Social behaviour in stallion groups (Equus przewalskii and Equus caballus) kept under natural and domestic conditions
Introduction
Animals living under natural conditions are regarded as showing the normal behaviour of that species, and deviations from normal behaviour in domestic animals may relate to changes resulting from domestication, or they may be caused by environmental factors. Price (1984) defined domestication as ‘that process by which a population of animals becomes adapted to man and to the captive environment by some combination of genetic changes occurring over generations and environmentally induced developmental events reoccurring during each generation’. Domestic animals generally show reduced responsiveness to environmental changes, which may not only be expressed in their reaction towards humans, but also in intra-specific reactions and investigative behaviours. Domestication may additionally induce a change in group spacing behaviour since management practices, which minimise competition for resources, should permit captive populations to exist at relatively high densities (Price, 1984).
Studies of feral horses living under natural conditions indicate that the species-specific behaviour of equids has remained relatively unaffected by the domestication process (Tyler, 1972, Feist and McCullough, 1976, Waring, 1983), and thus domestication seems to have influenced the quantitative rather than the qualitative nature of social behaviour in horses. Quantitative changes may include a reduction in aggressive behaviour because the domestic environment reduces the selective advantages of aggressive potential in securing resources.
Direct comparisons of social behaviour in domestic and non-domestic horses are lacking in the literature, and are made difficult by the fact that domestic horses are kept under conditions, which are very different to the natural environment. Typical rearing conditions for domestic horses include early weaning of foals, commonly at the age of 5–6 months, after which they may experience either individual housing conditions or enter groups with other similarly aged horses. In sharp contrast to these conditions, studies on horses in natural environments show that horses tend to live in mixed age groups and stay with their natal herd till approximately 2 years of age (for a review see Waring, 1983). The rearing environment and the environment in which an animal lives is likely to affect the behaviour of the individual. For instance, it has been shown that social restrictions affect the behaviour of horses (Christensen et al., 2002), and rearing animals in physical isolation from older, socially dominating conspecifics may preclude the development of normal adult-like behaviour patterns of agonistic behaviours and may result in the retention of juvenile social behaviours (Price, 1984).
Feh (1988) and Keiper and Receveur (1992) reported more aggressive behaviour in groups of Przewalski horses, which is the closest wild relative of domestic horses, than in feral horse groups. However, these findings were based upon a limited number of stallions (three Przewalski stallions versus eight Camargue stallions by Feh, 1988), or upon comparisons between different studies (Keiper and Receveur, 1992). The present study aims to extend these earlier investigations by studying social behaviour in Przewalski and domestic stallions within the same study, with a large number of stallions per group, and using the same observational techniques and observers on both groups.
Investigations on behavioural differences between domestic horses and their wild relatives—displayed in their respective home environments—are of great importance in understanding the span of equid behaviour. The present study investigates and discusses social and group spacing behaviour in domestic stallions and in non-domestic Przewalski stallions, reared under typical domestic and natural conditions, respectively.
Section snippets
Animals
The domestic group consisted of 19 2-year-old stallions, reared under typical domestic conditions at the Danish Institute of Agricultural Sciences. The stallions were weaned at approximately 4 months of age and housed either individually (n=7) or in groups of three (n=12) with daily access to paddocks during the autumn, winter and spring season and were pastured together during the summer season. The stallions were pastured in two separate groups (n=7 and n=12), which were joined into one group
Results
Table 2 shows the occurrence of the recorded social interactions in the two stallion groups. The naturally reared Przewalski stallions tended to show more agonistic interactions than the domestically reared stallions (Table 2), significant only for kick threats (t(30)=3.54; P=0.001) and kicks (MWU, P=0.001). The domestic stallions showed significantly more pushing behaviour (MWU, P=0.016), although the occurrence was very low in both groups. They likewise tended to show more mouth clapping
Discussion
A higher number of social interactions was recorded in the naturally reared and mixed age group of Przewalski stallions than in the domestically reared and similarly aged group of domestic stallions. Consistently, a significantly higher level of social grooming was recorded in the Przewalski group. Other authors have found that levels of social grooming vary with season, with peaks during the spring period, due to the shedding of the winter coat (Tyler, 1972, Kimura, 1998, Zharkikh, 2000). The
Conclusions
The qualitative nature of social behaviours was alike in domestic and Przewalski stallions, and the same behavioural definitions could be applied to register social behaviour in both groups. Play and play fight behaviour was both qualitatively and quantitatively very similar in the two stallion groups. Quantitative differences were found in social grooming and in investigative behaviours but these may be attributed to environmental factors and in the time since group formation. Quantitative
Acknowledgements
Thanks to Jens Malmkvist and Eva Sondergaard, Department of Animal Health and Welfare, DIAS, for comments on earlier versions of the manuscript, and to Erik L. Decker, DIAS, for technical assistance. Thanks to Julia Alexeyeva and Alla Nikitina, Kyiv State Zoological Park and to Helen Kraevoj, Biosphere Reserve Askania-Nova.
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