Abstract: Both humans and group-living animals associate and behave affiliatively more with some individuals than others. Human friendship has long been acknowledged, and recently scientists studying animal behaviour have started using the term friendship for close social associates in animals. Yet, while biologists describe friends as social tools to enhance fitness, social scientists describe human friendship as unconditional. We investigate whether these different descriptions reflect true differences in human friendship and animal close social associations or are a by-product of different research approaches: namely social scientists focussing on proximate and biologists on ultimate explanations. We first stress the importance of similar measures to determine close social associations, thereafter examine their ultimate benefits and proximate motivations, and discuss the latest findings on the central-neural regulation of social bonds. We conclude that both human friendship and animal close social associations are ultimately beneficial. On the proximate level, motivations for friendship in humans and for close social associations in animals are not necessarily based on benefits and are often unconditional. Moreover, humans share with many animals a similar physiological basis of sociality. Therefore, biologists and social scientist describe the same phenomenon, and the use of the term friendship for animals seems justified.

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: Domestic horses are faced with social challenges throughout their lives due to limitations in social contact, space restrictions and frequent changes in social companionship. This is in contrast to natural conditions where horses live in relatively stable harem bands. Currently, little is known about how repeated regrouping affect horse behaviour and welfare, and it is unknown whether horses may adapt to regrouping. In this study, we aimed to investigate the effects of an unstable group structure, caused by weekly regroupings, on behaviour and frequency of injuries in young horses. Forty-five horses were included in the study and were randomly assigned to the treatments; Stable (S; seven groups of three horses) or Unstable (U; eight groups of three horses). The experimental period lasted 7 weeks, during which horses in Stable groups remained in the same group, whereas one horse was exchanged between Unstable groups every week. The groups were kept in 80m×80m grass-covered enclosures and were fed additional roughage on the ground daily. Social interactions were recorded in Unstable groups immediately after each regrouping (30min), and in both Stable and Unstable groups on day 1, 3 and 6 after each regrouping (2×20min/group/day). Injuries were scored by the end of the experimental period. The level of aggression shown by horses in Unstable groups immediately after regrouping was not affected by week (F5,35=0.42, P=0.83), indicating that horses neither habituated, nor sensitized, to repeated regrouping. Compared to horses in Stable groups, more agonistic behaviour was shown by horses in Unstable groups (i.e. non-contact agonistic; F1,65=5.60, P=0.02), whereas there was no treatment effect on other variables. The level of play behaviour appeared, however, to be more variable in Unstable groups. There was a significant effect of week on the level of contact agonistic interactions as well as greeting behaviour, due to a high occurrence in weeks 4-6. Non-contact agonistic interactions constituted the major part of agonistic interactions (66%). Possibly as consequence, no serious injuries were registered and there was no treatment effect (U=184; P=0.11). We conclude that the behaviour of young horses is affected by group management, and that horses appear not to adapt to weekly regroupings.

Abstract: Group housed horses at a stud farm/riding stable in Belgium were observed on 17 days between 21 February and 25 April 2008, totalling 54hr25min of detailed data. The original group consisted of 8 Irish Cob mares, 1 Warmblood mare, 1 Arabian gelding and 2 Arabian mares. The group had been established in December 2007. During the course of the study 5 horses were removed from the group and 2 foals were born. 3 highly pregnant mares were housed adjacent to the group for part of the period. Horses were regularly used for lessons. Available surface area differed with the group on pasture at the end. Continuous all occurrence sampling of 10 agonistic and 2 affiliative behaviours was carried out for all group members present. Overall the group showed a frequency of 44.75 agonistic interactions per hour and 11.25 affiliative per hour. Of those agonistic interactions 46.3% were threats while 47% were less active interactions (displacement, being avoided), leaving only 6.7% more aggressive interactions ( mainly biting, some kicking and chasing). The effect on acting agonistically was not significant for age (p=0.1591) and borderline significant for density (p=0.0627). The analysis of the frequency of affiliative interactions showed there is no significant effect of age (p=0.1865) or density (p=0.7923). Agonistic and affiliative interactions were not significantly correlated (p=0.72). Affiliative behaviour a horse received showed a borderline effect (p=0.0787) on agonistic behaviour, as did the interaction between received agonistic and affiliative interactions (p=0.0725). Received agonistic interactions had a borderline negative effect (p=0.0656) on affiliative behaviour. A dominance hierarchy was calculated based on agonistic interactions using Empirical Bayes’ estimates based on Poisson regression with random effects. Agonistic behaviour expressed to other horses was significantly effected by relative rank (p=0.0243). Overall horses tended to be 3.7 times more aggressive towards lower ranking horses compared to higher ranking horses. Affiliative behaviour shown to other horses was not significantly influenced by the rank of the social partner (p=0.7915). Some individuals did show a significant effect whereby they showed more affiliative behaviour towards lower ranking individuals. This study was a small project to look at a practical situation of riding horses being kept in group housing. The frequent changes in group composition and available surface made it possible to look at agonistic and affiliative interactions in such circumstances. This is useful as instability in group composition is often used as main reason not to keep horses in social groups. The results from this study showed a borderline effect of density on agonistic behaviour. In reality it was also influenced by practical details, like a narrow paddock with only 2 hay crates on the smallest surface. Rank in a dominance hierarchy, based on agonistic behaviour, had a significant effect on the agonistic behaviour expressed towards higher or lower ranking horses. No injuries or escalating fights were observed. This study shows it is possible to keep a group of riding horses in a social context without excessive aggression.

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.