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: Social learning is taxonomically widespread and can provide distinct behavioural advantages, such as in finding food or avoiding predators more efficiently. Although extensively studied in bony fishes, no such empirical evidence exists for cartilaginous fishes. Our aim in this study was to experimentally investigate the social learning capabilities of juvenile lemon sharks, Negaprion brevirostris. We designed a novel food task, where sharks were required to enter a start zone and subsequently make physical contact with a target in order to receive a food reward. Naive sharks were then able to interact with and observe (a) pre-trained sharks, that is, ‘demonstrators’, or (b) sharks with no previous experience, that is, ‘sham demonstrators’. On completion, observer sharks were then isolated and tested individually in a similar task. During the exposure phase observers paired with ‘demonstrator’ sharks performed a greater number of task-related behaviours and made significantly more transitions from the start zone to the target, than observers paired with ‘sham demonstrators’. When tested in isolation, observers previously paired with ‘demonstrator’ sharks completed a greater number of trials and made contact with the target significantly more often than observers previously paired with ‘sham demonstrators’. Such experience also tended to result in faster overall task performance. These results indicate that juvenile lemon sharks, like numerous other animals, are capable of using socially derived information to learn about novel features in their environment. The results likely have important implications for behavioural processes, ecotourism and fisheries.

Abstract: Horses are highly social animals that have evolved to live in social groups. However, in modern husbandry systems, single housing prevails where horses experience social isolation, a challenge-to-welfare factor. One major reason for this single housing is the owners’ concerns that horses may injure each other during aggressive encounters. However, in natural conditions, serious injuries due to aggressive encounters are rare. What could therefore explain the claimed risks of group living for domestic horses? Basing our questioning on the current knowledge of the social life of horses in natural conditions, we review different practices that may lead to higher levels of aggression in horses and propose practical solutions. Observations of natural and feral horses mostly indicate a predominance of low frequencies and mild forms of aggression, based on subtle communication signals and ritualized displays and made possible by group stability (i.e. stable composition), dominance hierarchy and learning of appropriate social skills by young horses. Obviously, adults play a major role here in canalizing undesirable behaviours, and social experience during development, associated with a diversity of social partners, seems to be a prerequisite for the young horse to become socially skilled. Given the natural propensity of horses to have a regulation of aggression in groups, the tendency to display more aggression in groups of domestic horses under some management practices seems clearly related to the conditions offered. We therefore review the managing practices that could trigger aggressiveness in horses. Non social practices (space, resource availability) and social practices (group size, stability of membership, composition and opportunities for social experiences during development) in groups of domestic horses are discussed here. Finally, we propose simple practical solutions leading to more peaceful interactions in groups of domestic horses, based on the knowledge of horses’ natural social life which therefore should be enhanced (e.g. ensuring roughage availability, favouring group stability, introducing socially experienced adults in groups of young horses, etc.). The state of the art indicates that many questions still need to be answered. Given the importance of the associated welfare issues and the consequences on the use of horses, further research is required, which could benefit horses… and humans.

Abstract: Removing a horse from its social group may be considered risky, both for the handler and the horse, because other horses can interfere in the catching process. The main aim of this study was to identify where and when these risk situations occur while removing a horse from its group. A potential risk situation was defined by the closeness of loose horses in the group or by any physical contact with them. Whether the number of horses following would be influenced by the social rank of the horse being led out, and whether more horses would follow to the gate when a larger proportion of the group was removed compared to when a single horse was taken out were also investigated. Thirty-two mares (1–2 years) were kept in groups of four. All horses were taken out of their home paddock twice alone (64 tests) and twice with a companion (32 tests). One handler (or two handlers when two horses were removed) was asked to approach (phase 1) and catch the target horse (phase 2), walk it to the centre of the paddock and remain stationary at a post for 30 s (phase 3), walk to the paddock entrance (phase 4) and through the gate (phase 5). The number of horses following, and the number of loose horses in proximity (<2 m, 2–5 m) to the target horse and handler was estimated, and horse–horse and horse–human interactions were recorded continuously for the five scoring phases. Significantly more loose horses were within 2 m of a single target horse during the phases approach (mean ± SD: 1.5 ± 0.8), catch (1.6 ± 0.9) and post (1.7 ± 0.7) than during walk (1.0 ± 0.5) and gate (1.1 ± 0.6). Rank did not influence the number of horses following to the gate (high rank: 2.4 ± 0.7; lower rank: 2.0 ± 1.0; P = 0.396) and interactions between horses were rare. A greater proportion of the loose horses followed when two horses (0.9 ± 0.2) were removed compared to when a single horse (0.7 ± 0.3) was taken out (P = 0.011). In conclusion, maintaining a distance to other horses in the group by reducing the time being relatively stationary, so giving loose horses fewer chances to approach, is likely to contribute to improved handler's safety. Removing a small proportion of the group may also decrease the probability of the other horses following.

Abstract: Many group-living animals show social preferences for relatives, familiar conspecifics or individuals of similar attributes such as size, personality or sex. How such preferences could affect the collective motion of animal groups has been rather unexplored. We present a general model of collective animal motion that includes social connections as preferential reactions between individuals. Our conceptual examples illustrate the possible impact of underlying social networks on the collective motion of animals. Our approach shows that the structure of these networks could influence: (1) the cohesion of groups; (2) the spatial position of individuals within groups; and (3) the hierarchical dynamics within such groups. We argue that the position of individuals within a social network and the social network structure of populations could have important fitness implications for individual animals. Counterintuitive results from our conceptual examples show that social structures can result in unexpected group dynamics. This sharpens our understanding of the way in which collective movement can be interpreted as a result of social interactions.