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Taillon, J., & Côté, S. (2008). Are faecal hormone levels linked to winter progression, diet quality and social rank in young ungulates ? An experiment with white-tailed deer ( Odocoileus virginianus ) fawns. Behav. Ecol. Sociobiol., 62(10), 675–677.
Abstract: Abstract Hormones play a central role in the physiology and behaviour of animals. The recent development of noninvasive techniques has increased information on physical and social states of individuals through hormone measurements. The relationships among hormones, life history traits and behaviours are, however, still poorly known. For the first time, we evaluated natural winter glucocorticoid and testosterone levels in young ungulates in relation to winter progression, diet quality and social rank. Overwinter, levels of glucocorticoid and testosterone decreased, possibly due to the decline of fawns" body mass. The relationships between hormone levels and diet quality were surprising: Fawns fed the control diet presented higher glucocorticoid and lower testosterone levels then fawns fed the poor diet, suggesting that control fawns faced a higher nutritional stress than those on the poor diet. Similarly to other studies on social mammals, we found no relationship between faecal glucocorticoid levels and social rank, suggesting that social stress was similar for dominant and subordinate fawns during winter. Testosterone levels were not correlated to social rank as found previously in groups of individuals forming stable social hierarchies and maintaining stable dominance relationships. The simultaneous suppression of glucocorticoid and testosterone levels suggests for the first time that young ungulates present a hormonal strategy to prevent fast depletion of limited proteins and fat resources during winter.
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Hemelrijk, C. K., Wantia, J., & Gygax, L. (2005). The construction of dominance order: comparing performance of five methods using an individual-based model. Behaviour, 142(8), 1043–1064.
Abstract: In studies of animal behaviour investigators correlate dominance with all kinds of behavioural
variables, such as reproductive success and foraging success. Many methods are used to
produce a dominance hierarchy from a matrix reflecting the frequency of winning dominance
interactions. These different methods produce different hierarchies. However, it is difficult to
decide which ranking method is best. In this paper, we offer a new procedure for this decision:
we use an individual-based model, called DomWorld, as a test-environment. We choose this
model, because it provides access to both the internal dominance values of artificial agents
(which reflects their fighting power) and the matrix of winning and losing among them and,
in addition, because its behavioural rules are biologically inspired and its group-level patterns
resemble those of real primates. We compare statistically the dominance hierarchy based on
the internal dominance values of the artificial agents with the dominance hierarchy produced
by ranking individuals by (a) their total frequency of winning, (b) their average dominance
index, (c) a refined dominance index, the David`s score, (d) the number of subordinates each
individual has and (e) a ranking method based on maximizing the linear order of the hierarchy.
Because dominance hierarchies may differ depending on group size, type of society, and the
interval of study, we compare these ranking methods for these conditions.We study complete
samples as well as samples randomly chosen to resemble the limitations of observing real
animals. It appears that two methods of medium complexity (the average dominance index
and David`s score) lead to hierarchical orders that come closest to the hierarchy based on
internal dominance values of the agents. We advocate usage of the average dominance index,
because of its computational simplicity.
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Vervaecke, H., Vries, H. D., & Elsacker, L. V. (1999). An Experimental Evaluation Of The Consistency Of Competitive Ability And Agonistic Dominance In Different Social Contexts In Captive Bonobos. Behaviour, 136(4), 423–442.
Abstract: Bonobos have been described as a relatively egalitarian and female dominant species. The exact nature and quality of their dominance relationships and the existence of female dominance are current topics of dispute. We investigated the consistency across social contexts, the stability in time, and the degree of expression of the competitive feeding ability and agonistic dominance in a captive group of bonobos. First, we examined whether the competitive feeding ranks and agonistic ranks differed in different dyadic contexts, triadic contexts and the whole group context. For some pairs of animals the dominance relationships with respect to competitive feeding altered with different group compositions. The agonistic dominance relationships changed accordingly. The competitive feeding ranks and agonistic ranks in the experiments correlated strongly with each other. The alpha position was occupied by a female, but not all females outranked all males. We suggest that females can profit from each others presence to gain inter-sexual dominance. Second, although the agonistic rank order in the whole group remained the same over at least five years, some dyadic competitive feeding ranks changed over time, resulting in a stronger female intersexual dominance. Third, the degree of expression of the behaviors used to quantify dyadic competitive and agonistic dominance was not high, in line with the popular 'egalitarian' epithet. Notwithstanding its low consistency across contexts, the dominance hierarchy in the whole group has a strong predictive value for other social relationships such as grooming. Given this strong effect of rank on other behaviours and given the strong dependency of rank on social context, the choice of the right party members may be a crucial factor in the fission-fusion processes of free-ranging bonobos.
