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Wolter, R., & Krueger, K. (2015). The analysis of social bonds in feral horses. In Proceedings of the 3. International Equine Science Meeting.
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.
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A. Lanata, A. Guidi, G. Valenza, P. Baragli, & E. P. Scilingo. (2016). Quantitative heartbeat coupling measures in human-horse interaction. In 2016 38th Annual International Conference of the IEEE Engineering in Medicine and Biology Society (EMBC) (pp. 2696–2699). 2016 38th Annual International Conference of the IEEE Engineering in Medicine and Biology Society (E.
Abstract: Abstract— We present a study focused on a quantitative estimation of a human-horse dynamic interaction. A set of measures based on magnitude and phase coupling between heartbeat dynamics of both humans and horses in three different conditions is reported: no interaction, visual/olfactory interaction and grooming. Specifically, Magnitude Squared Coherence (MSC), Mean Phase Coherence (MPC) and Dynamic Time Warping (DTW) have been used as estimators of the amount of coupling between human and horse through the analysis of their heart rate variability (HRV) time series in a group of eleven human subjects, and one horse. The rationale behind this study is that the interaction of two complex biological systems go towards a coupling process whose dynamical evolution is modulated by the kind and time duration of the interaction itself. We achieved a congruent and consistent
statistical significant difference for all of the three indices. Moreover, a Nearest Mean Classifier was able to recognize the three classes of interaction with an accuracy greater than 70%. Although preliminary, these encouraging results allow a discrimination of three distinct phases in a real human-animal interaction opening to the characterization of the empirically proven relationship between human and horse.
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Bonnie, K. E., & de Waal, F. B. M. (2006). Affiliation promotes the transmission of a social custom: handclasp grooming among captive chimpanzees. Primates, 47(1), 27–34.
Abstract: Handclasp grooming is a unique social custom, known to occur regularly among some, but not all populations of chimpanzees (Pan troglodytes). As with other cultural behaviors, it is assumed that this distinctive grooming posture is learned socially by one individual from another. However, statistical comparisons among factors thought to influence how a behavior spreads within a group have never, to our knowledge, been conducted. In the present study, the origination and spread of handclasp grooming in a group of captive chimpanzees was followed throughout more than 1,500 h of observation over a period of 12 years. We report on the frequency, bout duration, and number and demography of performers throughout the study period, and compare these findings to those reported for wild populations. We predicted that dyads with strong affiliative ties, measured by time spent in proximity to and grooming one another, were likely to develop a handclasp grooming partnership during the study period. A quadratic assignment procedure was used to compare correlations among observed frequencies of grooming and proximity with handclasp grooming in all possible dyads within the group. As predicted, the formation of new handclasp grooming dyads was positively correlated with the rate of overall grooming and proximity within a dyad. In addition, in nearly all dyads formed, at least one individual had been previously observed to handclasp groom. We concluded that affiliation and individual experience determines the transmission of handclasp grooming among captive chimpanzees.
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de Waal, F. B. M. (2003). Darwin's legacy and the study of primate visual communication. Ann N Y Acad Sci, 1000, 7–31.
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.
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de Waal, F. B., Aureli, F., & Judge, P. G. (2000). Coping with crowding. Sci Am, 282(5), 76–81.
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Aureli, F., Preston, S. D., & de Waal, F. B. (1999). Heart rate responses to social interactions in free-moving rhesus macaques (Macaca mulatta): a pilot study. J Comp Psychol, 113(1), 59–65.
Abstract: Heart rate telemetry was explored as a means to access animal emotion during social interactions under naturalistic conditions. Heart rates of 2 middle-ranking adult females living in a large group of rhesus macaques (Macaca mulatta) were recorded along with their behavior. Heart rate changes during 2 types of interactions were investigated, while controlling for the effects of posture and activity. The risk of aggression associated with the approach of a dominant individual was expected to provoke anxiety in the approachee. This prediction was supported by the heart rate increase after such an approach. No increase was found when the approacher was a kin or a subordinate individual. The tension-reduction function of allogrooming was also supported. Heart rate decelerated faster during the receipt of grooming than in matched control periods.
