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Keiper, R. R. (1986). Social structure. Vet Clin North Am Equine Pract, 2(3), 465–484.
Abstract: Socially feral horses live in stable social groups characterized by one adult male, a number of adult females, and their offspring up to 2 years of age. Extra males either live by themselves or with other males in bachelor groups. The bands occupy nondefended home ranges that often overlap. Many abnormal behaviors seen in domestic horses occur because some aspect of their normal social behavior cannot be carried out in captivity.
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Kitchen, D. M., Cheney, D. L., & Seyfarth, R. M. (2005). Male chacma baboons (Papio hamadryas ursinus) discriminate loud call contests between rivals of different relative ranks. Anim. Cogn., 8(1), 1–6.
Abstract: Males in multi-male groups of chacma baboons (Papio hamadryas ursinus) in Botswana compete for positions in a linear dominance hierarchy. Previous research suggests that males treat different categories of rivals differently; competitive displays between males of similar rank are more frequent and intense than those between disparately ranked males. Here we test whether males also respond differently to male-male interactions in which they are not directly involved, using playbacks of the loud 'wahoo' calls exchanged between competing males in aggressive displays. We played paired sequences of vocal contests between two adjacently ranked and two disparately ranked males to ten subjects, half ranking below the signalers in the call sequences and half above. Subjects who ranked above the two signalers showed stronger responses than lower-ranking subjects. Higher-ranking subjects also responded more strongly to sequences involving disparately ranked, as opposed to adjacently ranked opponents, suggesting that they recognized those individuals' relative ranks. Strong responses to sequences between disparately ranked opponents might have occurred either because such contests typically involve resources of high fitness value (defense of meat, estrous females or infants vulnerable to infanticide) or because they indicate a sudden change in one contestant's condition. In contrast, subjects who ranked lower than the signalers responded equally strongly to both types of sequences. These subjects may have been able to distinguish between the two categories of opponents but did not respond differently to them because they had little to lose or gain by a rank reversal between males that already ranked higher than they did.
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Bergman, T. J., Beehner, J. C., Cheney, D. L., & Seyfarth, R. M. (2003). Hierarchical classification by rank and kinship in baboons. Science, 302(5648), 1234–1236.
Abstract: Humans routinely classify others according to both their individual attributes, such as social status or wealth, and membership in higher order groups, such as families or castes. They also recognize that people's individual attributes may be influenced and regulated by their group affiliations. It is not known whether such rule-governed, hierarchical classifications are specific to humans or might also occur in nonlinguistic species. Here we show that baboons recognize that a dominance hierarchy can be subdivided into family groups. In playback experiments, baboons respond more strongly to call sequences mimicking dominance rank reversals between families than within families, indicating that they classify others simultaneously according to both individual rank and kinship. The selective pressures imposed by complex societies may therefore have favored cognitive skills that constitute an evolutionary precursor to some components of human cognition.
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Lehmann, K., Kallweit, E., & Ellendorff, F. (2006). Social hierarchy in exercised and untrained group-housed horses--A brief report. Appl. Anim. Behav. Sci., 96(3-4), 343–347.
Abstract: Changes in social hierarchy were evaluated in a herd of 3-year-old Hanoverian geldings. One group (n = 5) was exposed to a training programme, the other (n = 5) remained untrained. After 6 months, the groups were reversed. Hierarchical positions were evaluated by field observations and/or paired-feeding tests at the beginning, the middle, the end of the first and at the end of the second training period. Both methods yielded identical results. Almost all horses changed position in only one direction: either up or down. Neither increase in aggression nor mutual injuries were recorded during the whole experiment. No statistically verified differences in dominance ranking occurred between trained and untrained groups, but apparent differences were consistent. Thus, if horses are kept in the same group for a longer period of time, exercise induced changes in hierarchy are probably of minor importance and are unlikely to increase the incidence of injuries. This may have implications for the promotion of group-housing for sport horses.
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Dunbar, R. I., & Dunbar, E. P. (1976). Contrasts in social structure among black-and-white colobus monkey groups. Anim. Behav., 24(1), 84–92.
Abstract: Three types of Colobus guereza groups may be distinguished on the bases of size and composition, namely small one-male groups, large, one-male groups and multi-male groups. The social structure of each type of group is described in terms of the distribution of non-agonistic interactions, the frequency and distribution of agonistic behaviour and the organization of the roles of vigilance, territorial defence and leadership. A number of differences are found between the group types which appear to be related to the differences in group size and composition. It is suggested that these group types represent stages in the life-cycle of colobus groups, and that such an interpretation may help to resolve some of the conflicting reports in the literature.