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Heitor, F., do Mar Oom, M., & Vicente, L. (2006). Social relationships in a herd of Sorraia horses: Part II. Factors affecting affiliative relationships and sexual behaviours. Behav. Process., 73(3), 231–239.
Abstract: The influence of age, dominance rank, kinship and aggressiveness over affiliative relationships and sexual behaviours were analysed in a herd of Sorraia horses, Equus caballus, kept under extensive management. Subjects were 10 adult mares 5-18 years old that had known each other since birth, and a stallion introduced into the group for breeding for the first time. Kinship coefficient and dominance rank were the most important factors affecting affiliative relationships. Bonds were reciprocal and stronger among mares with higher kinship. Mares spent more time in proximity to close-ranking and lower-ranking females. Mares with stronger affiliative relationships or higher relatedness were not less aggressive towards each other. Affiliative relationships between the stallion and the mares were not reciprocal: lower-ranking mares formed stronger bonds with the stallion but he preferred the less genetically related mares for proximity. However, the stallion was involved in sexual behaviours more frequently with the mares that were more genetically related to him. These results suggest that kinship beyond close relatives may affect affiliative relationships both among familiar and among unfamiliar horses. However, the influence of kinship does not imply that horses possess a kin recognition system and alternative explanations are discussed.
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Krueger, K., Schneider, G., Flauger, B., & Heinze, J. (2015). Context-dependent third-party intervention in agonistic encounters of male Przewalski horses. Behav. Process., 121, 54–62.
Abstract: Abstract One mechanism to resolve conflict among group members is third party intervention, for which several functions, such as kin protection, alliance formation, and the promotion of group cohesion have been proposed. Still, empirical research on the function of intervention behaviour is rare. We studied 40 cases of intervention behaviour in a field study on 13 semi-wild bachelor horses (Equus ferus przewalskii) in (a) standard social situations, and (b) when new horses joined the group (i.e. introductions). Only interventions in agonistic encounters were analysed. Eight of 13 animals directed intervention behaviour toward threatening animal in agonistic encounters of group members. One stallion was particularly active. The stallions did not intervene to support former group mates or kin and interventions were not reciprocated. In introduction situations and in standard social situations, the interveners supported animals which were lower in rank, but targeted, threatening animals of comparable social rank. After introductions, stallions received more affiliative behaviour from animals they supported and thus appeared to intervene for alliance formation. In standard social situations, interveners did not receive more affiliative behaviour from animals they supported and may primarily have intervened to promote group cohesion and to reduce social disruption within the group.
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Krueger, K., & Flauger, B. (2008). Social feeding decisions in horses (Equus caballus). Behav. Process., 78(1), 76–83.
Abstract: Like many other herbivores, in a natural environment equids feed on rather evenly distributed resources. However, the vegetation in their vast habitats constantly changes. If food is plentiful only little competition occurs over food, and in non-competitive situations domestic horses tend to return to the same feeding site until it is overgrazed. In contrast, they compete over limited food for which the social status of the individuals appears to be important. Especially in ruminants several studies have proved an influence of social organisations, rank, sex and the depletion of feeding sites on the feeding behaviour of individuals. However, it is not yet understood whether and how social aspects affect horses“ feeding decisions. Curiosity about the influence of social rank on the horses” feeding decisions between two, equally with high-quality surplus food-filled buckets placed in different social feeding conditions, led us to create the test below. The observer horses were alternately tested with a dominant and a subordinate demonstrator placed in one of the three different positions. We conclude that domestic horses use social cognition and strategic decision making in order to decide where to feed in a social feeding situation. When possible they tend to return to the same, continuously supplied feeding site and switch to an “avoidance tendency” in the presence of dominant horses or when another horse is already feeding there. Thus, the social rank and the position of conspecifics affect the feeding strategy of horses.