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de Waal, F. B., & Seres, M. (1997). Propagation of handclasp grooming among captive chimpanzees. Am. J. Primatol., 43(4), 339–346.
Abstract: A grooming posture previously reported for two wild chimpanzee (Pan troglodytes) communities developed spontaneously in a captive group of the same species. This offered a unique opportunity to follow the propagation of a new social custom. The posture consists of two partners grasping hands--either both right hands or both left hands--and raising the arms in an A-frame above their heads while mutually grooming with their free hands. The propagation of this pattern was followed over a 5 year period. In the beginning, handclasps were always initiated by the same adult female. This female initiated the posture mainly with her adult female kin. In subsequent years, these relatives became frequent participants in the posture with each other as well as with nonrelatives. Over the years the posture increased in frequency and duration and spread to the majority of adults and also to a few adolescents and older juveniles. The pattern persisted after removal of the apparent originator.
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Judge, P. G., & de Waal, F. B. (1994). Intergroup grooming relations between alpha females in a population of free-ranging rhesus macaques. Folia Primatol (Basel), 63(2), 63–70.
Abstract: Intergroup affiliation among female rhesus macaques, Macaca mulatta, was examined in the captive free-ranging colony of Morgan Island, S.C., USA. The provisioned colony has many social groups (35) and is maintained at a relatively high population density (21 animals/ha) with a relatively low adult male to female ratio (1:8.8). Focal and ad libitum samples were collected on 32 adults (3 males and 29 females) from two groups. Although infrequent, grooming was observed between adult females from different groups, and alpha females were the main participants in these interactions. Colony records indicated that none of the intergroup grooms was between females formerly from a common group. Relations between familiar neighboring groups may be maintained by a combination of both affiliative and aggressive behavior.
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de Waal, F. B., & Johanowicz, D. L. (1993). Modification of reconciliation behavior through social experience: an experiment with two macaque species. Child Dev, 64(3), 897–908.
Abstract: Reconciliation, defined as a friendly reunion between former opponents shortly after an aggressive encounter, is common in the stumptail macaque (Macaca arctoides) but rare in the rhesus macaque (M. mulatta). Juveniles of the two species were cohoused for 5 months, after which they were observed with conspecifics only. Control rhesus monkeys, matched in age and sex to the experimental subjects, went through the same procedure without exposure to the other species. A threefold increase in the proportion of reconciled fights was measured in the rhesus subjects. The difference emerged gradually during cohousing with the tutor species and was sustained following removal of this species. Other behavior, such as grooming and aggression, decreased over time. It is suggested that the social attitude of the subjects was affected through contact with a species characterized by a more relaxed dominance style.
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de Waal, F. B., & Luttrell, L. M. (1986). The similarity principle underlying social bonding among female rhesus monkeys. Folia Primatol (Basel), 46(4), 215–234.
Abstract: Twenty adult female rhesus monkeys (Macaca mulatta) were observed over a three-year period. They lived in a mixed captive group with kinship relations known for three generations. The study's aim was to test Seyfarth's [J. theor. Biol. 65: 671-698, 1977] model of rank-related grooming and to investigate two other possible determinants of social bonding, i.e. relative age and the group's stratification into two social classes. Data on affiliation, coalitions, and social competition were collected by means of both focal observation and instantaneous time sampling. Whereas certain elements of the existing model were confirmed, its explanatory principles were not. Social competition did not result in more contact among close-ranking females (the opposite effect was found), and the relation between affiliative behavior and coalitions was more complex than predicted. Based on multivariate analyses and a comparison of theoretical models, we propose a simpler, more encompassing principle underlying interfemale attraction. According to this 'similarity principle', rhesus females establish bonds with females whom they most resemble. The similarity may concern genetical and social background, age, hierarchical position and social class. Effects of these four factors were independently demonstrated. The most successful model assumed that similarity factors influence female bonding in a cumulative fashion.
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