Abstract: For some time now, social network analysis provides tools to describe concisely the social structure of animals. Nevertheless, the factors that shape the social network and determine the frequency of different types of affiliations are often unclear. For example, the effects of relatedness on social networks have been only studied on relatively few species. Both among social or sub-social species the effects ranged from no effect to a substantial effect. Here we test the hypothesis that relatedness has an effect on the social network of horses (Equus caballus) that live freely in semi-natural conditions, and specifically, that individuals with high values of relatedness to their neighbors in the network will have fewer links Wolf et al, 2011). This hypothesis is based on the notion that related individuals have common interests and are likely to assist each other, and may need less links to other, unrelated individuals. In addition, individuals of similar age and sex are likely to have common needs, and thus are more likely to be associated. We thus tested a second hypothesis: homophily to individuals with similar age and sex will have a role in determining the associations within the social network. The field study was conducted on 27 horses in the Blauwe Kamer reserve in the Netherlands (1.1 sq km). We videotaped horses and their groups and used the information from 22 horses, after excluding the five foals from the analysis. Relatedness was calculated from the pedigree, which was based on parentage, determined by DNA analysis. The social network was constructed based on spatial proximity data. We assessed the influence of relatedness, age-homophily and sex-homophily on the network structure with Multiple Regression Quadratic Assignment Procedure (MRQAP) (Krackhardt 1988), with the R package sna. The results show that there was no significant effect of relatedness on the network, nor an effect of age-homophily. Nevertheless, we found a significant effect of sex-homophily, the tendency of individuals to associate with individuals of the same sex. We argue that the lack of a relatedness effect is not likely to have been caused due to the inability to detect who is kin. The structured social system in horses includes strong associations between often unrelated individuals, alongside with young individuals leaving their natal harem as part of the inherent inbreeding avoidance (Linklater & Cameron 2009; Boyd et al. in press). The significant effect of sex-homophily could stem from the protection females get from associations with other females, e.g., in the reduction of harassment from males. Previous studies on feral horses showed that mares that were better connected with other females in their harem benefited from higher survival rates to their foals (Cameron et al. 2009). The associations among bachelor males could also contribute to the strength of the sex-homophily effect.
In order to generalize from our results, one needs to examine additional populations of horses, because the conditions in the Blauwe Kamer reserve may not be representative, mainly due to the limited opportunity for dispersal in a restricted area.
Keywords: Long-term affiliation; spatial proximity, kin detection
Boyd, L., Scorolli, A. L., Nowzari, H., & Bouskila, A. (2016). Chapter 2: Social organization. In J. I. Ransom, & P. Kaczensky (Eds.), Wild equids. Baltimore, Maryland (in press): The Johns Hopkins University Press.
Cameron, E. Z., Setsaas, T. H., & Linklater, W. L. (2009). Social bonds between unrelated females increase reproductive success in feral horses. Proceedings of the National Academy of Sciences, 106(33), 13850-13853
Krackhardt, D. (1988). Predicting with networks: Nonparametric multiple regression analysis of dyadic data. Social Networks, 10(4), 359-381.
Linklater, W. L., & Cameron, E. Z. (2009). Social dispersal but with philopatry reveals incest avoidance in a polygynous ungulate. Animal Behaviour, 77(5), 1085-1093.
Wolf, J. B., Traulsen, A., & James, R. (2011). Exploring the link between genetic relatedness r and social contact structure k in animal social networks. The American Naturalist, 177(1), 135-142.

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: 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 the existence of a social dominance hierarchy in the captive group of six adult bonobos at the Planckendael Zoo. We quantified the pattern of dyadic exchange of a number of behaviors to examine to what extent each behavior fits a linear rank order model. Following de Waal (1989), we distinguish three types of dominance: agonistic dominance, competitive ability and formal dominance. Fleeing upon aggression is a good measure of agonistic dominance. The agonistic dominance hierarchy in the study group shows significant and strong linearity. The rank order was: 1. female (22 yr), 2. female (15 yr)., 3. male (23 yr.), 4. female (15 yr.), 5. male (9 yr.), 6. male (10 yr.). As in the wild, the females occupy high ranks. There is prominent but nonexclusive female agonistic dominance. Teeth-baring does not fulfil the criteria of a formal submission signal. Peering is a request for tolerance of proximity. Since its direction within dyads is consistent with that of fleeing interactions, it is a useful additional measure to determine agonistic ranks in bonobos. In competitive situations, the females acquire more food than other group members do. The rank obtained from access to food resources differs from the agonistic rank due to female intrasexual social tolerance, expressed in food sharing. We typify the dominance styles in the group as female intrasexual tolerance and male challenging of rank differences. The agonistic rank order correlates significantly with age and has a strong predictive value for other social behaviors.

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.