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.

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: Summary
Many animals are lateralized when using sensory organs such as the eyes, ears or nostrils. Sensory laterality is not, as previously believed, caused by adjustment to motor laterality, but rather by one sided information processing in the particular brain hemi-spheres. While the right hemisphere predominantly analyses emotional information, the left hemisphere governs controlled rational, cognitive decisions. Since the brain hemi-spheres are largely connected with contralateral sensory organs, it is possible to infer how the information may be being interpreted by the side of preferred eye, ear or nostril used. The left eye usually dominates in emotional situations, i.e. fear or positive ex-citement, and the right eye in rational situations. Moreover, laterality increases when animals are stressed, e.g. when animals are confronted with anthropogenic or natural factors they can not handle, such as unsuitable housing or training conditions or un-avoidable predation pressure and social competition. A strong or increasing laterality could therefore potentially indicate welfare issues.
Zusammenfassung
Viele Tiere zeigen eine eindeutige sensorische Lateralität, sprich sie benutzen bevor-zugt ein Auge, ein Ohr, oder eine Nüster zur Aufnahme von Sinneseindrücken. Dies korreliert in den meisten Fällen nicht mit der motorischen Lateralität, sondern wird viel mehr durch die einseitige Verarbeitung von Informationen in den jeweiligen Gehirnhe-misphären bedingt. So werden emotionale Reaktionen von der rechten, reaktiven Ge-hirnhemisphäre und rationale Reaktionen von der linken, kognitiven Gehirnhemisphäre gesteuert. Da die Gehirnhälften zum Großen Teil mit den kontrolateralen Sinnesorga-nen verbunden sind lässt die Seite mit welcher Sinneseindrücke aufgenommen werden Schlüsse auf deren Informationsgehalt zu. So zeigen Tiere bei linksseitiger Aufnahme von Sinneseindrücken vermehrt reaktive, emotionalen Reaktionen, wie etwa bei Angst oder freudige Erregung, und bei rechtsseitig aufgenommene Sinneseindrücke eher rationales, gesteuertes Verhalten. Zudem verstärkt sich die sensorische Lateralität wenn Tiere Stress erfahren, sprich wenn sie wiederholt mit Situationen anthropogenen oder natürlichen Ursprungs konfrontiert werden denen sie nicht gewachsen sind, wie etwa bei unpassenden Haltungs- und Trainingsbedingungen, oder bei unausweichli-chem Raubtierdruck und sozialer Konkurrenz. Eine stark ausgeprägte, zunehmende sensorische Lateralität kann daher auf ein beeinträchtigtes Wohlergehen der Tiere hinweisen.

Abstract: Current definitions of horse personality traits are rather vague, lacking clear, universally accepted guidelines for evaluation in performance tests. Therefore, the aim of the present study was to screen behavioural and physiological measurements taken during riding for potential links with scores the same horses received in the official stallion performance test for rideability and personality traits. Behaviour, heart rate (HR) and HR variability from thirty-six stallions participating in a performance test were recorded repeatedly during their performance test training. Using the coefficient of determination, regression analysis revealed that about 1/3 of variation (ranging between r = 0.26 (“constitution” (i.e. fitness, health)) and r = 0.46 (rideability)) in the personality trait scores could be explained by selecting the three most influential behaviour patterns per trait. These behaviour patterns included stumbling (with all traits except character), head-tossing (temperament, rideability), tail-swishing (willingness to work), involuntary change in gait (character) and the rider's use of her/his hands (constitution, rideability), voice (temperament) or whip (constitution). Subsequent mixed model analysis revealed a significant (P < 0.05) influence of the behaviour pattern “horse-induced change in gait” on character (-0.98 ± 0.31 scores per additional occurrence of change in gaits), of head-tossing (-0.25 ± 0.08 scores) and rider's use of voice (-0.51 ± 0.25; P = 0.0594) on temperament, and of stumbling on each of the following: willingness to work (-2.5 ± 1.2), constitution (-2.5 ± 1.2 scores; P = 0.0516) and rideability scores (-3.3 ± 1.4). In addition, constitution scores tended (P = 0.0889) to increase with higher low frequency/high frequency heart rate variation ratios (LF/HF), indicating a shift towards sympathetic dominance and thus a higher stress load in horses with higher scores for constitution. Rideability scores from the training phase were also significantly influenced by head-tossing (-0.5 ± 0.1), and in addition rideability scores from the final test were influenced by the training rider, ranging between average estimated rideability scores of 6.8 ± 0.4 for one training rider and 8.36 ± 0.3 scores for another training rider. Horses ridden with their nose-line predominantly behind the vertical received higher scores for rideability (8.3 ± 0.3) than horses ridden with their nose-line at the vertical (7.7 ± 0.2). These findings indicate that either judges perceive horses to have a better rideability when they readily offer a more extreme poll flexion, or that riders make use of horses’ better rideability by imposing a more extreme poll flexion. Several of the above described associations, but also of the non-existing links (e.g. no association between shying or heart rate and temperament) between behaviour patterns and scores for personality traits are rather surprising, warranting further investigation regarding the underlying causes of these relationships. Some of these behaviour patterns should be considered when redesigning the current guidelines for evaluation of personality traits during breeding horse performance tests, ultimately leading to improved genetic selection for equine personality traits. However, ethical implication of defining aversive behaviour such as head-tossing as an indicator of, for example, poor temperament, should not be neglected when devising new guidelines: such aversive behaviour may in fact be an indication of inadequate training techniques rather than poor horse personality.