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: Social bond analysis is of major importance for the evaluation of social relationships in group housed horses. However, in equine behaviour literature, studies on social bond analysis are inconsistent. Mutual grooming (horses standing side by side and gently nipping, nuzzling, or rubbing each other), affiliative approaches (horses approaching each other and staying within one body length), and measurements of spatial proximity (horses standing with body contact or within two horse-lengths) are commonly used. In the present study, we assessed which of the three parameters is most suitable for social bond analysis in horses, and whether social bonds are affected by individual and group factors. We observed social behaviour and spatial proximity in 145 feral horses, five groups of Przewalskiâ&#65533;&#65533;s horses (N = 36), and six groups of feral horses (N = 109) for 15 h per group, on three days within one week. We found grooming, friendly approaches, and spatial proximity to be robust parameters, as their correlation was affected only by the animalsâ&#65533;&#65533; sex (GLMM: N = 145, SE = 0.001, t = â&#65533;&#65533;2.7, p = 0.008) and the group size (GLMM: N = 145, SE < 0.001, t = 4.255, p < 0.001), but not by the horse breed, the aggression ratio, the social rank, the group, the group composition, and the individuals themselves. Our results show a trend for a correspondence between all three parameters (GLMM: N = 145, SE = 0.004, t = 1.95, p = 0.053), a strong correspondence between mutual grooming and friendly approaches (GLMM: N = 145, SE = 0.021, t = 3.922, p < 0.001), and a weak correspondence between mutual grooming and spatial proximity (GLMM: N = 145, SE = 0.04, t = 1.15, p = 0.25). We therefore suggest either using a combination of the proactive behaviour counts mutual grooming and friendly approaches, or using measurements of close spatial proximity, for the analysis of social bonds in horses within a limited time frame.

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.