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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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Collery, L. (1974). Observations of equine animals under farm and feral conditions. Equine Vet J, 6(4), 170–173.
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Kiley, M. (1972). The vocalizations of ungulates, their causation and function. Z. Tierpsychol., 31(2), 171–222.
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Klingel, H. (1982). Social organization of feral horses. J Reprod Fertil Suppl, 32, 89–95.
Abstract: The basic social unit in feral horses is the family group consisting of one stallion, one to a few unrelated mares and their foals. Surplus stallions associate in bachelor groups. Stallions are instrumental in bringing mares together in a unit which then persists even without a stallion. The similarity of social organization in populations living in a variety of different habitats indicates that feral horses have reverted to the habits of their wild ancestors, and that domestication has had no influence on this basic behavioural feature.
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Feist, J. D., & McCullough, D. R. (1975). Reproduction in feral horses. J Reprod Fertil Suppl, (23), 13–18.
Abstract: A behavioural study of feral horses was conducted on the Pryor Mountain Wild Horse Range in the western United States. All 270 horses on the Range were identified individually. The sex ratio was nearly balanced. Foal to adult female ratio was 43-2:100. Morality was concentrated among foals and old horses. Horses were organized as forty-four harem groups each with a dominant stallion, one to two immature stallions, one to three immature mares, one to three adult mares and their yearling and foal offspring, and 23 bachelor groups of one to eight stallions. Harem groups were quite stable year-round because of dominance and leadership by the stallions and group fidelity by mares and their offsring. Most changes occurred during the breeding season and involved immature females. Defeat of dominant stallions was infrequent. Immature males were tolerated because of their submissive behaviour. Bachelor stallion groups were inherently unstable. Mares came into heat after foaling in May/June, and were mated by harem stallions only.
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Kawamura, S. (1967). Aggression as studied in troops of Japanese monkeys. UCLA Forum Med Sci, 7, 195–223.
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Fricke, H. W. (1973). Individual partner recognition in fish: field studies on Amphiprion bicinctus. Naturwissenschaften, 60(4), 204–205.
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Ganswindt, A., Palme, R., Heistermann, M., Borragan, S., & Hodges, J. K. (2003). Non-invasive assessment of adrenocortical function in the male African elephant (Loxodonta africana) and its relation to musth. Gen Comp Endocrinol, 134(2), 156–166.
Abstract: Adult male elephants periodically show the phenomenon of musth, a condition associated with increased aggressiveness, restlessness, significant weight reduction and markedly elevated androgen levels. It has been suggested that musth-related behaviours are costly and that therefore musth may represent a form of physiological stress. In order to provide data on this largely unanswered question, the first aim of this study was to evaluate different assays for non-invasive assessment of adrenocortical function in the male African elephant by (i) characterizing the metabolism and excretion of [3H]cortisol (3H-C) and [14C]testosterone (14C-T) and (ii) using this information to evaluate the specificity of four antibodies for determination of excreted cortisol metabolites, particularly with respect to possible cross-reactions with androgen metabolites, and to assess their biological validity using an ACTH challenge test. Based on the methodology established, the second objective was to provide data on fecal cortisol metabolite concentrations in bulls during the musth and non-musth condition. 3H-C (1 mCi) and 14C-T (100 microCi) were injected simultaneously into a 16 year old male and all urine and feces collected for 30 and 86 h, respectively. The majority (82%) of cortisol metabolites was excreted into the urine, whereas testosterone metabolites were mainly (57%) excreted into the feces. Almost all radioactive metabolites recovered from urine were conjugated (86% 3H-C and 97% 14C-T). In contrast, 86% and >99% of the 3H-C and 14C-T metabolites recovered from feces consisted of unconjugated forms. HPLC separations indicated the presence of various metabolites of cortisol in both urine and feces, with cortisol being abundant in hydrolysed urine, but virtually absent in feces. Although all antibodies measured substantial amounts of immunoreactivity after HPLC separation of peak radioactive samples and detected an increase in glucocorticoid output following the ACTH challenge, only two (in feces against 3alpha,11-oxo-cortisol metabolites, measured by an 11-oxo-etiocholanolone-EIA and in urine against cortisol, measured by a cortisol-EIA) did not show substantial cross-reactivity with excreted 14C-T metabolites and could provide an acceptable degree of specificity for reliable assessment of glucocorticoid output from urine and feces. Based on these findings, concentrations of immunoreactive 3alpha,11-oxo-cortisol metabolites were determined in weekly fecal samples collected from four adult bulls over periods of 11-20 months to examine whether musth is associated with increased adrenal activity. Results showed that in each male levels of these cortisol metabolites were not elevated during periods of musth, suggesting that in the African elephant musth is generally not associated with marked elevations in glucocorticoid output. Given the complex nature of musth and the variety of factors that are likely to influence its manifestation, it is clear, however, that further studies, particularly on free-ranging animals, are needed before a possible relationship between musth and adrenal function can be resolved. This study also clearly illustrates the potential problems associated with cross-reacting metabolites of gonadal steroids in EIAs measuring glucocorticoid metabolites. This has to be taken into account when selecting assays and interpreting results of glucocorticoid metabolite analysis, not only for studies in the elephant but also in other species.
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