Abstract: Group living commonly helps organisms face challenging environmental conditions. Although a known phenomenon in humans, recent findings suggest that a benefit of group living in animals generally might be increased innovative problem-solving efficiency. This benefit has never been demonstrated in a natural context, however, and the mechanisms underlying improved efficiency are largely unknown. We examined the problem-solving performance of great and blue tits at automated devices and found that efficiency increased with flock size. This relationship held when restricting the analysis to naive individuals, demonstrating that larger groups increased innovation efficiency. In addition to this effect of naive flock size, the presence of at least one experienced bird increased the frequency of solving, and larger flocks were more likely to contain experienced birds. These findings provide empirical evidence for the “pool of competence” hypothesis in nonhuman animals. The probability of success also differed consistently between individuals, a necessary condition for the pool of competence hypothesis. Solvers had a higher probability of success when foraging with a larger number of companions and when using devices located near rather than further from protective tree cover, suggesting a role for reduced predation risk on problem-solving efficiency. In contrast to traditional group living theory, individuals joining larger flocks benefited from a higher seed intake, suggesting that group living facilitated exploitation of a novel food source through improved problem-solving efficiency. Together our results suggest that both ecological and social factors, through reduced predation risk and increased pool of competence, mediate innovation in natural populations.

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: Horses are one-sided, not only on a motor level, but they also prefer to use one eye, ear or nostril over the other under particular circumstances. Horses usually prefer using the left eye to observe novel objects and humans. This preference is more marked in emotional situations and when confronted with unknown persons. Thus the horse’s visual laterality provides a good option for assessing its mental state during training or in human-horse interactions. A strong preference for the left eye may signal that a horse cannot deal with certain training situations or is emotionally affected by a particular person.
Pferde benutzen für die Begutachtung von Objekten und Menschen bevorzugt eine bestimmte Nüster, ein Ohr oder ein Auge. So betrachten die meisten Pferde Objekte und Menschen mit dem linken Auge. Die Lateralitätsforschung erklärt diese sensorische Lateralität mit der Verarbeitung von Informationen unterschiedlicher Qualität in verschiedenen Gehirnhälften und zeigt auf, dass positive und negative emotionale Informationen sowie soziale Sachverhalte mit dem linken Auge aufgenommen und vorwiegend an die rechte Gehirnhälfte weitergegeben werden. In diesem Zusammenhang ermöglicht die visuelle Lateralität, den Gemütszustand des Pferdes im Training und im therapeutischen Fördereinsatz zu erkennen und zu berücksichtigen.

Abstract: In apes, four criteria are set to explore referential and intentional communication: (1) successive visual orienting between a partner and distant targets, (2) the presence of apparent attention-getting behaviours, (3) the requirement of an audience to exhibit the behaviours, and (4) the influence of the direction of attention of an observer on the behaviours. The present study aimed at identifying these criteria in behaviours used by dogs in communicative episodes with their owner when their toy is out of reach, i.e. gaze at a hidden target or at the owner, gaze alternation between a hidden target and the owner, vocalisations and contacts. In this study, an additional variable was analysed: the position of the dog in relation to the location of the target. Dogs witnessed the hiding of a favourite toy, in a place where they could not get access to. We analysed how dogs engaged in communicative deictic behaviours in the presence of their owner; four heights of the target were tested. To control for the motivational effects of the toy on the dogs’ behaviour and for the referential nature of the behaviours, observations were staged where only the toy or only the owner was present, for one of the four heights. The results show that gazing at the container and gaze alternation were used as functionally referential and intentional communicative behaviours. Behavioural patterns of dog position, the new variable, fulfilled the operational criteria for functionally referential behaviour and a subset of operational criteria for intentional communication: the dogs used their own position as a local enhancement signal. Finally, our results suggest that the dogs gazed at their owner at optimal locations in the experimental area, with respect to the target height and their owner’s (or their own) line of gaze.

Abstract: In recent years, considerable literature has been published on cognition in horses; however, much less is known about the cognitive abilities of domestic donkey (Equus asinus). This study aimed to expand our knowledge of donkey cognition by assessing their short-term memory capacity. We employed a detour problem combined with the classic delayed-response task, which has been extensively used to compare working memory duration in a variety of different species. A two-point choice apparatus was used to investigate location recall and search behaviour for a food target, after a short delay following its disappearance. Four donkeys completed the task with a 10 sec delay, while four others were tested with a 30 sec delay. Overall, each group performed above chance level on the test, showing that subjects had successfully encoded, maintained, and retrieved the existence and location of the target despite the loss of visual contact.