Abstract: Understanding the factors that influence horse learning is critical to ensure horse welfare and rider safety. In this study, data were obtained from horses (n=96) training to step backwards through a corridor in response to bit pressure. Following training, learning ability was determined by the latency to step backwards through the corridor when handled on the left and right reins. Additionally, horse owners were questioned about each horse's management, training, behavior and signalment (such as horse breed, age and sex). Factors from these four broad domains were examined using a multiple logistic regression (MLR) model, following an Information Theoretic approach, for associations between horses' behavioral attributes and their ability to learn the task. The MLR also included estimates of the rider's ability and experience as well as owner's perceptions of their horse's trainability and temperament. Results revealed several variables including explanatory variables that correlated significantly with rate of learning. Horses were faster at backing, a behavioral trait, when handled on the right (t = 3.65, df = 94, P < 0.001) than the left side. Thoroughbred horses were slower at completing the tests than other breeds of horses when handled on the left side (LM, F1,48=4.5, P=0.04) and right side (LM, F1,45=6.0, P=0.02). Those in regular work, a training factor, did not learn faster than their unworked counterparts on the right rein but completed the task faster on the left rein (F1,44=5.47, P=0.02). This may reflect differences in laterality and habituation effects. In contrast, more anxious horses were faster at completing the test when handled from the right (Spearman, r=-0.22, P=0.04). It is possible that these horses have an increased arousal level when interacting with handlers, resulting in more engagement with the lesson, accounting for the improved performance results. The findings of this study will help clarify how horse behavior, training and management may influence learning and how their application may optimize learning outcomes. Future equine behavior assessment and research questionnaires should include items that assess these qualities.

Abstract: This series of studies investigated horses' ability to learn the concept of sameness under several different conditions. Before experimentation began, three horses were shaped to touch individually presented stimuli with their muzzles, and then to make two responses to two matching cards from an array of three. A modified version of the identity matching-to-sample (IMTS) procedure was used to present stimuli in a variety of configural arrangements on a barn wall (Experiment 1 and Experiment 2), and on a flat panel mounted to a barn door (Experiment 3). The task in each experiment was to select the two stimulus cards that were the same (either circles or Xs) and to avoid the nonmatching stimulus card (either a star or a square). In Experiment 1, the mean accuracy rate for selecting the matching alternatives was 74%. The horses' accuracy levels reached a mean level of 83% during Experiment 2, in which they received additional trials and an intermittent secondary reinforcement schedule. In Experiment 3, when the stimuli were moved further apart from each other within arrangements and were presented on a novel background, the mean accuracy rate was 73%. These data demonstrate that horses can learn complex discrimination problems involving the concept of sameness, and that they are able to generalize this learning to a novel stimulus presentation situation. These results also suggest that a relational discrimination test may be useful for assessing horses' learning ability and the level of training appropriate for individual horses.

