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Keiper, R. R., & Keenan, M. A. (1980). Nocturnal activity patterns of feral horses. J. Mammal, 61, 116–118.
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Keiper, R. R., Moss, M., & Zervanos, S. (1980). Daily and seasonal patterns of feral ponies on Assateague Island. In 2nd Conference on Scientific Research in the National Parks.
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Keiper, R. R. (1979). The behaviour, ecology and social organization of the feral ponies of Assateague Island. Proc. I. Conf. Sci. Res. Nat. Parks,, .
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Keiper, R. R. (1988). Social interactions of the Przewalski horse (Equus przewalskii Poliakov, 1881) herd at the Munich Zoo. Appl. Anim. Behav. Sci., 21(1-2), 89–97.
Abstract: Data were collected on 972 aggressions and 233 acts of mutual grooming in a herd of 9 Przewalski horses in the Hellabrunn Tierpark in Munich, West Germany. The herd was composed of 1 adult stallion, 5 adult mares and 3 foals. A distinct linear dominance hierarchy was present in the herd, with the stallion being the top-ranking animal. Age was significantly correlated with rank. Almost 40% of all aggressions consisted of herding actions by the stallion. Threats to bite (20% of all aggressions) and threats to kick (11.4%) were next in frequency of occurrence. Most mutual grooms (71%) involved grooming the front part of the body. Although mutual grooming may be used to appease higher-ranked animals, most grooming bouts were between related horses. Foals initiate 47.6% of all allogrooming. Mutual grooming may reduce weaning conflict between a mare and her foal or may result in female coalitions that defend against predators or aggression by the herd stallion.
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Keiper, R. R., & Sambraus, H. H. (1986). The stability of equine dominance hierarchies and the effects of kinship, proximity and foaling status on hierarchy rank. Appl. Anim. Behav. Sci., 16(2), 121–130.
Abstract: Dominance hierarchies were determined in four bands of feral horses living on Assateague Island. The bands varied in size from 10 to 16 horses, and consisted of one stallion, several mares and their offspring. The animals ranged in age from less than 1 to over 18 years. Field observation of all social interactions during the summer of 1981 was used to determine dominance. 1981 hierarchies for three of the bands were compared with hierarchies determined for the same bands in 1978, and showed that hierarchies change over time. Age was significantly correlated with rank. Mares with foals did not rank any higher in the hierarchies than mares without foals. Kinship did not appear to have an effect on dominance rank either, since neither juvenile nor adult offspring ranks correlated with the ranks of their mothers. The band stallion was not the highest-ranking animal of any band, but the location of the stallion peripheral to the main body of the band, the nature of his interactions with band members, and his length of residence in the band may have contributed to his low rank.
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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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Rutberg, A. T., & Keiper, R. R. (1993). Proximate causes of natal dispersal in feral ponies: some sex differences. Anim. Behav., 46(5), 969–975.
Abstract: Abstract. Fifteen years of data on natal dispersal age and the context of dispersal for the feral ponies of Assateague Island, Maryland are presented. Ninety-seven per cent of males and 81% of females dispersed from their natal groups by 5 years of age. For animals that left their natal group, average age of dispersal was 20[middle dot]8 months for males and 24[middle dot]6 months for females. Male dispersal age was strongly and significantly correlated with number of peers in the natal group, and males dispersing with peers were significantly older than males dispersing without peers, suggesting that males delayed dispersal when peers were available for interaction. Female dispersal age was not influenced by number of peers, but was correlated with age of first reproduction. Factors not influencing dispersal age in either sex were presence of a younger sibling, maternal band transfers, and maternal age and dominance rank. The relatively high frequency of females failing to disperse from their natal groups is puzzling in light of data showing diminished fecundity in non-dispersing pony mares.
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