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Puppe, B., Langbein, J., Bauer, J., & Hoy, S. (2008). A comparative view on social hierarchy formation at different stages of pig production using sociometric measures. Livestock Science, 113(2-3), 155–162.
Abstract: A standardised and comprehensive approach to describe dominance relationships in gregarious farm animals quantitatively was recently developed, incorporating a combination of appropriate sociometric measures. The present study applied this approach to a comparative analysis of the social hierarchies within 57 groups of domestic pigs at different age/production stages with a total of 496 animals. Unacquainted pigs were grouped to three age categories which correspond to the typical production stages: weaned pigs (PIG28, 12 groups), growing pigs (PIG80, 16 groups), and reproductive sows (SOW, 29 groups). Based on observed agonistic interactions, sociometric values were calculated both at the dyadic and at the group level and may be considered as preliminary reference values for further studies. As indicated by the respective values of the Kendall index (PIG28: 0.66, tested as significant in 69.0% of the observed groups; PIG80: 0.71, 87.5%; SOW: 0.61, 69.0%), and the improved Landau index (PIG28: 0.70, 75.0%; PIG80: 0.72, 93.7%; SOW: 0.71, 72.4%), a social organisation towards a quasi-linear social hierarchy was predominantly developed throughout all age/production categories. However, compared to weaned and growing pigs, sows were characterised by significant differences concerning establishment (fewer agonistic interactions) and kind (more unknown dyads, fewer two-way and significant dyads, higher directional consistency index) of their social hierarchy. It seems that sows have effectively adapted their agonistic behaviour towards pen-mates to regulate social dominance relationships, whereas younger pigs frequently display agonistic interactions also to gain additional experience on social cues (e.g. the fighting ability of an opponent). Hence, it is concluded that the effective experience of socialisation during sensitive periods may increase the social skills of pigs which in turn can improve their welfare and health, e.g. by adjusted aggressive behaviour. The consideration of comparable and standardised sociometric measures in livestock breeding may help to improve husbandry conditions.
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Langbein, J., & Puppe, B. (2004). Analysing dominance relationships by sociometric methods--a plea for a more standardised and precise approach in farm animals. Appl. Anim. Behav. Sci., 87(3-4), 293–315.
Abstract: Social dominance is a multidimensional phenomenon occurring in all gregarious farm animals and finds its reflection in a dominance hierarchy. Hence, numerous studies have tried to analyse dominance relationships as well as to correlate outcoming results (mostly individual ranks) with other behavioural and/or physiological features of the animals. Although the concept of dominance, once established, has been developed continuously and several sociometric measures were cumulatively introduced, a consistent analysing approach has not been achieved, especially in farm animals. Thus, considerable inconsistencies in the used methodology may impair obtained results and interpretations. The present paper is a plea for a more standardised and complex approach when analysing dominance relationships, not only in farm animals. First, derived from a structural definition of dominance, we suggest in detail the preferably consistent use of appropriate sociometric measures at all social levels of analysis: the dyad as the starting level, the group as the highest level, and the individual as the basic level. Second, we applied this procedures in a case study to analyse social dominance in a group of dwarf goats (n=12) and pigs (n=10), respectively, to comparatively demonstrate benefits and problems of such an approach in two different farm animal species. It is concluded that the use of individual ranks is actually only reasonable when fundamental sociometric measures both at the dyadic level (e.g. percentage of dyads which have a significant asymmetric outcome) and at the group level (e.g. the strength of hierarchy) are successfully tested by statistical methods as also presented in this paper. The calculated sociometric measures deliver not only a more comprehensive “picture” of the social relationships within a group as simple ranks do, but also indicate possible reasons of differences in the behavioural development. For instance, whereas the dwarf goats maintained a quasi-linear dominance hierarchy over time with a high rate of overt agonistic behaviour, pigs after the establishment of their hierarchy showed a reduced agonistic behaviour which makes it questionable to calculate reliable sociometric measures. These species-dependent variations may be primarily caused by different kinds of the fighting behaviour in goats (i.e. ritualised, low costs) and pigs (i.e. more seriously, high costs). Overall, a more consistent and standardised approach of analysing social dominance in (farm) animals may improve the scientific value of single studies and makes it easier to compare various studies within a species and between species.