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: Viele Pferdebesitzer bevorzugen aus Angst vor aggressiven Interaktionen und Verletzungsgefahr der Tiere untereinander die Einzelhaltung, obwohl von Tierschutzorganisationen die Gruppenhaltung für Pferde empfohlen wird. In dieser Studie beobachteten wir während des alltäglichen Soziallebens als auch bei der Eingliederung von neuen Gruppenmitgliedern das Sozialverhalten, insbesondere das Aggressionsverhalten, von elf Gruppen domestizierter Pferde (Equus caballus) verschiedener Größe und Zusammensetzung. Während des alltäglichen Soziallebens hatten die Gruppe und der Paddock-Typ (Gras / kein Gras) keinen Einfluss auf die Verhaltensweisen, wohingegen die Paddockgröße unter 10000 m2 einen signifikanten Einfluss auf die submissiven Verhaltensweisen (GzLM; n=56; t=-2.061, P=0.044) und einen nicht signifikanten Einfluss auf die aggressiven Verhaltensweisen (GzLM; n=56; t=-1.782, P=0.081) hatte. Allerdings verringerten sich sowohl die aggressiven als auch die submissiven Verhaltensweisen mit steigendem Platzangebot bis zu 10000 m2 (Spearman rank Korrelation; n=56; aggressive Verhaltensweisen: r = -0.313, P = 0.019; submissive Verhaltensweisen: r = -0.328, P = 0.014). Während den Eingliederungen reduzierten sich die Aggressionen pro Stunde mit der Vergrößerung des Platzangebotes (Spearman rank Korrelation; n=28; r=-0.402, P=0.034). Dies zeigte sich noch deutlicher, wenn Beobachtungen mit einem Platzangebot von über 10000 m2 ausgeschlos- sen wurden (Spearman rank Korrelation; n=23; r=-0.549, P=0.007). Während des alltäglichen Soziallebens näherte sich der Aggressionslevel der Nulllinie an, wenn das Platzangebot pro Pferd mehr als 331 m2 betrug. Deshalb empfehlen wir zur Reduzierung des Aggressionslevels und des Verletzungsrisikos von sozial gehaltenen Pferdegruppen ein Platzangebot von mindestens 331 m2 pro Pferd.
[Even though animal welfare organisations propose group housing for horse welfare, many owners stable horses individually for fear of aggressive interactions and injury risks. In the present study we observed social behaviour, and especially aggressiveness, in eleven domestic horse groups (Equus caballus) of different size and composition, in basic social situations and when new group members were introduced. During basic social situations, the group and the type of paddock (grass / no grass) had no effect on any of the behaviours, where- as the enclosure size below 10,000 m2 had a significant effect on submissive behaviour (GzLM; n=56; t=-2.061, P=0.044) and an insignificant effect on aggressive behaviour (GzLM; n=56; t=-1.782, P=0.081). However, aggressive and submissive behaviour dimi- nished with the increase of enclosure sizes up to 10,000 m2 (Spearman rank correlation; n = 56; aggressive behaviour: r = -0.313, P=0.019; submissive behaviour: r=-0.328, P=0.014). During introductions, aggression levels per hour decreased with any increase of enclosure size (Spearman rank correlation; n=28; r=-0.402, P=0.034) and even more when enclosure sizes above 10,000 m2 were excluded (Spearman rank correlation; n=23; r=-0.549, P=0.007). During basic social situations the aggression level approached zero when the space allowance was more than 331 m2 per horse. We therefore recommend keeping horse groups in an enclosure with at least 331 m2 per horse to reduce aggression and injuries.]

Abstract: Management of grazed grasslands for production and/or conservation objectives requires a thorough understanding of the choices of feeding sites by herbivores, and of the biological processes involved. Most models of the feeding strategies of herbivores are based on the principle that optimising the intake of energy (or some nutrient) is the primary goal of foragers but other selective forces, such as parasitism, could be important. Gastrointestinal parasites (including cyathostome nematodes) have powerful effects on the fitness of herbivores and may act as a major selection pressure favouring host behaviour that reduces the risk of encountering parasites. Among large herbivores, horses have perhaps the most marked tendency to select particular feeding sites within grasslands. We test here: (1) whether horses select feeding patches with relatively low parasite densities and (2) if their choice is affected by their parasite load. We used 10 two-year old saddle-horses and three periods. In the first period, the horses were under natural parasitism which varied strongly among individuals; in the second period they were all dewormed, and in the third, a sub-set of the horses was experimentally infected with cyathostome larvae. Ninety-eight percent of the infective larvae in the pasture were found <1 m from faeces. The main determinant of the choice of feeding patch by horses was the availability of patches of different parasite risk and grass height. Controlling for availability, the horses used tall grasses (>16 cm) less than expected, whether the grass was contaminated or not, and they selected for short patches >1 m from faeces, where the risk of encountering parasites was low. These results suggest that selection of feeding sites by horses is driven by an interaction between their nutritional and anti-parasite strategies: the horses avoid the patches of tall grass which are generally of low quality and areas contaminated by parasite larvae which leads them to prefer the patches of short grass far from faeces. The parasite status of the horses at the time of the experiment had no effect on their feeding choices. However, before concluding that the challenge by cyathostomes has no effect on the selection of feeding sites in horses, it will be necessary to test whether the history of parasitism of the individuals, rather than the current status, is important.