Abstract: Single species aggregations are a commonly observed phenomenon. One potential explanation for these aggregations is provided by the selfish herd hypothesis, which states that aggregations result from individual efforts to reduce personnel predation risk at the expense of group-mates. Not all movement rules based on the selfish herd hypothesis are consistent with observed animal behavior. Previous work has shown that herd-like aggregations are not generated by movement rules limited to local interactions between nearest neighbors. Instead, rules generating realistic herds appear to require delocalized interactions. To date, it has been an open question whether or not the necessary delocalization can emerge from local interactions under natural selection. To address this question, we study an individual-based model with a single quantitative genetic trait that controls the influence of neighbors as a function of distance. The results indicate that predation-based selection can increase the influence of distant neighbors relative to near neighbors. Our results lend support for the idea that selfish herd behavior can arise from localized movement rules under natural selection.

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: Gegenstand und Ziel: Überprüfung der Tiergerechtheit von Abrufstationen
für Kraftfutter und Heu in der Gruppenhaltung von Pferden anhand
ethologischer und physiologischer Parameter im Vergleich zu konventionellen
Fressständen. Material und Methoden: Die Verhaltensbeobachtungen
fanden in 11 bzw. 10 Offenlaufställen mit Abrufstationen
bzw. Fressständen statt. 270 Pferde wurden individuell unterschieden.
Untersuchungsparameter waren: Drohverhalten mit und ohne Verletzungsrisiko,
Meideverhalten, Verdrängung aus dem Fressstand sowie
Herzfrequenz und Integumentverletzungen. In jedem Betrieb erfolgten
kontinuierliche Direktbeobachtungen für je einen 24-Stunden-Tag (Tortenstückverfahren).
Ergebnisse: Im Wartebereich der Abrufautomaten
traten signifikant mehr Drohgesten ohne bzw. mit Verletzungsgefahr je
Pferd und Tag (8,6 bzw. 3,0) auf als im Wartebereich der Fressstände
(3,4 bzw. 0,9). Demgegenüber konnten die Pferde in Abrufstationen
(1,4 Drohgesten je Pferd und Tag) ungestörter fressen als in Fressständen
(3,2 Drohgesten je Pferd und Tag). Insgesamt betrachtet ergab sich
jedoch bei beiden Fütterungssystemen eine geringe Anzahl an sozionegativen
Interaktionen im Bereich der Futtereinrichtungen. Die Herzfrequenz
lag im Warteareal im Durchschnitt im physiologischen Bereich
(44,59 ± 11,73 Schläge/min). Integumentverletzungen im Zusammenhang
mit dem Fütterungssystem traten nicht auf. Der wichtigste Einflussfaktor
auf die Untersuchungskriterien war der Betrieb (Fläche, Konzeption,
Management). Schlussfolgerung: Unter dem Aspekt der Tiergerechtheit
hinsichtlich des Stress- und Verletzungsrisikos durch soziale
Interaktionen eignen sich bei ordnungsgemäßer Gruppenhaltung mit
fachgerechtem Management sowohl Fressstände als auch Abrufstationen
für Pferde im Offenlaufstall.
[Objective: A comparison with regard to animal welfare of feeding
stalls and automatic feeding systems for hay and concentrates in group
housing systems for horses using parameters of ethology and physiology.
Material and method: The observations of animal behaviour took
place in 10 stables with feeding stalls and in 11 stables with automatic
feeding systems. The field around the feeding systems was divided into
three areas with comparable dimensions (waiting area, exit area and
inside of the feeding system). 270 horses were individually observed.
Parameters of research comprised: threatening behaviour with and
without risk of injury, avoiding behaviour, ”chasing away from the feeding
place”, as well as heart rate and injuries of the integument. Every
group of horses was continuously observed for 24 hours. This observation
took place on four different days and comprised six sessions, each
of 4 hours. Results: In the waiting area of the automatic systems there
were significantly more threatening gestures with and without risk of
injuries for each horse and day (8.6 and 3.0, respectively) than in the
waiting area of the feeding stalls. In contrast, the horses could eat more
relaxed in the automatic systems (1.4 threatening gestures for each
horse and day) than in the feeding stalls (3.2 threatening gestures for
each horse and day). All together the number of negative interactions in
the feeding area of both feeding systems was relatively low. The heart
rate was within the physiological range (44,59 ± 11,73 beats/min) in
the waiting area. There were no injuries of the integument in correlation
with the feeding systems. The most important factor of the observation
criteria was the individual group housing system with its different dimension,
conception and management. Conclusion: Under the aspect
of animal welfare both feeding systems are suitable for horses with respect
to the risk of stress and injuries by social interactions and under
the condition of proper group housing with professional management.]

Abstract: Domestic horses are kept in so-called “fate societies” where they have to deal with frequent mixing. Several studies have evaluated and discussed the aggression level and injury risk during the introduction of horses into new groups, but nothing is known about the endocrine responses and thus if horses experience stress during introduction.
In this study we analysed the efficiency of four approved introduction techniques and evaluated the introduction of 30 horses into 11 different groups. Horses were introduced: 1) immediately, 2) after observing the new group for several days, 3) together with an “integration horse” after several days of observation, or 4) with a mixed strategy. Aggressive as well as positive social behaviour between the introduced horses and the group members were analysed the two hours following the introduction event. In addition, we focussed on the glucocorticoid production of the newcomer horses by measuring faecal cortisol metabolites (FCM) on the day of the introduction as well as the following three days.
For the four introduction techniques we found significant differences in the horses’ aggressive and submissive behaviour as well as in their total interactions. The introduction together with an integration horse led to significantly lower levels of aggression and less total interactions than the immediate introduction of single horses.
Horses which were introduced immediately or after an observation period showed significantly elevated levels of FCM on the first, second and third day after the introduction. For horses introduced together with an integration horse FCM were already significantly higher on the day of the introduction, indicating a stressful event before the introduction itself. In contrast, FCM levels were always very low when using the mixed technique.
In sum, horses have the ability to deal with conflict when they are introduced to new group members. The introduction event itself appears not to be as stressful as previously assumed. Standing together with an “integration horse” on a separate paddock and not being able to integrate immediately into a new group appears to be stressful for the newcomer. Based on the findings of our study we suggest to introduce new horses in group management together with a new group mate, a so-called “integration horse”. This would reduce the number of total social interactions as well as the aggression level. While this technique may be stressful for the newcomer, it lowers aggressive behaviour between the introduced horse and the group members and consequently reduces injury risks.

Abstract: Revealing risk factors of abnormal stereotypic behaviour (ASB) in horses can help in the design of protective measures. Previous epidemiological studies indicate that social isolation, housing, management conditions, and feeding regime have a strong effect on developing ASB. The common belief that exposure to a stereotypic horse increases the risk of ASB has never been substantiated. Here we report that a generalised linear mixed models (GLMM) analysis of data on 287 horses of nine riding schools revealed that exposure to a stereotypic neighbour is a significant risk factor for performing stereotypy. Also, aggressive behaviour towards other horses increased the odds of stereotypy in the aggressor. These correspondences are unlikely to be a riding-school effect, because riding schools were treated as random factor in the GLMM. Risk factors identified by epidemiological studies cannot be treated as causal agents without independent evidence. Our aim in presenting these findings was to draw attention to the possibility of neighbour effects so that other researchers would include this variable in their surveys.