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Vervaecke, H., Stevens, J., Vandemoortele, H., Sigurjönsdöttir, H., & De Vries, H. (2007). Aggression and dominance in matched groups of subadult Icelandic horses (Equus caballus). J. Ethol., 25(3), 239–248.
Abstract: Abstract We studied sex differences in the nature of aggression and dominance behaviour in two newly formed groups of 1-year-old Icelandic horses. One herd contained nine geldings, the other nine mares. The groups were matched with regard to dominance-determining traits such as age, weaning age, composition of native herd, social experience, genetic origin, body condition and maternal dominance status. High-ranking individuals of both sexes were more aggressive, high-ranking males were older, and high-ranking females had a better body condition. Frequencies of aggressions were similar in both groups. The mares reacted significantly more by showing submission upon an aggression rather than by not responding or by escalating the aggression. For the geldings, this difference was not observed due to a lower tendency to submit. A linear dominance hierarchy was found in both groups. David`s scores provided additional information regarding cardinal rank distances and were used to calculate steepness of hierarchies. The female hierarchy was somewhat steeper compared to the male hierarchy, suggesting somewhat lower despotism among males. This was mainly a consequence of the lower unidirectionality in male submission. Male contests occurred predominantly between dyads at top and mid positions, suggesting a low degree of acceptance of the hierarchy.
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Pluhacek, J., Bartos, L., & Culik, L. (2006). High-ranking mares of captive plains zebra Equus burchelli have greater reproductive success than low-ranking mares. Appl. Anim. Behav. Sci., 99(3-4), 315–329.
Abstract: Plains zebra live in harems that include one to six adult mares. Between these mares is a strong order of social hierarchy. The social rank of an equid mare is typically correlated with her age. Further, high-ranking captive plains zebra mares produce more surviving offspring than low-ranking mares. The objectives of this study were to, first, examined the factors that influence social rank of captive plains zebra mares, and second, test if high-ranking mares conceive earlier and if they have shorter inter-birth intervals than low-ranking ones. We observed three herds of captive plains zebra (a total of 18 mares) at the Dvur Kralove Zoo, Czech Republic. During the 831 h of observation, we recorded 1713 aggressive interactions (biting and offensive kicking) between the mares. These data were used to determine, for each mare, the total number of mares that dominated her in each period of social stability. The GLMM model revealed that older mares were dominated by a lower number of mares than the younger mares. We also found that the probability that a mare would conceive declined with the increasing number of dominant mares. Further, we tested the relationship between the number of dominant mares and the inter-birth interval using 29 intervals for 15 mares. These inter-birth intervals were divided into two groups. When a stallion was continuously present in the herd, the intervals lasted from one birth to the next birth (natural intervals). When a herd was without a stallion, the intervals lasted from the release of the stallion into the herd to the birth of foal (stallion-influenced intervals). The analysis revealed that the inter-birth intervals decreased with an increasing number of dominant mares and the natural intervals decreased with an increasing number of offspring successfully reared by a mare. This finding is the first one in equids and contributes to the previous findings that suggest that social status influences reproductive success.
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Overli, O., Korzan, W. J., Hoglund, E., Winberg, S., Bollig, H., Watt, M., et al. (2004). Stress coping style predicts aggression and social dominance in rainbow trout. Horm Behav, 45(4), 235–241.
Abstract: Social stress is frequently used as a model for studying the neuroendocrine mechanisms underlying stress-induced behavioral inhibition, depression, and fear conditioning. It has previously been shown that social subordination may result in increased glucocorticoid release and changes in brain signaling systems. However, it is still an open question which neuroendocrine and behavioral differences are causes, and which are consequences of social status. Using juvenile rainbow trout of similar size and with no apparent differences in social history, we demonstrate that the ability to win fights for social dominance can be predicted from the duration of a behavioral response to stress, in this case appetite inhibition after transfer to a new environment. Moreover, stress responsiveness in terms of confinement-induced changes in plasma cortisol was negatively correlated to aggressive behavior. Fish that exhibited lower cortisol responses to a standardized confinement test were markedly more aggressive when being placed in a dominant social position later in the study. These findings support the view that distinct behavioral-physiological stress coping styles are present in teleost fish, and these coping characteristics influence both social rank and levels of aggression.
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