Abstract: Using heart rate (HR) measurements we investigated whether potential calming effects of social licking were evident for both active (performers) and passive (receivers) licking partners. A HR decline was assumed to indicate relaxation and thus the experience of positive emotions. Effects of the licking category (spontaneous, solicited), the animals' basic activity (standing, lying) and the licked body region (head, neck, rest) were also considered. Two studies (A, B) were carried out in the same loose housed Austrian Simmental dairy herd. HR was recorded in up to 20 focal animals on 16 and 18 days, respectively. Using either direct observations (A) or video recordings (B), social licking interactions were continuously observed. The cow's basic activity was recorded using scan sampling at 5 min intervals. Linear mixed effects models were applied separately for Study A and B in order to compare the mean HR of the licking bouts with the mean of the respective 5 min pre- and post-licking periods. In receivers we found a significant calming effect in terms of a HR decline during allogrooming in both studies (A: -1.3 beats per minute, B: -1.1 bpm). This effect was more pronounced when animals were standing (A/B: -2.4 bpm/-3.8 bpm). However, it was not affected by the licked body region. In dairy cows performing social licking, we did not find an overall calming effect. On the contrary, in Study B, HR significantly increased during licking in lying performers (+2.5 bpm). This reaction was even stronger, when licking was directed to the receivers' head (+3.5 bpm) or neck (+3.0 bpm) as compared to the rest of the body (+1.4 bpm). The licking category had no effect on HR changes during the licking events. Our findings suggest that relaxation effects induced by social licking differ between performers and receivers and are affected by the cows' basic activity. In receivers, there were clear indications of a calming effect implying the experience of positive affective states. In performers, such calming effects during social licking were not identified.

Abstract: In many social mammals, individuals preferentially affiliate with a small subset of available partners instead of distributing their social behaviors equally among all group members. The resulting social bonds have been investigated in several mammalian taxa, especially in primate societies, but also in other taxa such as birds, dolphins, rodents and ungulates. In feral horses, a great number of studies on social bonds can be found, but with a huge variety between methods for the analysis. There seems to be a lack of a clear and common definition of social bonds in horses and of comparable analyses. For example, there are irregularities between the studies regarding the research designs, the selection of recording methods and the interpretation of the measurements. Mutual grooming is used most often for the analysis of social relationships in many species. As mutual grooming is rare in horses, especially measurements of spatial proximity are commonly used for the analysis of social bonds in addition to other behavioral patterns. However, the combination of mutual grooming and nearest neighborhood analyses for the analysis of social bond is debatable, as in contrast to mutual grooming, which must occur deliberately by both grooming partners, the spatial distribution can be influenced by one partner alone, which may even force the other horse to keep a certain distance or to stay in close proximity.
In this study, we investigated the comparability of mutual grooming and nearest neighborhood data for social bond analyses in feral horses. Therefore, we observed five groups of semi-wild living Przewalski’s horses and six groups of feral horses.
We analysed the horses’ social ranks by applying an Average Dominance Index, we recorded the distances between the animals and observed the number of mutual grooming events as well as friendly approaches.
Our results show that there was only a weak correlation between the frequency of staying in nearest neighborhood and mutual grooming in all observed horse groups. In contrast to this, the correlation between the number of friendly approaches and mutual grooming events was higher in most groups.
Hierarchies did not affect social bonds, as mutual grooming was similarly induced by higher and lower ranking animals and the social rank did not affect the choice of the grooming partner. Similarly, likelihoods of staying in the neighborhood of particular animals were not affected by the animals’ social rank.
The grooming frequencies differed between the different horse groups and between the individual horses living in the particular groups. They seem to be effected by individual predisposition.
Altogether we suggest that the ratio of mutual grooming seems to be a better indicator for social bonds in feral horses than the frequency of staying in the nearest neighborhood. Mutual grooming occurs deliberately and is bidirectional, whereas nearest neighborhoods could be enforced and unidirectional. For the calculation of social bonds in horses, we consider it to be more reliable to combine the frequency of mutual grooming with the frequency of friendly approaches.

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: Simple Summary
Because of their social nature, horses need to have plenty of opportunities to interact with others to establish bonds and learn from their elders. Comparison of social behaviour of 426 horses in 20 groups of Icelandic horses in pastures, showed that aggression was lowest where the group composition was like the natural system, i.e., with a stallion, mares and their young foals. In groups without a stallion, the presence of foals is also associated with low aggression. Stability of the group with respect to group composition is of great importance; the horses are less aggressive in the more stable groups. The highest aggression was found in groups of unfamiliar yearlings. The horses allogroomed more in groups with relatively more young horses, which suggests they are forming bonds. Later, they groom less but prefer certain individuals. Horse owners should all be aware of the importance of planning the composition of horse groups and to keep the membership as stable as possible in order to ensure good welfare.
Abstract
We explore how herd composition and management factors correlate with frequencies of social interactions in horse groups. Since the welfare of horses correlates with low aggression levels and social contact opportunities, information of this kind is important. The data are a collection of records of social interactions of 426 Icelandic horses in 20 groups of at least eight horses. The complexities and limitations of the data prohibit useful statistical modelling so the results are presented descriptively. Interesting and informative patterns emerge which can be of use both in management and in future studies. Of special interest are the low levels of agonistic behaviours in breeding groups where one stallion was present. The horses were less agonistic when in groups with young foals and where group membership was stable. Unfamiliar yearlings in peer groups were especially aggressive. Allogrooming was most frequent in groups with relatively more young horses and in unstable and small groups. Interestingly, the horses allogroomed more if they had few preferred allogrooming partners. The findings show that composition (age/sex) and stability of groups are of great importance with respect to aggression levels and opportunities for establishing bonds.

Abstract: After Charles Darwin's The Expression of the Emotions in Man and Animals, published in 1872, we had to wait 60 years before the theme of animal expressions was picked up by another astute observer. In 1935, Nadezhda Ladygina-Kohts published a detailed comparison of the expressive behavior of a juvenile chimpanzee and of her own child. After Kohts, we had to wait until the 1960s for modern ethological analyses of primate facial and gestural communication. Again, the focus was on the chimpanzee, but ethograms on other primates appeared as well. Our understanding of the range of expressions in other primates is at present far more advanced than that in Darwin's time. A strong social component has been added: instead of focusing on the expressions per se, they are now often classified according to the social situations in which they typically occur. Initially, quantitative analyses were sequential (i.e., concerned with temporal associations between behavior patterns), and they avoided the language of emotions. I will discuss some of this early work, including my own on the communicative repertoire of the bonobo, a close relative of the chimpanzee (and ourselves). I will provide concrete examples to make the point that there is a much richer matrix of contexts possible than the common behavioral categories of aggression, sex, fear, play, and so on. Primate signaling is a form of negotiation, and previous classifications have ignored the specifics of what animals try to achieve with their exchanges. There is also increasing evidence for signal conventionalization in primates, especially the apes, in both captivity and the field. This process results in group-specific or “cultural” communication patterns.