Abstract: Individual recognition is considered a complex process and, although it is believed to be widespread across animal taxa, the cognitive mechanisms underlying this ability are poorly understood. An essential feature of individual recognition in humans is that it is cross-modal, allowing the matching of current sensory cues to identity with stored information about that specific individual from other modalities. Here, we use a cross-modal expectancy violation paradigm to provide a clear and systematic demonstration of cross-modal individual recognition in a nonhuman animal: the domestic horse. Subjects watched a herd member being led past them before the individual went of view, and a call from that or a different associate was played from a loudspeaker positioned close to the point of disappearance. When horses were shown one associate and then the call of a different associate was played, they responded more quickly and looked significantly longer in the direction of the call than when the call matched the herd member just seen, an indication that the incongruent combination violated their expectations. Thus, horses appear to possess a cross-modal representation of known individuals containing unique auditory and visual/olfactory information. Our paradigm could provide a powerful way to study individual recognition across a wide range of species.

Abstract: Video playback has been used to explore many issues in animal communication, but the scope of this work has been constrained by the lack of stimulus-subject interaction. In many natural contexts, each participant's signalling behaviour is dependent from moment-to-moment on that of the other. Analyses of acoustic communication demonstrate the value of reproducing such social contingencies. We assessed the utility of interactive playback for studies of visual signalling by comparing the responses of male Jacky dragons, Amphibolurus muricatus, to interactive and non-interactive digital video playbacks of a life-sized conspecific. Displays produced by lizards in the interactive condition had the effect of suppressing the aggressive display of their simulated opponent. Each stimulus sequence generated during an interactive playback was subsequently played to a size-matched control animal. Males that could interact with the video stimulus responded principally with aggressive displays, while those that could not produced a mixture of aggressive and appeasement signals. Adding a degree of receiver responsiveness is hence sufficient to alter the type of signal evoked, even when video stimuli are physically identical. Interactive playback permits the experimental study of a broader range of theoretical topics and can enhance the realism of video stimuli.

Abstract: An ethogram of play behavior among equids was developed. Several key English-language studies on equids were reviewed to derive a preliminary inventory of specific behaviors to be included in the ethogram. Our primary observations were based on a herd of semi-feral Shetland-type ponies kept at New Bolton Center, University of Pennsylvania School of Veterinary Medicine, Kennett Square, PA. Greater than 100 h of direct observation and photo-documentation focused specifically on play in order to identify play behaviors to be added to the preliminary inventory and to obtain detailed descriptions of each behavior. Additionally, these observations were supplemented with photographs obtained during several years of observational study of this herd for other purposes, and with the cumulative equid observational experience and study notes of the principal investigator with other equid species. An initial draft was sent out to 18 equine behavior colleagues for review. A total of 38 individual behaviors classified into four distinct categories were included in the ethogram. These included object play (14 entries), play sexual behavior (3 entries), locomotor play (14 entries) and play fighting (7 entries). All of the behaviors catalogued from direct observation of the herd were also found in the equid literature. The resulting ethogram offers a practical tool as a field guide or reference for quantitative research and other studies of equid play behavior as well as for teaching of equid behavior.

Abstract: Under certain models of animal competition, individuals are expected to gather information about opponent quality in order to determine whether they should fight or withdraw. However, the ability to process complex information differs between individuals and across brain hemispheres: a feature of vertebrate cognition known as lateralization that is not anticipated by contest models. I investigated the relationship between aggressive behaviour and mating success during the fallow deer, Dama dama, rut and a measure of lateralization derived from eye preference during parallel walking. Results show that there was no relationship between the tendency to escalate to fighting or predictability in the tendency to engage in fighting and lateralization. Conversely, there was a quadratic relationship between third-party intervention behaviour and lateralization: the greater the tendency to intervene in ongoing fights the lower the degree of lateralization. However, individuals that showed lateralization for right-eye use were least likely to be targeted by the intervening male; thus lateralization is beneficial in this context because targeted males are highly likely to lose this subsequent encounter. The relationship between lateralization and mating success was also nonlinear: males that showed little evidence for an eye bias during lateral displays had the greatest mating success. Taken together, individuals that showed lateralization benefited from avoiding being targeted after third-party intervention; conversely, individuals that showed little evidence for lateralization actively intervened during ongoing fights and had higher mating success. These results suggest that, although lateralization does appear to confer a fitness advantage on individuals, this is not as extensive as anticipated.

Abstract: Some behaviours remain a mystery and their determinants are still uncertain. Here we propose a critical interpretation of two puzzling behaviours, i.e. yawning and adult play, recorded in large samples of domestic horses kept in sub-optimal conditions (e.g. time-restricted feeding practices, social isolation). Yawning is involved in behavioural state changes, especially in quiet contexts of motor relaxation, but seems also to be triggered by stress and emotional contexts. Thus, we investigated specific correlates of yawning and stereotypic behaviours in two large samples of working horses (n1 = 87, n2 = 59). A clear co-occurrences of yawning and stereotypic behaviours appeared: stereotypic horses yawned more than the non-stereotypic horses (Fisher test, p < 0.02), yawning increased at the same time periods as stereotypic behaviours did (Wilcoxon tests, p < 0.05 to 0.001) and yawning frequency was even sometimes positively correlated with stereotypic behaviour frequencies (Spearman correlations, p < 0.005). Play, more typical of young stages in healthy individuals, occurs rarely at adult stages but then more often in captive/domestic animals (which can be challenging to welfare) than in animals living in natural conditions. Thus, we tested the hypothesis that adult play may reflect altered welfare states in horses, where, as in several species, play rarely occurs at adult stages in natural conditions. We observed the behaviour (in particular social play) of 29 adult riding school horses during occasional outings in a paddock and measured several stress indicators (health-related, behavioural and physiological) when these horses were in their home individual boxes. Results revealed that the number of horses and rates of adult play appeared very high compared to field report data and that most stress indicators measured differed between “players” and “non-players”, revealing that most “playful” animals were more prone to suffer from chronic stress contrarily to “non-playful” horses (Fisher, Chi-square, Mann-Whitney and Spearman correlation tests, p < 0.05 to 0.001). Altogether, these results show that yawning and adult play can both reflect altered welfare states in horses, highlighting that these puzzling behaviours should not be considered, as currently thought, as reliable indicators of good welfare states and / or expression of positive emotions.