Abstract: Horses are likely to exhibit aggression when meeting for the first time. Therefore, this study compared 3 methods for mixing horses to evaluate their effectiveness in reducing aggressive interactions: (1) mixing pairs of horses in a paddock (P, 10 minutes, 15 tests), (2) introducing 1 unfamiliar horse to a pair of familiar, resident horses in a paddock (PP, 10 minutes, 15 tests), (3) allowing limited physical contact between pairs of horses for a short period of pre-exposure in neighboring boxes (B, 5 minutes, 16 tests) before mixing them in a paddock (BP, 10 minutes 16 tests). A total of 16 Swedish Standardbred mares, aged 6-18 years (mean age ± SD: 11 ± 4.4), were included in the study. Half of the horses were familiar with each other (resident horses, n = 8), whereas the other half were bought in from a variety of sources (unfamiliar horses, n = 8). Social interactions, consisting of behaviors from the sender, the receiver, and the subsequent sender's response, were recorded continuously as frequencies. There were no differences in the frequencies of aggressive behaviors between the 3 mixing methods, including those aggressive behaviors in which physical contact had been attempted (kick, strike). Although resident horses were overall more aggressive (median number of aggressive behaviors per horse, 62; Q1, 36; Q3, 68.5) than unfamiliar horses (median per horse, 4; Q1, 2; Q3, 12.5) during all tests (U = 97, P = 0.003), none of the 62 tests needed to be terminated. Unfamiliar horses did not receive more aggression from resident horses in PP (mean per test ± SD: 5.1 ± 3.1) than in P (mean per test ± SD: 6.4 ± 4.9) (t = 0.63, P = 0.544). However, the behavior “attack” was more frequent in PP (median per test, 2; Q1, 0; Q3, 5) than in P (median per test, 0; Q1, 0; Q3, 1) (U = 282, P = 0.042), and “flee” was more frequent in PP (median per test, 6; Q1, 4; Q3, 8) than in P (median per test, 1; Q1, 0; Q3, 6) (U = 290, P = 0.018). Pre-exposure in boxes did not reduce aggression in BP (median per test, 7; Q1, 4.3; Q3, 11.8) as compared with P (median per test, 6; Q1, 2; Q3, 16) (U = 264, P = 0.767), but during pre-exposure in B tests, horses exchanged more nonaggressive (median per test, 2; Q1, 0.3; Q3, 4) than aggressive (median frequency of aggressive behavior, 0; Q1, 0; Q3, 1) (W = 71, P = 0.013) and mixed interactions (median per test, 0; Q1, 0; Q3, 1) (W = 92, P = 0.016) through the opening. Results suggest mixing an unfamiliar horse with 2 resident horses at the same time instead of one by one may be preferable. In this way, the total aggression received by the unfamiliar horse will potentially be less, even though aggressive interactions may be more intense.

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: Gregarious animals living in permanent social groups experience intra-group competition. Conflicts over resources can escalate into costly aggression and, in some conditions, non-dispersive forms of conflict resolution may be favoured. Post-conflict friendly reunions, hence reconciliation, have been described in a variety of species. The aim of this study was to explore, for the first time, the occurrence of reconciliation in a group of domestic horses (Equus caballus) and learn more about strategies used to maintain group cohesion. The behaviour of seven horses living as permanent group in an enclosure for at least 2 years was observed by video for 108 h from June to August 2007. We used a Post-Conflict/Matched Control method to assess the existence of reconciliation and third-party affiliation. Behaviours recorded Post-Conflict, or during Matched Control periods, were classified as affiliative based on previous descriptions of visual communication patterns in horses. The proportion of attracted pairs over total post-conflict situations was significantly greater than the proportion of dispersed pairs, both during dyadic interactions (p < 0.001) and during triadic interactions (p = 0.002). The results of the present study show that both dyadic reconciliation and third-party post-conflict affiliative interactions form important social mechanisms for managing post-conflict situations in horses.

Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.

Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.