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BASHORE, T. L., KEIPER, R., TURNER, J. W. J. R., & KIRKPATRICK J. F. (1990). The accuracy of fixed-wing aerial surveys of feral horses on a coastal barrier island. J. coast. res, 6, 53–56.
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Boyd, L., & Keiper, R. (2005). Behavioural ecology of feral horses. In D. S. Mills, & McDonnell S. M. (Eds.), The domestic horse: the origins, development, and management of its behaviour. Cambridge: Cambridge University Press.
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Houpt, K. A., & Keiper, R. (1982). The position of the stallion in the equine dominance hierarchy of feral and domestic ponies. J. Anim Sci, 54(5), 945–950.
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Keiper, R. (1976). Social organization of feral ponies. Proc Pennsyl Acad Sci, 50, 69–70.
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Keiper, R., & Houpt, K. (1984). Reproduction in feral horses: an eight-year study. Am J Vet Res, 45(5), 991–995.
Abstract: The reproductive rate and foal survival of the free-ranging ponies on Assateague Island National Seashore were studied for 8 years, 1975 to 1982. Most (52%) of the 86 foals were born in May, 13% were born in April, 22.6% in June, 10.4% in July, and less than 1% in August and September. The mean foaling rate was 57.1 +/- 3.9% and the survival rate was 88.3 +/- 3.6%. Forty-eight colts and 55 fillies were born (sex ratio 53% female). Mares less than 3 years old did not foal and the foaling rate of 3-year-old mares was only 23%, that of 4-year-old mares was 46%, that of 5-year-old mares was 53%, and 6-year-old mares was 69%. The relatively poor reproduction rate was believed to be a consequence of the stress of lactating while carrying a foal when forage quality on the island was low. The hypothesis was supported by the higher reproductive rate (74.4 +/- 2.4%) of the ponies in the Chincoteague National Wildlife Refuge on the southern part of the island. Their foals are weaned and sold in July each year. Despite the low reproductive rate on Assateague Island National Seashore , the number of ponies increased from 43 to 80, a 90% increase in the 8-year period or greater than 10%/yr. There were 24 deaths and 8 dispersals from the study area.
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Keiper, R., & Receveur, H. (1992). Social interactions of free-ranging Przewalski horses in semi-reserves in the Netherlands. Appl. Anim. Behav. Sci., 33(4), 303–318.
Abstract: Social interactions were recorded in two bands of free-ranging Przewalski horses living on large (greater than 30 ha) pastures in the Netherlands. The average number of aggressive interactions per hour was 8.86 at Lelystad and 10.36 at Noorderheide. The most common aggressive interactions were lower intensity, lower cost displacements (17.2% of all aggressive acts at Lelystad, 13.2% at Noorderheide), threats to bite (42.3% and 40.7%, respectively) and threats to kick (15.4% and 23.9%, respectively). Analysis of aggression revealed that a clear, linear dominance hierarchy was present in each band. For each band there was a positive and highly significant correlation between the age of a horse and its rank in the hierarchy. In each band, the stallion was not the highest ranked horse. Non-agonistic behaviors exceeded the number of agonistic interactions (1253 vs. 558 for Lelystad; 1257 vs. 995 at Noorderheide). There was a negative correlation between the rank of a horse in the dominance hierarchy and the number of non-agonistic behaviors displayed. The group displaying the highest number of non-agonistic interactions were foals (48.9% of total non-agonistic behaviors at Lelystad; 51.1% at Noorderheide). The non-agonistic/agonistic ratio was greater than 1 for yearlings and the band stallion, as was also the case for foals, but was less than 1 for males.
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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. (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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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., & Keenan, M. A. (1980). Nocturnal activity patterns of feral horses. J. Mammal, 61, 116–118.
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