Abstract: Equine welfare can be enhanced by applying species specific training. This may incorporate social learning, as horses are highly social and social stimuli are of primary importance. Social learning is comparable to individual learning in its learning mechanisms, differing primarily in the way it is stimulated. Our initial study showed that horses of different breeds (N = 38) follow humans after observing other horses doing so, but only if the observed horse was familiar to and higher ranking than the observer (Fisher's exact test: N = 12, P = 0.003). A second study showed that horses and ponies (N = 25) learned to pull a rope to open a feeding apparatus after observing demonstrations by conspecifics, again, only if the demonstrating horse was older and higher ranking than the observer (Fisher's combination test, N = 3, v2 = 27.71, p = 0.006). Our third approach showed that horses and ponies (N = 24) learned to press a switch to open a feeding apparatus after observing a familiar person (GzLM: N = 24, z = 2.33, P = 0.02). Most recently, we confronted horses and ponies (N = 50) with persons demonstrating different techniques for opening a feeding apparatus. In this study we investigated whether the horses would copy the demonstrators' techniques or apply their own. Here only some horses copied the technique, and most of the successful learners used their mouths irrespective of the demonstrators' postures (Chi Square Test: N = 40, df = 2, &#967;2 = 31.4, p < 0.001). In all the approaches social stimuli elicited learning processes in the test horses, while only a few individuals in the control groups mastered the tasks by individual learning. The following behaviour observed in the initial study may have been facilitated by a social stimuli (social facilitation), and the opening of the feed boxes in the subsequent studies appear to be mostly the result of enhancement (social enhancement). Some horses may have used the social stimuli at first and continued their learning process by individual trial and error. However, the horses were also selective in whom and some in how to copy. This may have been conditioned (socially conditioned) or the result of simple forms of reasoning on the reliability of the particular information provided by demonstrators of certain social ranks or social positions, as high ranking and familiar horses and familiar persons were copied and some imitated exactly.
Lay person message: Traditional riding instructions suggest that horses learn by observing other horses. For example, older, more experienced driving horses are used for initial training of young driving horses. We have shown that horses indeed use learning stimuli provided by other horse, as well as by humans. Horses readily accept stimuli observed in high ranking and familiar horses, and familiar persons. Such stimuli elicit learning processes which are comparable to individual learning. We suggest applying social learning whenever possible, as it is much faster and less stressful than individual learning, where learners experience negative outcomes in trial and error learning.

Abstract: The Eneolithic Botai culture of the Central Asian steppes provides the earliest archaeological evidence for horse husbandry, ~5,500 ya, but the exact nature of early horse domestication remains controversial. We generated 42 ancient horse genomes, including 20 from Botai. Compared to 46 published ancient and modern horse genomes, our data indicate that Przewalski's horses are the feral descendants of horses herded at Botai and not truly wild horses. All domestic horses dated from ~4,000 ya to present only show ~2.7% of Botai-related ancestry. This indicates that a massive genomic turnover underpins the expansion of the horse stock that gave rise to modern domesticates, which coincides with large-scale human population expansions during the Early Bronze Age.

Abstract: Stress responses play a key role in allowing animals to cope with change and challenge in the face of both environmental certainty and uncertainty. Measurement of glucocorticoid levels, key elements in the neuroendocrine stress axis, can give insight into an animal’s well-being and can aid understanding ecological and evolutionary processes as well as conservation and management issues. We give an overview of the four main biological samples that have been utilized [blood, saliva, excreta (feces and urine), and integumentary structures (hair and feathers)], their advantages and disadvantages for use with wildlife, and some of the background and pitfalls that users must consider in interpreting their results. The matrix of choice will depend on the nature of the study and of the species, on whether one is examining the impact of acute versus chronic stressors, and on the degree of invasiveness that is possible or desirable. In some cases, more than one matrix can be measured to achieve the same ends. All require a significant degree of expertise, sometimes in obtaining the sample and always in extracting and analyzing the glucocorticoid or its metabolites. Glucocorticoid measurement is proving to be a powerful integrator of environmental stressors and of an animal’s condition.

Abstract: Pair and social bonding has been documented in various taxa, where pair formations are often described as being driven by kinship or sexual motivation. However, pair-bonding between unrelated individuals where sexual motivation is not a factor is not well documented. Many social relationships and pair-bonds between members of a dyad are facilitated by each individual's ability to recognise their partner using cues which are characteristic of that particular individual. The aims of this study were i) to investigate the existence of pair-bonding in domestic donkeys and ii) to determine whether members of a dyad could recognise their companion during a Y-maze recognition test. Subjects were 55 unrelated donkeys (38 gelded males, 15 females) in seven groups of mixed or same sex, comprising 4?14 individuals. Spatial proximity (nearest-neighbour) was observed three times a day over a 22-day period. Using a simulation approach based on observed data to generate randomised nearest-neighbour matrices, the statistical significance of social relationships was estimated. Of these, 42 (79.2%) were involved in significantly (p<0.05) non-random nearest-neighbour relationships, most of which were reciprocal pair relationships. Based on the spatial data, 24 of the donkeys which had shown significant reciprocal nearest-neighbour preferences for one individual (companion) were then used in a Y-maze recognition test in which they were presented with a choice of their companion and either a familiar donkey from the same group or an unfamiliar donkey from a different group. Donkeys? spatial location in the Y-maze demonstrated a preference for their companion versus familiar (one sample Wilcoxon signed rank test, W=239, p=0.002) or unfamiliar donkeys (W=222, p=0.041). These results verify anecdotal evidence from donkey handlers that donkeys often form pair-bonds, and show that reciprocal social preference and recognition are the basis of these. Pair-bond formation and companionship among donkeys have potential implications for their management, husbandry and welfare.

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.