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: Few studies have examined whether non-human tool-users understand the properties that are relevant for a tool's function. We tested cotton-top tamarins (Saguinus oedipus) and rhesus macaques (Macaca mulatta) on an expectancy violation procedure designed to assess whether these species make distinctions between the functionally relevant and irrelevant features of a tool. Subjects watched an experimenter use a tool to push a grape down a ramp, and then were presented with different displays in which the features of the original tool (shape, color, orientation) were selectively varied. Results indicated that both species looked longer when a newly shaped stick acted on the grape than when a newly colored stick performed the same action, suggesting that both species perceive shape as a more salient transformation than color. In contrast, tamarins, but not rhesus, attended to changes in the tool's orientation. We propose that some non-human primates begin with a predisposition to attend to a tool's shape and, with sufficient experience, develop a more sophisticated understanding of the features that are functionally relevant to tools.

Abstract: Generalizations about neural function are often drawn from non-human animal models to human cognition, however, the assumption of cross-species conservation may sometimes be invalid. Humans may use different strategies mediated by alternative structures, or similar structures may operate differently within the context of the human brain. The transitive inference problem, considered a hallmark of logical reasoning, can be solved by non-human species via associative learning rather than logic. We tested whether humans use similar strategies to other species for transitive inference. Results are crucial for evaluating the validity of widely accepted assumptions of similar neural substrates underlying performance in humans and other animals. Here we show that successful transitive inference in humans is unrelated to use of associative learning strategies and is associated with ability to report the hierarchical relationship among stimuli. Our work stipulates that cross-species generalizations must be interpreted cautiously, since performance on the same task may be mediated by different strategies and/or neural systems.

Abstract: Grooming networks among adult female monkeys exhibit two similar features across a number of different species. High-ranking animals receive more grooming than others, and the majority of grooming occurs between females of adjacent rank. A theoretical model which duplicates these features is presented, and the properties of the model are used to explain the possible causation and function of female grooming behaviour. The model illustrates how relatively simple principles governing the behaviour of individuals may be used to explain more complex aspects of the social structure of non-human primate groups.

Abstract: Hawk-dove games have been extensively used to predict the conditions under which group-living animals should defend their resources against potential usurpers. Typically, game-theoretic models on aggression consider that resource defense may entail energetic and injury costs. However, intruders may also take advantage of owners who are busy fighting to sneak access to unguarded resources, imposing thereby an additional cost on the use of the escalated hawk strategy. In this article we modify the two-strategy hawk-dove game into a three-strategy hawk-dove-sneaker game that incorporates a distraction-sneaking tactic, allowing us to explore its consequences on the expected level of aggression within groups. Our model predicts a lower proportion of hawks and hence lower frequencies of aggressive interactions within groups than do previous two-strategy hawk-dove games. The extent to which distraction sneakers decrease the frequency of aggression within groups, however, depends on whether they search only for opportunities to join resources uncovered by other group members or for both unchallenged resources and opportunities to usurp.