Abstract: Observational field studies were conducted on two remote populations of feral horses in Australia to determine whether lateralization is a characteristic of Equus caballus as a species or results from handling by humans. Group 1 had been feral for two to five generations and Group 2 for 10–20 generations. In both groups, left-side biases were present during agonistic interactions and in reactivity and vigilance. Therefore, as in other vertebrates, the right hemisphere appears to be specialized to control agonistic behaviour and responses to potential threats. The leftwards bias was stronger in measures of behaviour involving more aggression and reactivity. Preferences to place one forelimb in front of the other during grazing were also determined. No population bias of forelimb preference was found, suggesting that such limb preferences present in domestic horses may be entrained. Since stronger individual limb preferences were found in immature than in adult feral horses, limb preference may be modified by maturation or experience in the natural habitat. Stronger limb preference was associated significantly with elevated attention to the environment but only in younger feral horses. No sex differences in lateralization were found. The findings are evidence that horses show visual lateralization, as in other vertebrates, not dependent on handling by humans. Limb preference during grazing, by contrast, does appear to depend on experience.

Abstract: Learning performances are influenced by many factors, not only breed, age and sex, but also temperament. The purpose of this study was to understand how different temperamental dimensions affect the learning performance of horses, Equus caballus. First, we carried out a series of behavioural tests on 36 Welsh ponies aged 5-7 years to measure five temperamental dimensions: fearfulness (novel area test and surprise test), gregariousness (social isolation test), reactivity to humans (passive human test), tactile sensitivity (von Frey filament test) and activity level (evaluation of locomotor activity during all the tests). We then presented them with two learning tasks (avoidance and backwards-forwards tasks). In the avoidance task they had to learn to jump over a fence when they heard a sound associated with an aversive stimulus (puff of air). In the backwards-forwards task they had to walk forwards or move backwards in response to a tactile or vocal command to obtain a food reward. There was no correlation between performances on the two learning tasks, indicating that learning ability is task-dependent. However, correlations were found between temperamental data and learning performance (Spearman correlations). The ponies that performed the avoidance task best were the most fearful and the most active ones. For instance, the number of trials required to perform 5 consecutive correct responses (learning criterion) was correlated with the variables aimed at measuring fearfulness (way of crossing a novel area: rs = -0.41, P = 0.01 and time to start eating again after a surprise effect: rs = -0.33, P = 0.05) and activity level (frequency of trotting during all the tests: rs = -0.40, P = 0.02). The animals that performed the backwards-forwards task best were the ones that were the least fearful and the most sensitive. For instance, the learning criterion (corresponding to the number of trials taken to achieve five consecutive correct responses) was correlated with the variables aimed at measuring fearfulness (latency to put one foot on the area: rs = 0.43, P = 0.01; way of crossing a novel area: rs = 0.31, P = 0.06; and time to start eating again after a surprise effect: rs = 0.43, P = 0.009) and tactile sensitivity (response to von Frey filaments: rs = -0.44, P = 0.008). This study revealed significant links between temperament and learning abilities that are highly task-dependent.

Abstract: Knowledge about social recognition and memory in animals can help us to determine appropriate management and husbandry techniques. In this study, we used a habituation–discrimination procedure to investigate the ability of horses (Equus caballus) to distinguish between the body odour samples of unfamiliar conspecifics. To pick up body odour, we rubbed material on the coat of horses and presented these unknown body odours to 16 different conspecifics of the same sex and similar age. The test consisted of two successive two-min presentations of a sample from one individual (e.g. individual ‘A’) and a simultaneous presentation of samples from individual ‘A’ and a novel individual (e.g. individual ‘B’) during a final third presentation. The results showed that horses, regardless of sex, decreased the time they spent investigating conspecific body odour across the initial two presentations—demonstrating habituation. In the final presentation, the results demonstrated successful discrimination of the previously experienced odour because horses investigated the novel olfactory sample (‘B’) significantly more than the pre-exposed sample (‘A’). Taken together, these findings suggest, for the first time, that horses are able to discriminate two stimuli derived from body odours of unfamiliar conspecifics over short period of time.

Abstract: Contraception is often considered for population control of wildlife that is otherwise protected from more traditional management strategies, though little is known about the behavioral consequences of contraception in wild populations of socially complex animals. Feral horses, Equus caballus, in the western United States are ideal candidates for contraceptive management due to broad scale federal protection, reported herd growth rates of 15-25% per year, and finite public land allocated for them. We investigated influences of the immunocontraceptive porcine zona pellucida (PZP) on social behavior within bands of feral horses in three discrete populations for 4 years. Each band consisted of a single stallion and a harem of adult females (>1 years old) and their foals, as well as associated yearling males and females. Four mutually exclusive behaviors (herding, reproductive, harem tending, and agonism) were analyzed to investigate the difference between behaviors initiated by band stallions toward control mares and contracepted mares within the bands. Additionally, spatial relationships between each stallion and each harem female were analyzed to assess possible passive interactions. A candidate set of 22 hierarchical mixed effects models, using the discrete populations as a random effect on various fixed effects, was analyzed by restricted maximum likelihood estimation. The most supported models were selected by corrected Akaike"s Information Criteria (AICc). Analyses were done on 3 female age cohorts based on distinct fecundity rates: 2 to 5 year olds, 6 to 14 year olds, and > 14 year olds. Stallions herded control mares in the 6 to 14 year cohort more than contracepted mares (n=128, P=0.037) with treatment being the most supported effect, but foal presence (dependent foal) also contributed significantly to the model. Contracepted mares received more reproductive behaviors than control mares in the 6 to 14 year cohort (n = 151, P=0.020). No differences were detected in herding or reproductive behaviors in
the least fecund groups, the 2 to 5 year olds and &#8805; 15 year olds. The only independent variable in the most supported model for reproductive behavior was treatment, and the covariates of foal presence, band residence (resident or transient female), band size, and body condition did not contribute. There were no differences in any age cohort for harem tending or agonism. Spatially, stallions maintained closer relationships with 2 to 5 year old contracepted females than with the same age control females (n=136, P < 0.001) while the group was feeding (at its most dispersed structure), but there were no differences while the band was resting or in locomotion. There were no spatial differences detected in the other age cohorts. Given the polyestrous nature and high fecundity of feral horses, the observed difference in reproductive behaviors between treatment groups was not surprising; however, the difference detected in herding rates was an unexpected behavioral modification. This change in herding behavior suggests that further investigation is needed to determine if PZP immunocontraception has an underlying influence on mare social rank, band structure, and band stability, as well as the scope and long-term importance of these behavioral dynamics.

Abstract: In the present study, we sought to determine the influence of stress and temperament on working memory for disappearing food in horses. After assessment of five dimensions of temperament, we tested working memory of horses using a delayed-response task requiring a choice between two food locations. Delays ranging from 0 to 20 s were tested. The duration of working memory for disappearing food was first characterized without stressors (N = 26). The horses were then divided into two groups and their performance was assessed under stressful (exposure to acute stressors prior to testing, N = 12) or control conditions (N = 12). Results showed that the duration of working memory for disappearing food lasted at least 20 s under nonstressful conditions, and that under stressful conditions this duration lasted less than 12 s. This stress-induced impairment confirms in a nonrodent species that working memory performance is very sensitive to exposure to stressors. In addition, working memory performance in horses is influenced by the temperamental dimension of fearfulness according to the state of stress: fearful horses showed better performance under control conditions and worse performance under stressful conditions than nonfearful horses. These findings are discussed in the context of the Yerkes–Dodson law of stress and performance.