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Fischhoff, I. R., Sundaresan, S. R., Cordingley, J., Larkin, H. M., Sellier, M. - J., & Rubenstein, D. I. (2007). Social relationships and reproductive state influence leadership roles in movements of plains zebra, Equus burchellii. Anim. Behav., 73(5), 825–831.
Abstract: In animal groups, collective movements emerge from individual interactions. Biologists seek to identify how characteristics of actors in these groups, and their relationships, influence the decision-making process. We distinguished two basic factors determining leadership in group choices: identity and state. We hypothesized that identity is more important to leadership in groups with stable relationships, which permit the development of habitual roles. In groups with fluid membership, particular individuals or subgroups are less likely to emerge as consistent leaders. Instead, we predicted that movement initiation in unstable groups depends on individual state at the time of the decision. We characterized how identity and reproductive state influenced leadership patterns in the movements of plains zebra. As in many other mammals, lactation in this species significantly alters water and energy needs. We investigated leadership in tightly knit harems and loosely bonded herds of multiple harems. Harem females tended to have habitual roles in the initiation of harem movement. In herds, however, we found no consistent leaders among harems. At both levels of social organization, lactation was a key determinant of leadership. In harems, lactating females were more likely to initiate movement than nonlactating females. In turn, harems containing lactating females were more likely to lead herd movements. Thus, we conclude that social relationships and reproductive state together shape the interactions that produce group behaviours. One benefit to lactating females of leading herd movements is preferential access to scarce water. Thus, leadership roles in group decisions may have fitness consequences.
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Heitor, F., & Vicente, L. (2008). Maternal care and foal social relationships in a herd of Sorraia horses: Influence of maternal rank and experience. Appl. Anim. Behav. Sci., 113(1-3), 189–205.
Abstract: The influence of maternal rank and experience on patterns of maternal care and social relationships of foals were investigated in a managed herd of Sorraia horses, Equus caballus. Social interactions and spatial relationships of 13 foals (seven females and six males) born to seven mares were examined from birth to 10 months of life, within the three major periods of foal development. Conflict over suckling between dam and foal was not generally affected by rank and experience, but higher-ranking mothers allowed more suckling during late lactation than lower-ranking mothers. Foals of higher-ranking mares spent more time in proximity to the mother during socialization. Maternal rank and experience did not significantly affect maternal protectiveness, foal independence from the mother or the development of affiliative relationships between foals and group members. Foals of higher-ranking mares received lower frequencies of aggression from other horses only in the first month of life. Dominance relationships among foals depended mainly on aggressiveness and were not associated with maternal rank. The large variability in maternal behaviour, the absence of a significant association between maternal rank and body condition at parturition and the stable social environment within this herd may partly account for the reported results.
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Krueger, K., Farmer, K., & Heinze, J. (2014). The effects of age, rank and neophobia on social learning in horses. Anim. Cogn., 17(3), 645–655.
Abstract: Social learning is said to meet the demands of complex environments in which individuals compete over resources and co-operate to share resources. Horses (Equus caballus) were thought to lack social learning skills because they feed on homogenously distributed resources with few reasons for conflict. However, the horse’s social environment is complex, which raises the possibility that its capacity for social transfer of feeding behaviour has been underestimated. We conducted a social learning experiment using 30 socially kept horses of different ages. Five horses, one from each group, were chosen as demonstrators, and the remaining 25 horses were designated observers. Observers from each group were allowed to watch their group demonstrator opening a feeding apparatus. We found that young, low ranking, and more exploratory horses learned by observing older members of their own group, and the older the horse, the more slowly it appeared to learn. Social learning may be an adaptive specialisation to the social environment. Older animals may avoid the potential costs of acquiring complex and potentially disadvantageous feeding behaviours from younger group members. We argue that horses show social learning in the context of their social ecology, and that research procedures must take such contexts into account. Misconceptions about the horse’s sociality may have hampered earlier studies.
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Mitani, J. C. (2009). Male chimpanzees form enduring and equitable social bonds. Anim. Behav., 77(3), 633–640.
Abstract: Controversy exists regarding the nature of primate social relationships. While individual primates are frequently hypothesized to form enduring social bonds with conspecifics, recent studies suggest that relationships are labile, with animals interacting only over short periods to satisfy their immediate needs. Here I use data collected over 10 years on a community of chimpanzees, Pan troglodytes, at Ngogo, Kibale National Park, Uganda, to investigate whether male chimpanzees establish long-term social relationships and to determine the factors that affect variation in relationship quality and the stability of social bonds. Kinship and dominance rank influenced the quality of relationships. Maternal brothers and males of the same dominance rank class groomed each other more equitably than did unrelated males and males that were dissimilar in rank. In addition, males that formed strong social bonds groomed more equitably than did males that displayed weaker bonds. Social bonds were stable over time, with relationships in one year predicting those in subsequent years. Kinship and the quality of social relationships affected bond stability. Maternal half siblings and males that groomed each other equitably maintained longer-lasting bonds than did nonkin and males that groomed each other unevenly. Virtually all of the males established at least one enduring relationship with another individual. The most enduring bonds formed between a few pairs of maternal brothers and dyads that maintained balanced grooming interactions. These results indicate that male chimpanzees maintain long-lasting and equitable social bonds whose formation is affected by maternal kinship and the quality of social relationships.
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Schwab, C., Bugnyar, T., Schloegl, C., & Kotrschal, K. (2008). Enhanced social learning between siblings in common ravens, Corvus corax. Anim. Behav., 75(2), 501–508.
Abstract: It has been suggested that social dynamics affect social learning but empirical support for this idea is scarce. Here we show that affiliate relationships among kin indeed enhance the performance of common ravens, Corvus corax, in a social learning task. Via daily behavioural protocols we first monitored social dynamics in our group of captive young ravens. Siblings spent significantly more time in close proximity to each other than did nonsiblings. We subsequently tested birds on a stimulus enhancement task in model-observer dyads composed of both siblings and nonsiblings. During demonstration the observer could watch the model manipulating one particular object (target object) in an adjacent room. After removing the model, the observer was confronted with five different objects including the former target object. Observers from sibling dyads handled the target object for significantly longer periods of time as compared with the other four available objects, whereas observers from nonsibling dyads did not show a preference for the target object. Also, siblings matched the model's decision to cache or not to cache objects significantly more often than did nonsiblings. Hence, siblings were likely to attend to both, the behaviour of the model (caching or noncaching) and object-specific details. Our results support the hypothesis that affiliate relations between individuals affect the transmission of information and may lead to directed social learning even when spatial proximity has been experimentally controlled for.
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VanDierendonck, M. C. (2006). General Introduction (Vol. Chapter 1). Ph.D. thesis, , .
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.
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VanDierendonck, M. C. (2006). Summary (Vol. Chapter 8). Ph.D. thesis, , Universiteit Utrecht.
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.
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VanDierendonck, M. C., de Vries, H., Schilder, M. B. H., Colenbrander, B., & Þorhallsdóttir, A. G. and S., H. (2009). Interventions in social behaviour in a herd of mares and geldings. Appl. Anim. Behav. Sci., 116(1), 67–73.
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.
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