Abstract: We investigated perseveration and detour behaviour in 36 equids (Equus caballus, E. asinus, E. caballus × E. asinus) and compared these data to those of a previous study on domestic dogs (Canis familiaris). The animals were required to make a detour through a gap at one end of a straight barrier in order to reach a visible target. After one, two, three or four repeats (A trials), the gap was moved to the opposite end of the barrier (B trials). We recorded initial deviations from the correct solution path and the latency to crossing the barrier. In the A trials, mules crossed the barrier significantly faster than their parental species, the horses and donkeys. In the B trials, following the change of gap location, all species showed a reduction in performance. Both dogs and horses exhibited significant spatial perseveration, going initially to the previous gap location. Donkeys and mules, however, performed at chance level. Our results suggest that hybrid vigour in mules extends to spatial abilities.

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: In the present study, we sought to determine the influence of stress and temperament on working memory for disappearing food in horses. After assessment of five dimensions of temperament, we tested working memory of horses using a delayed-response task requiring a choice between two food locations. Delays ranging from 0 to 20 s were tested. The duration of working memory for disappearing food was first characterized without stressors (N = 26). The horses were then divided into two groups and their performance was assessed under stressful (exposure to acute stressors prior to testing, N = 12) or control conditions (N = 12). Results showed that the duration of working memory for disappearing food lasted at least 20 s under nonstressful conditions, and that under stressful conditions this duration lasted less than 12 s. This stress-induced impairment confirms in a nonrodent species that working memory performance is very sensitive to exposure to stressors. In addition, working memory performance in horses is influenced by the temperamental dimension of fearfulness according to the state of stress: fearful horses showed better performance under control conditions and worse performance under stressful conditions than nonfearful horses. These findings are discussed in the context of the Yerkes–Dodson law of stress and performance.

Abstract: An ethogram of agonistic and related behaviors among equid bachelor band members was developed. Several key English-language studies on equids were reviewed to derive a preliminary inventory of specific behaviors to be included in the ethogram. A bachelor band of domestic pony stallions pastured together was observed for approximately 50 daylight hours to obtain detailed descriptions of each behavior, enable photographic and video documentation of behaviors, and identify any behaviors to be added to the preliminary inventory. An initial draft of the ethogram was sent to 65 equine researchers for review. Twenty-eight critical reviews were received and their suggestions considered for the final draft. A total of 49 elemental behaviors including five distinct vocalizations was included in the ethogram. Three complex behavioral sequences were also included. Most of the behaviors catalogued from the direct observation of pastured pony stallions were also found in the equid literature. For many, references to these behaviors specifically among males or bachelor band members were not found. The results offer a practical tool for quantitative research and other studies of equid inter-male behavior as well as for teaching of equid behavior, and should facilitate progress toward development of a complete ethogram for the horse and other equids.

Abstract: In contrast to horses in pastures, it is thought that free-ranging horses do not perform latrine behaviour, i.e. a behavioural pattern whereby the animals graze and defecate in separate areas. However, few studies deal with this particular subject, reporting contrasting conclusions. We hypothesize that horses free-ranging in large heterogeneous areas do not perform latrine behaviour. Thus, we believe that grazing and elimination behaviour are spatially related: where horses graze, they will also defecate. Behavioural data were collected from Konik horses, Haflinger horses, Shetland ponies and donkeys, grazing in different nature reserves (54-80 ha). Data for the different equids were analyzed separately, as well as data for mares and stallions (Konik and donkey stallions only). We investigated the proportion of the number of defecations/urinations while grazing on the total number of defecations/urinations; furthermore, we searched for the sequence of behaviours representing latrine behaviour in the strict sense. Additionally, we analyzed the correlation between grazing behaviour and eliminative behaviour on both vegetation type level and patch level. All the female equids often continued grazing while defecating. During urination, grazing ceases in the majority of instances. Cases where a mare terminated grazing in a certain vegetation type and sward height to eliminate in another vegetation type or in another sward height within the same vegetation type were rarely observed. On the vegetation type level as well as on the patch level, there was a highly significant (P<0.001) positive correlation between grazing time and number of eliminations (or eliminating time). The high values of the correlation coefficients (in case of the defecation variables r ranges between 0.553 and 0.955; in case of the urination variables r ranges between 0.370 and 0.839) illustrate that the spatial distribution of the eliminative behaviour can be explained to a high degree by the spatial distribution of the grazing behaviour. Results in the case of the stallions are preliminary, but indicate the same pattern. Horses, free-ranging in large heterogeneous areas, do not perform latrine behaviour, but defecate where they graze. Possibly, animal density is of major importance to explain this behavioural difference with horses in pastures. We suggest that also spatial vegetation heterogeneity and plant productivity of the grazed area, as well as parasite status of the grazing animals could play a role.