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P. K. McGregor,, & T. M. Peake,. (2000). Communication networks: social environments for receiving and signalling behaviour. Acta. Ethol., 2(2), 71–81.
Abstract: Communication and social behaviour are inextricably linked, with communication mediating important social behaviours such as resource defence and mate attraction. However, the social environment in which communication occurs is often ignored in discussions of communication behaviour. We argue that networks of several individuals are the common social environment for communication behaviour. The consequences for receivers and signallers of communicating in a network environment are the main subjects of this review. Eavesdropping is a receiving behaviour that is only possible in the environment of a network and therefore we concentrate on this behaviour. The main effect of communication networks on signallers is to create competition with other signallers for receiver attention. We discuss the consequences of such competition. To conclude, we explore the role of signals and signalling interactions as sources of information that animals exploit to direct their behaviour.
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VanDierendonck, M. C. (2006). Social relationships in a group of horses without a mature stallion (Vol. Chapter 4). 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. (2006). Diff erences in social behaviour between late pregnant, post-partum and barren mares in a herd of Icelandic horses (Vol. Chapter 5). 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. (2006). Interventions in social behaviour in a herd of mares and geldings (Vol. Chapter 6). 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. (2006). General Discussion (Vol. Chapter 7). 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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Wey, T., Blumstein, D. T., Shen, W., & Jordán, F. (2008). Social network analysis of animal behaviour: a promising tool for the study of sociality. Anim. Behav., 75(2), 333–344.
Abstract: Social animals live and interact together, forming complex relationships and social structure. These relationships can have important fitness consequences, but most studies do not explicitly measure those relationships. An approach that explicitly measures relationships will further our understanding of social complexity and the consequences of both direct and indirect interactions. Social network analysis is the study of social groups as networks of nodes connected by social ties. This approach examines individuals and groups in the context of relationships between group members. Application of social network analysis to animal behaviour can advance the field by identifying and quantifying specific attributes of social relationships, many of which are not captured by more common measures of sociality, such as group size. Sophisticated methods for network construction and analysis exist in other fields, but until recently, have seen relatively little application to animal systems. We present a brief history of social network analysis, a description of basic concepts and previous applications to animal behaviour. We then highlight relevance and constraints of some network measures, including results from an original study of the effect of sampling on network parameter estimates, and we end with promising directions for research. By doing so, we provide a prospective overview of social network analysis' general utility for the study of animal social behaviour.
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Whishaw, I. Q., Sacrey, L. - A. R., & Gorny, B. (2009). Hind limb stepping over obstacles in the horse guided by place-object memory. Behav. Brain. Res., 198(2), 372–379.
Abstract: An animal that has stepped over an obstacle with its forelimbs uses a memory of the obstacle to guide the hind limbs so that they also clear the obstacle, even in situations in which long pauses are introduced between forelimb and hind limb stepping. To further clarify the features of hind limb obstacle clearance memory, the present study examined hind limb obstacle clearance in the horse. A rider guided horses over obstacles and paused the horse over obstacles in tests that examined the relationship between forelimb and hind limb stepping, with the following results. First, the horses displayed memory for an obstacle as measured by hind limb lifting over the obstacle for durations lasting as long as 15Â min. The response was not dependent upon ongoing visualization of the obstacle, as limb lifting was unaffected by visual occlusion with blinders, a blindfold, or by removing the obstacle during the pause. Second, previous experience of stepping over an obstacle led to pause-related hind limb lifting at the object's previous location even on trials for which there was no obstacle and so no preceding forelimb lifting. Third, whereas a horse would lift its hind limbs to clear two successively presented obstacles, replacing an obstacle before the horse after the forelimbs had cleared the obstacle prevented subsequent hind limb lifting at the obstacle's previous location. Taken together the results show that hind limb obstacle clearance is guided by a place-object memory. The results are discussed in relation to the differential sensory and memonic control of forelimb and hind limb stepping with the suggestion that place-object memory can guide hind stepping as well as overshadow working memory from front leg stepping.
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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.
Keywords: Horses; Social relationships; Interventions; Mares; Geldings; Play; Allogrooming; Social network
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Sueur, C., & Petit, O. (2008). Organization of Group Members at Departure Is Driven by Social Structure in Macaca. Int. J. Primatol., 29(4), 1085–1098.
Abstract: Abstract Researchers have often explained order of progression of group members during joint movement in terms of the influence of ecological pressures but rarely that of social constraints. We studied the order of joining by group members to a movement in semifree-ranging macaques with contrasting social systems: 1 group of Tonkean macaques (Macaca tonkeana) and 1 group of rhesus macaques (M. mulatta). We used network metrics to understand roles and associations among individuals. The way the macaques joined a movement reflected the social differences between the species in terms of dominance and kinship. Old and dominant male rhesus macaques were more often at the front of the movement, contrary to the Tonkean macaques, which exhibited no specific order. Moreover, rhesus macaques preferred to join high-ranking or related individuals, whereas Tonkean macaques based associations during joining mostly on sexual relationships with a subgroup of peripheral males.
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Krause, J., Croft, D., & James, R. (2007). Social network theory in the behavioural sciences: potential applications. Behav. Ecol. Sociobiol., 62(1), 15–27.
Abstract: Abstract Social network theory has made major contributions to our understanding of human social organisation but has found relatively little application in the field of animal behaviour. In this review, we identify several broad research areas where the networks approach could greatly enhance our understanding of social patterns and processes in animals. The network theory provides a quantitative framework that can be used to characterise social structure both at the level of the individual and the population. These novel quantitative variables may provide a new tool in addressing key questions in behavioural ecology particularly in relation to the evolution of social organisation and the impact of social structure on evolutionary processes. For example, network measures could be used to compare social networks of different species or populations making full use of the comparative approach. However, the networks approach can in principle go beyond identifying structural patterns and also can help with the understanding of processes within animal populations such as disease transmission and information transfer. Finally, understanding the pattern of interactions in the network (i.e. who is connected to whom) can also shed some light on the evolution of behavioural strategies.
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