Abstract: Primates give alarm calls in response to the presence of predators. In some species, such as the Thomas langur (Presbytis thomasi), males only emit alarm calls if there is an audience. An unanswered question is whether the audience's behaviour influences how long the male will continue his alarm calling. We tested three hypotheses that might explain the alarm calling duration of male Thomas langurs: the fatigue, group size and group member behaviour hypotheses. Fatigue and group size did not influence male alarm calling duration. We found that males only ceased calling shortly after all individuals in his group had given at least one alarm call. This shows that males keep track of and thus remember which group members have called.

Abstract: We investigated dyadic grooming relationships in a captive group of bonobos (Pan paniscus) and questioned what social function grooming fulfils in the 'market of services and favors'. Hereto we examined which of two theoretical models – grooming for support (Seyfarth, 1977, 1980) or grooming according to the similarity principle (de Waal & Luttrell, 1986) – best accounted for the observed grooming distribution. Similarity in traits did not correlate with increased grooming or close proximity among the individuals. Therefore, the similarity hypothesis was rejected. Seyfarth's model of rank-related grooming was largely confirmed. The animals distributed their grooming according to the rank of the receivers. We found an exchange between grooming and receipt of support. There was more grooming up than down the hierarchy. However, not all predictions about rank-related competition over grooming were confirmed. We found that dyadic grooming reciprocity indeed increased with decreasing rank distance. Yet, there was no increase of grooming within the dyad with decreasing rank distance and high ranking individuals were not competed over at the highest rates. The observed correlation between grooming and support received represents an important fit with Seyfarth's prediction, but does not allow for conclusions about underlying causal processes. Other causal explanations, besides the 'groom to receive support' hypothesis, that could explain a similar correlation are discussed.

Abstract: Social dynamics and maintenance of social cohesion were studied by analysing social interventions in two groups of horses consisting of adult mares, their offspring, adult geldings and sub-adults. The animals were observed for a total of 1316 h. All relevant dyadic and triadic social interactions, including initial behaviour, possible intervention and outcome, were recorded. The main question was: do horses use interventions in affiliative interactions to safeguard their social network? Horses were significantly more likely to intervene in allogrooming or play interactions that involved a preferred partner. The stronger the preferred association in allogrooming, the higher the likelihood the intervener took over allogrooming with an initial dyad member. Interveners originating from two newly introduced groups (n = 3 and 5), intervened significantly more often when a member of their own group allogroomed with an unfamiliar horse. In play, no correlation with unfamiliarity was found. Overall, the intervening horses stopped more than half of the initial allogrooming interactions, and a third of all interactions. Therefore, social facilitation cannot sufficiently explain interference behaviour. This study shows that maintaining relationships with preferred partners is important to horses and has implications for equine husbandry and management.

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: Biological market models explain variability in reciprocity and interchange between groups. In groups with a shallow dominance gradient, grooming will be mostly exchanged for itself (i.e. exchange will occur). In groups with steep dominance hierarchies, interchange is expected: individuals will groom higher ranking individuals to get access to limited resources or commodities such as support in conflicts, and grooming will be traded for these commodities. We examine patterns of reciprocity in grooming and support, and of interchange of grooming for support or for tolerance in six captive groups of bonobos. We test whether differences between groups in patterns of reciprocity and interchange can be attributed to differences in a measure of steepness of dominance hierarchies, which is based on dyadic agonistic interactions. We found that grooming was reciprocal in some, but not all groups. Support was highly reciprocal, but this was a side effect of dominance in most groups. Interchange between grooming and support was observed in some groups. Corroborating earlier findings, this was a side effect of individuals preferring high ranking individuals as grooming and support partners, possibly because these high-ranking individuals provide more efficient support in conflicts. There was no evidence for interchange of grooming for tolerance. Variability in grooming reciprocity was explained by differences in steepness of dominance hierarchies, as predicted by the biological market models. In groups with a shallow dominance hierarchy, grooming was more reciprocal. This was not true for reciprocity in support. There was some evidence that individuals groomed dominants more frequently in groups with a steep dominance hierarchy. The variation in interchange relations between grooming and support did not depend on the steepness of dominance hierarchies. We suggest that grooming in itself is a valuable commodity in bonobos, especially under captive conditions, which can be exchanged reciprocally. Bonobos may interchange grooming for another value equivalent, with food sharing as a very likely candidate. This interchange effects seem more dependent on potential to monopolise food than on steepness of dominance hierarchies per se.