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Krueger, K., Flauger, B., Farmer, K., & Hemelrijk, C. (2014). Movement initiation in groups of feral horses. Behav. Process., 103, 91–101.
Abstract: Abstract Herds of ungulates, flocks of birds, swarms of insects and schools of fish move in coordinated groups. Computer models show that only one or very few animals are needed to initiate and direct movement. To investigate initiation mechanisms further, we studied two ways in which movement can be initiated in feral horses: herding, and departure from the group. We examined traits affecting the likelihood of a horse initiating movement i.e. social rank, affiliative relationships, spatial position, and social network. We also investigated whether group members join a movement in dominance rank order. Our results show that whereas herding is exclusive to alpha males, any group member may initiate movement by departure. Social bonds, the number of animals interacted with, and the spatial position were not significantly associated with movement initiation. We did not find movement initiation by departure to be exclusive to any type of individual. Instead we find evidence for a limited form of distributed leadership, with higher ranking animals being followed more often.
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Heitor, F., & Vicente, L. (2010). Dominance relationships and patterns of aggression in a bachelor group of Sorraia horses (Equus caballus). J. Ethol., 28(1), 35–44.
Abstract: Abstract The influence of individual factors on dominance rank and the relationship between rank distance and patterns of aggression predicted by models of evolutionarily stable strategies (ESS) of animal conflict were investigated in a managed bachelor group of Sorraia horses, Equus caballus. The group was composed of four to six stallions 3- to 12-years-old during the study period. The dominance hierarchy was significantly linear and rank was not related to age, weight, height or aggressiveness. Frequency and intensity of agonistic interactions were low, but higher-ranking stallions did not receive lower aggressiveness than lower-ranking stallions. There was some evidence that dominance relationships were more contested among close-ranking stallions, as predicted. Agonistic-related interactions among close-ranking stallions served similar functions to those among distant-ranking stallions, but the latter interacted more frequently than expected for access to resting sites and/or resting partners. Therefore, we found some evidence that agonistic-related interactions among distant-ranking stallions play a larger role in providing access to valuable and defendable resources than those among close-ranking stallions. Nevertheless, the fact that space to escape from aggression was limited and breeding access was independent from dominance rank may have reduced the benefits relative to costs of aggression and therefore limited the occurrence of contests over dominance and resources.
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VanDierendonck, M. C., de Vries, H., & Schilder, M. B. H. (1995). An Analysis of Dominance, Its Behavioural Parameters and Possible Determinants in a Herd of Icelandic orses in Captivity. Netherl. J. Zool., 45(3-4), 362–385.
Abstract: Th e applicability of the concept of dominance was investigated in a captive herd of  Icelandic
horses and  ponies of diff erent breeds. Eight out of  behaviours possibly related
to dominance occurred frequently enough to be investigated in detail. For these eight agonistic
behaviours the coverage, the unidirectionality in the exchange, and the degree of
transitivity (Landau`s linearity index) were calculated. Four off ensive behaviours, together
with avoidance, were suitable for further analysis with regard to dominance. Th e patterns
of asymmetries with which these behaviours were exchanged were suffi ciently similar as to
justify the application of the dominance concept and to construct a (nearly) linear dominance
hierarchy. Th e rank order of the castrated stallions was completely linear, the hierarchy
of the mares was almost completely linear. Th e results suggest that off ensive and defensive
aggressive behaviours should be treated separately and that the concept of dominance
is applicable. However, ritualized formal dominance signals between adult horses appear to
be (almost) absent. Th e rank positions of the individuals were correlated with age and residency
in the herd but not with height. Middle ranking horses tended to be more frequently
in the close vicinity of another horse than high ranking or low ranking horses. Over and
above this correlation at the individual level, it was found that pairs of horses close in rank
to each other were more often also spatially close to each other. Being in oestrus did not infl
uence the dominance relationships between mares. For castrated stallions the rank positions
were correlated with the age at which they were castrated. Th is suggests that in male
horses experience prior to neutering infl uences the behaviour afterwards.
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VanDierendonck, M. C., de Vries, H., Schilder, M.B.H. (1995). An Analysis of Dominance, Its Behavioural Parameters and Possible Determinants in a Herd of Icelandic horses in captivity. Netherl. J. Zool., 45(3-4), 362–385.
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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