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Klingel, H. (1975). Social organization and reproduction in equids. J Reprod Fertil Suppl, (23), 7–11.
Abstract: There are two distinct types of social organization and, accordingly, two types of mating systems in equids. In the horse, Plains zebra and Mountain zebra, the adults live in non-territorial and cohesive one-male groups and in stallion groups. The family stallions have exclusive mating rights which are respected by all others. In Grevy's zebra and in the African and Asiatic wild asses, the stallions are permanently territorial and have exclusive mating rights within their territories. Ecological and evolutionary aspects are discussed.
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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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Aurich, J., Wulf, M., Ille, N., Erber, R., von Lewinski, M., Palme, R., et al. (2015). Effects of season, age, sex and housing on salivary cortisol concentrations in horses. Domest. Anim. Endocrinol., .
Abstract: Abstract Analysis of salivary cortisol is increasingly used to assess stress responses in horses. Since spontaneous or experimentally induced increases in cortisol concentrations are often relatively small for stress studies proper controls are needed. This requires an understanding of factors affecting salivary cortisol over longer times. In this study, we have analysed salivary cortisol concentration over 6 mo in horses (n = 94) differing in age, sex, reproductive state and housing. Salivary cortisol followed a diurnal rhythm with highest concentrations in the morning and a decrease throughout the day (P < 0.001). This rhythm was disrupted in individual groups on individual days; however, alterations remained within the range of diurnal changes. Comparison between months showed highest cortisol concentrations in December (P < 0.001). Cortisol concentrations increased in breeding stallions during the breeding season (P < 0.001). No differences in salivary cortisol concentrations between non-pregnant mares with and without a corpus luteum existed. In stallions, mean daily salivary cortisol and plasma testosterone concentration were weakly correlated (r = 0.251, P < 0.01). No differences in salivary cortisol between female and male young horses and no consistent differences between horses of different age existed. Group housing and individual stabling did not affect salivary cortisol. In conclusion, salivary cortisol concentrations in horses follow a diurnal rhythm and are increased in active breeding sires. Time of the day and reproductive state of the horses are thus important for experiments that include analysis of cortisol in saliva.
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Kirkpatrick, J. F., & Turner, A. (2003). Absence of effects from immunocontraception on seasonal birth patterns and foal survival among barrier island wild horses. J Appl Anim Welf Sci, 6(4), 301–308.
Abstract: Despite a large body of safety data, concern exists that porcine zonae pellucidae (PZP) immunocontraception--used to manage wild horse populations--may cause out-of-season births with resulting foal mortality. Our study at Assateague, Maryland indicated the effects of immunocontraception on season of birth and foal survival between 1990 and 2002 on wild horses from Assateague Island. Among 91 mares never treated, 69 (75.8%) of foals were born in April, May, and June (in season). Among 77 treated mares, 50 (64.9%) were born in season. Of 29 mares foaling within 1 year after treatment (contraceptive failures), 20 (68.9%) were born in season. Of 48 mares treated for greater than 2 years then withdrawn from treatment, 30 (62.5%) of 48 foals were born in season. There were no significant differences (p <.05) between either treatment group or untreated mares. Survival did not differ significantly among foals born in or out of season or among foals born to treated or untreated mares. Data indicate a lack of effect of PZP contraception on season of birth or foal survival on barrier island habitats.
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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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Detmer, D. (1992). Response: of pigs and primitive notions. Between Species, 8(4), 203–208.
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Nelson, W. A., Keirans, J. E., Bell, J. F., & Clifford, C. M. (1975). Host-ectoparasite relationships. J Med Entomol, 12(2), 143–166.
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Henneke, D. R., Potter, G. D., Kreider, J. L., & Yeates, B. F. (1983). Relationship between condition score, physical measurements and body fat percentage in mares. Equine Vet J, 15(4), 371–372.
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Carlstead, K., & Brown, J. L. (2005). Relationships between patterns of Fecal corticoid excretion and behavior, reproduction, and environmental factors in captive black (Diceros bicornis) and white (Ceratotherium simum) rhinoceros. Zoo Biol., 24(3), 215–232.
Abstract: Mortality is high in zoo-housed black rhinoceros (Diceros bicornis), and the reproductive rates of captive white rhinoceros (Ceratotherium simum) are unsustainably low. To determine the possible role of stress in the causation of these problems, we analyzed weekly fecal samples collected for 1 year from black (10 males and 16 females) and white (six males and 13 females) rhinoceroses at 16 zoos for corticoid metabolite concentrations. Fecal corticoid profiles were examined in relation to behavior as rated by keepers in a questionnaire, luteal phase ovarian cycles of females (Brown et al., 2001), and socioenvironmental factors. We compared individual fecal corticoid profiles by examining hormone means and variability (i.e., standard deviation (SD) and coefficient of variation (CV)). For the black rhinos, higher mean corticoid concentrations were found at zoos where rhinos were maintained in enclosures that were exposed to the public around a greater portion of the perimeter. Higher variability in corticoid excretion was correlated with higher rates of fighting between breeding partners and higher institutional mortality rates. Black rhino pairs that were kept separated exhibited lower corticoid variability and less fighting activity when they were introduced during female estrous periods compared to pairs that were kept together every day. For white rhinos, significantly lower mean corticoids were found for individuals that rated higher on “friendliness to keeper.” Higher corticoid variability was found in noncycling as compared to cycling white rhino females. Noncycling females exhibited higher rates of stereotypic pacing and lower frequencies of olfactory behaviors. Interindividual differences in mean corticoids in both species appeared to be related to responsiveness to humans, whereas corticoid variability was related to intraspecific social relationships. More importantly, high corticoid variability appeared to be an indicator of chronic or “bad” stress, because of its association with potentially deleterious consequences in each species (i.e., fighting and mortality (black rhino), and reproductive acyclicity (white rhino)). Our results provide evidence that social stressors may cause chronic stress in black and white rhinos, and that this contributes to the captive-population sustainability problems observed in each species. Zoo Biol 0:1–18, 2005. © 2005 Wiley-Liss, Inc.
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Kaseda, Y., Ogawa, H., & Khalil, A. M. (1997). Causes of natal dispersal and emigration and their effects on harem formation in Misaki feral horses. Equine Vet J, 29(4), 262–266.
Abstract: Misaki feral horses were separated into 2 herds and the difference between dispersal from natal group (natal dispersal) and dispersal from natal area (natal emigration) was studied. The causes of dispersal and emigration and their effects on harem formation were studied 1979-1994. The number of horses ranged from 73 (mature males: 8, mature females: 26, young males: 8, young females: 3, colt foals: 6, filly foals: 10 and geldings: 12) in 1979 and 86 (mature males: 14, mature females: 37, young males: 12, young females: 7, colt foals: 5, filly foals: 7 and geldings: 4) in 1994 when the present study ended. All 29 males which survived to age 4 years and 58 females which survived to age 3 years left their natal or mother groups at age one to 3. Seventeen of 22 dispersing males and 29 of 39 dispersing females left their natal groups around the birth of their siblings and significant correlations were found between natal dispersal and birth of a sibling. The number of emigrating young males correlated negatively and significantly with the total number of young males in another herd and the number of emigrating young females correlated positively and significantly with the total number of young females in the natal herd. All 13 emigrating stallions which survived to age 5 years formed stable harem groups and a significant correlation was found between natal emigration and harem formation. Twenty-three of 35 resident mares formed stable consort relations with harem stallions and a significant correlation was found between residence and formation of stable consort relations.
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