Abstract: The behavior of feral populations of the African wild ass (Equus africanus) were studied in the Northern Panamint Range of Death Valley National Monument for 20 months from 1970 to 1973 [Moehlman, P.D., 1974. Behavior and ecology of feral asses (Equus asinus). PhD dissertation, University of Wisconsin, Madison, 251 pp.; Moehlman, P.D., 1979. Behavior and ecology of feral asses (Equus asinus). Natl. Geogr. Soc. Res. Reports, 1970: 405-411]. Maintenance behavior is described and behavior sequences that were used in social interactions are quantified by sex and age class. Agonistic, sexual, and greeting behavior patterns are described and analyzed in conjunction with the responses they elicited. Mutual grooming mainly occurred between adult males, and between females and their offspring. Five types of vocalizations were distinguished: brays, grunts, growls, snorts, and whuffles. A second population was studied for 1 month on Ossabaw Island, GA (Moehlman, 1979). This population had more permanent social groups and had a higher rate of mutual grooming and foal social play.

Abstract: Feral asses were studied on Volcano Alcedo, Galapagos Islands, Ecuador, during the wet season of 1980. On the volcano rim during March/April, two stable groups were observed to have a `female (harem) defense' polygynous mating system [Emlen, S.T., Oring, S.W., 1977. Ecology, sexual selection, and the evolution of mating systems. Science 197 (4300), pp. 215-223] and social behavior patterns and feeding ecology similar to feral asses living in a habitat where forage and climate are similar, e.g., Ossabaw Island, Georgia [Moehlman, P.D., 1979. Behavior and ecology of feral asses (Equus asinus). Nat. Geogr. Soc. Res. Rep., 1970, pp. 405-411; Moehlman, P.D., 1997. Feral asses (Equus africanus): intraspecific variation in social organization in arid and mesic habitats. J. Appl. Anim. Behav. Sci., this issue; McCort, W.D., 1980. The feral asses (Equus asinus) of Ossabaw Island, Georgia., PhD Dissertation, Pennsylvania State University, University Park, 219 pp.].

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.

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: Understanding how groups of individuals with different motives come to daily decisions about the exploitation of their environment is a key question in animal behaviour. While interindividual differences are often seen only as a threat to group cohesion, growing evidence shows that they may to some extent facilitate effective collective action. Recent studies suggest that personality differences influence how individuals are attracted to conspecifics and affect their behaviour as an initiator or a follower. However, most of the existing studies are limited to a few taxa, mainly social fish and arthropods. Horses are social herbivores that live in long-lasting groups and show identifiable personality differences between individuals. We studied a group of 38 individuals living in a 30-ha hilly pasture. Over 200 h, we sought to identify how far individual differences such as personality and affinity distribution affect the dynamic of their collective movements. First, we report that individuals distribute their relationships according to similar personality and hierarchical rank. This is the first study that demonstrates a positive assortment between unrelated individuals according to personality in a mammal species. Second, we measured individual propensity to initiate and found that bold individuals initiated more often than shy individuals. However, their success in terms of number of followers and joining duration did not depend on their individual characteristics. Moreover, joining process is influenced by social network, with preferred partners following each other and bolder individuals being located more often at the front of the movement. Our results illustrate the importance of taking into account interindividual behavioural differences in studies of social behaviours.