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.

Abstract: Abstract Scent marking is a common form of intraspecific communication in mammal species, and using faeces or urine is a cost-effective way of signalling competitive ability and resource holding power. Marking is ritually performed by male equids, and here we assess the function of male scent-marking behaviour in a recently introduced population of Przewalski horses Equus ferus przewalskii in Mongolia. Two forms of scent marking were observed: defecation on stud piles formed from repeated dunging in the same place, and overmarking of faeces and urine of mares. Stud piles were marked with dung by the harem holder and sniffed before and after dung was deposited. They were not found specifically at the periphery of harem ranges but occurred for the most part along routes used by the horses, and were more common in the core parts of harem ranges or where harem ranges overlapped. Thus, rather than being used to defend range boundaries, stud piles were placed predominantly where they would be encountered by male intruders. Mare excreta were covered with urine by the stallion, but were only sniffed before they were marked, not after. These marks appear to advertise to the mare and other, intruding stallions that the harem holder was the mare's consort and that the interloper should not risk trying to steal the mare or sneak a mating. Thus, the two forms of marking by harem holders appear to combine as first and second lines of defence of paternity rights in male intrasexual competition.

Abstract: The paper examines systematic relationships among primates between brain size (relative to body size) and differences in ecology and social system. Marked differences in relative brain size exist between families. These are correlated with inter-family differences in body size and home range size. Variation in comparative brain size within families is related to diet (folivores have comparatively smaller brains than frugivores), home range size and possibly also to breeding system. The adaptive significance of these relationships is discussed.

Abstract: Social learning is fundamental to social life across the animal kingdom, but we still know little about how natural selection has shaped social learning abilities on a proximate level. Sometimes, complex social learning phenomena can be entirely explained by Pavlovian processes that have little to do with the evolution of sociality. This implies that the ability to learn socially could be an exaptation, not an adaptation, to social life but not that social learning abilities have been left untouched by natural selection. I discuss new empirical evidence for associative learning in social information use, explain how natural selection might facilitate the associative learning process and discuss why such studies are changing the way that we think about social learning.