Abstract: Horse domestication revolutionized transport, communications, and warfare in prehistory, yet the identification of early domestication processes has been problematic. Here, we present three independent lines of evidence demonstrating domestication in the Eneolithic Botai Culture of Kazakhstan, dating to about 3500 B.C.E. Metrical analysis of horse metacarpals shows that Botai horses resemble Bronze Age domestic horses rather than Paleolithic wild horses from the same region. Pathological characteristics indicate that some Botai horses were bridled, perhaps ridden. Organic residue analysis, using δ13C and δD values of fatty acids, reveals processing of mare's milk and carcass products in ceramics, indicating a developed domestic economy encompassing secondary products.

Abstract: Additional measures of well-being would be beneficial to the management of a variety of species in human care, including elephants. Immunoglobulin A (IgA) is an immune protein associated with pathogen defense, which has been demonstrated to decrease during times of stress, and increase in response to positive stimuli. This paper describes the development and validation of an enzyme immunoassay (EIA) for the quantification of Asian elephant (Elephas maximus) IgA in feces, saliva, urine, and serum. Samples were collected weekly from four females for 6 months to assess IgA and glucocorticoid (GC) concentrations, establish relationships between these two biomarkers, and determine variability in IgA within and between individuals, and across sample types. IgA was quantified in all four sample types, although urinary concentrations were low and sometimes undetectable in individual samples. Concentrations were highly variable within and between individuals, with fecal, salivary and serum IgA, and fecal, salivary and urinary GCs all differing significantly across individuals. Contrary to previous findings, IgA and GC were generally not correlated. Serum IgA was less variable within individuals, with the exception of one female that experienced a brief illness during the study. However, marked inter-individual differences were still apparent. When data from all individuals were combined, fecal IgA was significantly predicted by salivary and urinary IgA; however, this relationship did not hold when individuals were analyzed separately. Analysis of a fifth female that exhibited a more severe systemic illness demonstrated clear increases in fecal IgA and GC, suggesting these may also be useful health biomarkers. Further investigation is needed to determine what sample type is most reflective of biological state in elephants, and how IgA concentrations are associated with health and positive and negative welfare states. Based on observed variability, a longitudinal approach likely will be necessary to use IgA as a measure of well-being.

Abstract: In recent years, Przewalski’s horses have been increasingly kept in semi-reserves. However,there areonly few studies ontheir behaviour and their ability to adaptto management interventions.In the main part of my dissertation, I focus on investigatingthe animals’ behaviour in different semi-reserves with varyinghabitats and living spaces. In addition, I investigate the horses’ behaviour during various management interventionsand analysetheensuing changes instress levels. Another aspect of my dissertation is the studyof social behaviour inPrzewalski’s horses. I investigate theparameters that should be used to demonstrate social bonds between individualsandassess whichdata provide the most meaningful results.In the commentary tochapter 1,several studies investigatingsocial bonds in horsesare discussed. Comparing the various studies, it is strikingthat no homogeneous analyses orevaluation criteria exist. While some authors only considersocial grooming, others include data onthe spatial proximity of the individuals in their evaluations, and various definitionsof proximity can also be found in the literature. Additionally, someauthors use friendly approaches between individuals asa furtherparameter wheninvestigating the social bonds.Continuing with this theme, in chapter 2I investigate the social behaviour of the horses and compare various analysis methods. I show that proactive behaviour, such as friendly approaches, is a good alternative to spatial proximity when investigating social bonds between group members, andis also useful for expanding the often very small data sets of mutual grooming in horses. Comparing Przewalski’s horses with wild living horses, I found no significant differences in the social behavior and the frequency of social interactions, regardlessof group size, group composition, habitat, and individual parameters such as age and gender.Inchapter 3,I investigate the behaviour of a Przewalski’s horse group when exploring a new area of their enclosure. Their behaviour changed, showing less resting and more feeding. Furthermore, the animals maintained greater distances from each other, and the alpha male, instead of herding the group from behind, led the group around the new area and walked in front of the other group members. Moreover, he showed a substantial increase in stress level during the first day.A general comparison of the behaviour of the Przewalski’s horses in different semi-reserves is provided in chapter 4. In it, the habitat choice of the animals and their reactions to various management interventions are investigated. It is shown that Przewalski’s horses prefer open grassland to dense woods, although keeping Przewalski’s horses in a pine forest does not influence the animals’ stress level. In contrast to habitat, food range, and changes in the group composition, which do not appear to change stress levels, individual factors, such as the hierarchy, influence the glucocorticoid level of the animals significantly. The largest increases in stress hormones were demonstrated when the horses were temporarily confined in smaller areas.The importance of the available space is also discussed in chapter 5, where it is shown that horses show less aggressive behaviour when more space is provided. In contrast, the husbandry system does not influence the animals’ aggression, but the way of feeding can additionally reduce agonistic behaviour, for example if food is offered ad libitum.In summary, the results of this study provide indications for the optimization of keeping Przewalski’s horses in semi-reserves. The animals can adapt themselves to the environment and thrive in habitats which do not correspond to their original steppe-like home. Nevertheless, the semi-reserves should provide sufficient grassland, as the horses prefer this type of habitat. General speaking, any types of habitat can only offer a suitable living space if the food range is sufficient for the number of horses. Otherwise, and especially during could winter months, supplementary feeding is necessary according to the body condition of the animals. This is particularly important for older, weakened, or very young animals, which are still adapting to life in the semi-reserve. Without sufficient food, stress hormones can increase and negatively influence the well-being of the horses. The same is true for management interventions: restricting the animals to small enclosures, for example, can adversely affect the horses’ well-being and should be only done if absolutely necessary. Targetedbehaviour observations allow the animals that have a special meaning for the group to be identified, and these should not be taken out of the group unless it is unavoidable, as young and unexperienced horses orientate themselves on those animals. This is especially true for the alpha male in a bachelor group, as these groups are often composed of young horses and the alpha-male provides the necessary stability and experience. Social bonds between individuals can be investigated by observing friendly and proactive behaviour, and social grooming and friendly approaches yield suitable data for such analysis.

Abstract: Italian wolf howls are described for the first time from observations between 2003–2008 of a population living in eastern Tuscany, central Italy. A sample of 37 howls selected among single responses and 128 howls included in the choruses of 7 free ranging packs was recorded and analysed. The mean fundamental frequency of the howls ranged between 274–908 Hz. Two main structures recognised by means of multivariate explorative analysis, in particular Principal Component and Cluster Analysis, were ascribed to breaking and flat howls. Discriminant Function Analysis was applied to the recognised groups with the aim to find a general rule for classification. Howls with different features were correctly assigned to the groups obtained by explorative analysis in 95.8% of cases. The analysis of the variables characterising the structure of the howls suggests that maximum frequency and range of fundamental frequency are the most important parameters for classification, while duration does not appear to play any significant role.