Abstract: Play behaviour is widespread in mammals, but benefits to play have been difficult to demonstrate. Physical training is one of the many proposed hypotheses, suggesting that males and females should play differently, that increased maternal investment should lead to increases in play, and that increases in play should result in physical advantages. In a population of feral horses, Equus caballus, males and females did not differ in their play behaviour except that males initiated more of their play bouts than females. Maternal condition influenced play behaviour only in males, with sons of mothers in good condition playing more. However, when we controlled for maternal effects by comparing a son and a daughter of the same mother, daughters played more when their mother was in poor condition and sons played more when their mother was in good condition. Mothers of foals that played more lost more condition. Therefore, the difference in play behaviour could not be explained by offspring sex or maternal condition alone, but play behaviour mirrored variation in maternal investment. In addition, those individuals that played more survived better and had better body condition as yearlings despite weaning earlier. Since increased activity has been linked to enhanced musculoskeletal development in domestic horses, we suggest that play provides a link between increased maternal investment, increased body condition and future reproductive success in feral horses, and probably in other species.

Abstract: A study was made of the behavior of young male Misaki feral horses in the developmental stage, by observing nine of them once a week from January 1988 to December 1996. The relationship between behavior before separation and in the developmental stage was also investigated. This stage begins just after young males separate from their natal band or mothers, and it continues until they start to form harems. The duration of the developmental stage in the study ranged from 0.6 to 3.9 years, depending on the age of the young males at the time of separation. Young males associated with three types of social groups at the beginning of the developmental stage, according to their social groups before separation. These were bachelor groups, harem groups and wandering female groups. During this period, males joined the three groups, mixed sex groups and sometimes were solitary. It was considered that these associations provided a good opportunity for males to acquire different behavioral patterns and experiences before they entered the next stage. Depending on the groups with which they associated, young males that spent more time with bachelor groups had the longest average developmental stage. They associated with harem groups more often during the breeding season and more frequently with other groups or were solitary during the non-breeding season. This may be a transition period because by the end of this stage all males had spent time in solitude before forming their own harem bands.

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: Over a 16-year period (1979-1994), long-term investigations were carried out on 14 Misaki feral stallions to analyze changes in harem size and the reproductive success. Harem size changed with the age of the stallions. Most stallions formed harem groups with four to five mares at the age of 4-6 and then the number of mares increased rapidly to the maximum at the age of 6-9 years. Thereafter, harem size decreased gradually to a minimum with advancing age. The harem size of 60 stable harem groups ranged from 1 to 9, and the average varied from a minimum mean of 1.8 in 1988 to a maximum mean of 5.3 in 1982. Mean harem size increased as adult sex ratio increased and a significant and positive correlation was found between them. One hundred and ninety-eight sire-foal pairs were determined by a paternity test with blood types and consort relations between stallions and mares during the study period. Out of 99 foals which were born in the stable harem groups, the true sires of 84 foals (85%) were the harem stallions in which the foals were born but the remaining 15 foals (15%) were sired by other harem stallions. Two out of three stallions which were studied throughout their lifetime produced 24 and 25 foals in 10 and 11 years of their reproductive lifespan, respectively. Another one produced only five foals in 6 years. The number of foals sired by the harem stallions was less than two over harem size 7 and some of the foals born in the harem were sired by other harem stallions. These results suggest that if a particular stallion monopolizes too many mares, he could not sire so many offspring because he could not always prevent his rival stallions from mating with his mares in wild or feral circumstances.

Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.