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: The aim of this study was to test the existence of a “reactivity-to-humans” trait which is stable over time and across situations. For this purpose, four test situations were repeated on the same animals at 8 months, 1.5 years (N = 110 horses) and 2.5 years of age (N = 54 horses). These situations involved a “familiar passive human test” during which a familiar person stayed motionless in the test pen, a “familiar active human test” and an “unfamiliar active human test” during which a familiar or unfamiliar person attempted to touch the horse, and a “halter-fitting and heart-rate measurement test”, during which the horse was haltered and equipped with a surcingle and then had its heart-rate measured. Many behavioural variables were recorded during the tests, and correlations between ages and variables measured during other situations were investigated. When correlations were found, we concluded that the variables corresponded to temperament traits which are stable over time and across situations. The horses tested were divided into four groups according to breed (Anglo-Arab and Welsh) and year of birth (2001 and 2002). Data for each group were analysed separately with Spearman rank correlations. Regarding stability over time, there was a strong correlation between ages in the frequency of licking/nibbling the passive human, the time taken by a familiar or unfamiliar person to touch the horse, and the time taken to fit a halter (e.g. the time taken to touch the horse's muzzle was significantly correlated between ages in 7 out of the 8 possible cases: 0.40 <= R <= 0.67). Regarding stability across situations, results indicate that the more frequently a horse sniffed, licked or nibbled a passive human, the easier it was for either a familiar or unfamiliar active human to touch and halter it. Moreover, a horse which could be touched easily by a familiar human could also be touched and haltered easily by an unfamiliar human. These behaviours, which are linked directly to humans, therefore indicate good stability over time and across situations. In conclusion, this stability suggests the existence of a “reactivity-to-humans” temperament trait, whether the human is passive or active, familiar or unfamiliar. When the animals do not have frequent contact with humans, this reactivity is stable over time and can be measured as early as 8 months of age using the behavioural variables cited above.

Abstract: Temperament is an important factor when working with horses. Behavioural tests have already been developed to measure certain dimensions of a horse's temperament (fearfulness, gregariousness, etc.). In order to measure the temperament more precisely, our work aimed to identify a dimension which has already been described in several species but not yet in horses, namely sensory sensitivity. Our study was based on the definition of a temperament dimension as “a behavioural characteristic stable across situations and over time”. We designed several tests for each sense and then determined whether the responses observed were correlated between situations and in time. The principle of the tests was to generate two stimuli of different intensities for each sense (e.g. two different sounds) and to measure the intensity of the horse's response (N = 26). Using Spearman rank correlations, we tested whether the responses to these different stimuli were inter-correlated. We repeated the same tests 5 months later to determine whether the responses were correlated over time. Within each sense, results show that the greater the horses' response to one stimulus, the greater their response to the other. For example, the reaction to the odour of cinnamon (time spent near the source of the odour) was significantly correlated to the reaction to lavender (R = 0.53, p = 0.004). The reactions to two different sounds or to two different tactile stimuli (von Frey filaments, or contact of a brush on the body), were also significantly correlated (R = 0.59, p < 0.0001; R = 0.38, p = 0.029). Finally, the reactions to two different tastes or to two visual stimuli tended to be correlated (R = 0.27, p = 0.09; R = 0.27, p = 0.09). However, there was no significant correlation between the responses to stimuli relating to different senses. Finally, except for the responses to odour, the responses to other sensory stimuli showed stability over a 5-month period (e.g. tactile stimulation: R = 0.71, p < 0.0001). In conclusion, our study revealed characteristics which were stable across situations and over time. The absence of links between the characteristics measured for the different senses suggests that a dimension for each sense exists (e.g. tactile sensitivity) rather than a general sensory sensitivity dimension covering all the senses.

Abstract: Selection pressures during domestication are thought to lead to an enhanced ability to use human-given cues. Horses fulfil a wide variety of roles for humans and have been domesticated for at least 5000 years but their ability to read human cues has not been widely studied. We tested the ability of 28 horses to attend to human-given cues in an object choice task. We included five different cues: distal sustained pointing, momentary tapping, marker placement, body orientation and gaze (head) alternation. Horses were able to use the pointing and marker placement cues spontaneously but not the tapping, body orientation and gaze alternation cues. The overall pattern of responding suggests that horses may use cues that provide stimulus enhancement at the time of choice and do not have an understanding of the communicative nature of the cues given. As such, their proficiency at this task appears to be inferior to that of domestic dogs, Canis lupus familiaris, but similar to that of domestic goats, Caprus hircus.

Abstract: The behavioral effects of the immunocontraceptive agent porcine zona pellucida (PZP) have not been adequately studied. Important managerial decisions for several species, including the wild horse (Equus caballus), have been based on this limited research. We studied 30 horses on Shackleford Banks, North Carolina, USA to determine the effects of PZP contraception on female fidelity to the harem male. We examined two classes of females: contracepts, recipients of the PZP vaccine (n = 22); and controls, females that have never received PZP (n = 8). We conducted the study during the non-breeding season from December 2005 to February 2006, totaling 102.2 h of observation. Contracepted mares changed groups more often than control mares (P = 0.04). Contracepts also visited more harem groups than did control mares (P = 0.02) and exhibited more reproductive interest (P = 0.05). For both contracepted and control females, the number of group changes (P = 0.01) and number of groups visited (P = 0.003) decreased with the proportion of years mares were pregnant. Our study shows that the application of PZP has significant consequences for the social behavior of Shackleford Banks horses. In gregarious species such as the horse, PZP application may disrupt social ties among individuals and inhibit normal social functioning at the population level.