfactory recognition. In chapter 6 it is shown that horses are able to distinguish their own from their conspecifics? faeces. In addition, they paid most attention to the faeces of those group members from which they received the highest amount of aggressive behaviour (chapter 6). Horses show cognitive abilities because they are able to use humans as local enhancement cues when searching for food, independently of their body posture or gaze consistency when the persons face them. Moreover, they seem to orientate on the attention of familiar persons more than of unfamiliar persons (chapter 7). Altogether, the results of this thesis provide further support for the view that horses show good conflict resolution strategies. They are perfectly able to deal with the conflict situation of being introduced to new group members, and the introduction event itself is not as stressful as previously assumed. It is rather suggested that standing together with an integration horse and not being able to integrate immediately into the complete group elicits stress in horses. All additional experimental set-ups could demonstrate that horses are well capable of social cognition.

Abstract: Horses fulfill many of the criteria for a species in which it would be adaptive to be capable of individual recognition: they are highly social, form strong and long lasting bonds, their affiliations are rarely kin based, they have a fission-fusion social structure and they possess inter and intra-group dominance hierarchies.
We used a novel cross-modal, expectancy violation paradigm to provide the first systematic evidence that a non-human animal – the domestic horse- is capable of cross modal recognition. We believe this paradigm could provide an ideal way to study individual recognition across a wide range of species.
For full published details see: Proops L, McComb K, Reby D (2009) Cross-modal individual recognition in domestic horses (Equus caballus). Proc Natl Acad Sci U S A 106: 947-951.

Abstract: In functional laterality research, most ungulate livestock species have until recently been mainly overlooked. However, there are many scientific and practical benefits of studying laterality in ungulate livestock. As social, precocial and domestic species, they may offer insight into the mechanisms involved in the ontogeny and phylogeny of functional laterality and help to better understand the role of laterality in animal welfare. Until now, most studies on ungulate livestock have focused on motor laterality, but interest in other lateralized functions, e.g., cognition and emotions, is growing. Increasingly more studies are also focused on associations with age, sex, personality, health, stress, production and performance. Although the full potential of research on laterality in ungulate livestock is not yet exploited, findings have already shed new light on central issues in cognitive and emotional processing and laid the basis for potentially useful applications in future practice, e.g., stress reduction during human-animal interactions and improved assessments of health, production and welfare. Future research would benefit from further integration of basic laterality methodology (e.g., testing for individual preferences) and applied ethological approaches (e.g., established emotionality tests), which would not only improve our understanding of functional laterality but also benefit the assessment of animal welfare.

Abstract: In the paper we investigate how owner personality, attitude and gender influence dog behavior, dyadic practical functionality and the level of dog salivary cortisol. In three meetings, 12 female and 10 male owners of male dogs answered questionnaires including the Neo-FFI human personality inventory. Their dyadic behavior was video-taped in a number of test situations, and saliva samples were collected. Owners who scored highly in neuroticism (Neo-FFI dimension one) viewed their dogs as social supporters and spent much time with them. Their dogs had low baseline cortisol levels, but such dyads were less successful in the operational task. Owners who scored highly in extroversion (Neo-FFI dimension two) appreciated shared activities with their dogs which had relatively high baseline cortisol values. Dogs that had female owners were less sociable-active (dog personality axis 1) than dogs that had male owners. Therefore, it appears that owner gender and personality influences dyadic interaction style, dog behavior and dyadic practical functionality.