Abstract: Prima facie, the acquisition of novel behaviors in animals through observation of conspecifics seems straightforward. There are, however, various mechanisms through which the behavior of animals can be altered from observing others. These mechanisms range from simple hard-wired contagious processes to genuine learning by observation, which differ fundamentally in cognitive complexity. They range from social facilitation and local enhancement to true social learning. The different learning mechanisms are the subject of this review, largely because research on learning by observation can be confounded by difficulties in interpretation owing to the looming possibility of associative learning infecting experimental results. While it is often assumed that horses are capable of acquiring new behavior through intra-species observation, research on social learning in horses includes a variety of studies some of which may overestimate the possession of higher mental abilities. Assuming such abilities in their absence can have welfare implications, e.g. isolating stereotypical horses on the assumption that these behaviors can be learned though observation by neighboring horses. This review summarizes the definitions and criteria for the various types of social transmission and social learning and reviews the current documentation for each type in horses with the aim of clarifying whether horses possess the ability to learn through true social learning. As social ungulates, horses evolved in open landscapes, exposed to predators and grazing most of the day. Being in close proximity to conspecifics may theoretically offer an opportunity to learn socially, however anti-predator vigilance and locating forage may not require the neural complexity of social learning. Given the significant energetic expense of brain tissue, it is likely that social facilitation and local enhancement may have been sufficient in the adaptation of equids to their niche. As a consequence, social learning abilities may be maladaptive in horses. Collectively, the review proposes a novel differentiation between social transmission (social facilitation, local and stimulus enhancement) and social learning (goal emulation, imitation). Horses are undoubtedly sensitive to intra-species transfer of information but this transfer does not appear to satisfy the criteria for social learning, and thus there is no solid evidence for true social learning in horses.

Abstract: A human-animal relationship can be developed through subsequent interactions, affected by the positive or negative emotional valence of the proceeding one. Horses implement a process of categorization to classify humans with whom they interact as positive, negative, or neutral stimuli by evaluating the kind of approach and the nature of the contact. In these terms, human-animal interactions are emotionally charged events, eliciting specific emotional states in both subjects involved. Although the human-horse relationship has been mainly investigated through behavioral analysis, physiological indicators are needed for a more objective assessment of the emotional responses. Heart rate variability (HRV) is a commonly used autonomic nervous system (ANS) correlate estimating the sympathovagal balance as a psychophysiological marker of emotion regulation in horses. We have assumed that long-term positive relationships with humans may have a positive and immediate impact on the emotional arousal of the horse, detectable, via ANS activity, during the interaction. We analyzed horses' heartbeat dynamics during their interaction with either familiar or unfamiliar handlers, applying a standardized experimental protocol consisting of three different conditions shifting from the absence of interaction to physical contact. The ANS signals were monitored through an innovative non-invasive wearable system, not interfering with the unconscious emotional response of the animal. We demonstrated that horses appeared to feel more relaxed while physically interacting (e.g., grooming on the right side) with some familiar handlers compared to the same task performed by someone unfamiliar. The shift of the sympathovagal balance toward a vagal predominance suggests that the horses experienced a decrease in stress response as a function not only of the handler's familiarity but also of the type of interaction they are experiencing. These results constitute the objective evidence of horses' capacity to individually recognize a familiar person, adding the crucial role of familiarity with the handler as a paramount component of human-animal interaction. Our rigorous methodological approach may provide a significant contribution to various fields such as animal welfare while further investigating the emotional side of the human-animal relationships.

Abstract: Domestic animals are highly capable of detecting human cues, while wild relatives tend to perform less well (e.g. responding to pointing gestures). It is suggested that domestication may have led to the development of such cognitive skills. Here, we hypothesized that because domestic animals are so attentive and dependant to humans' actions for resources, the counter effect may be a decline of self sufficiency, such as individual task solving. Here we show a negative correlation between the performance in a learning task (opening a chest) and the interest shown by horses towards humans, despite high motivation expressed by investigative behaviours directed at the chest. If human-directed attention reflects the development of particular skills in domestic animals, this is to our knowledge the first study highlighting a link between human-directed behaviours and impaired individual solving task skills (ability to solve a task by themselves) in horses.

Abstract: Generalist predators are expected to shape their diets according to the local availability of prey species. In turn, the extent of consumption of a prey would be influenced by the number of alternative prey species. We have tested this prediction by considering the wild boar and the grey wolf: two widespread species whose distribution ranges overlap largely in Southern Europe, e.g. in Italy. We have reviewed 16 studies from a total of 21 study areas, to assess whether the absolute frequency of occurrence of wild boar in the wolf diet was influenced by (i) occurrence of the other ungulate species in diet and (ii) the number of available ungulate species. Wild boar turned out to be the main prey of the wolf (49% occurrence, on average), followed by roe deer (24%) and livestock (18%). Occurrence of wild boar in the wolf diet decreased with increasing usage of roe deer, livestock, and to a lower extent, chamois and red deer. The number of prey species did not influence the occurrence of wild boar in the wolf diet. The wild boar is a gregarious, noisy and often locally abundant ungulate, thus easily detectable, to a predator. In turn, the extent of predation on this ungulate may not be influenced so much by the availability of other potential prey. Heavy artificial reductions of wild boar numbers, e.g. through numerical control, may concentrate predation by wolves on alternative prey (e.g. roe deer) and/or livestock, thus increasing conflicts with human activities.