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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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Stahlbaum, C. C., & Houpt, K. A. (1989). The role of the Flehmen response in the behavioral repertoire of the stallion. Physiol. Behav., 45(6), 1207–1214.
Abstract: The role of the Flehmen response in equine behavior was investigated under field and laboratory conditions. In Experiment 1, a field study made of five stallions on pasture with between three and eighteen mares each during the season indicated the following: 1) The Flehmen response was most frequently preceded by nasal, rather than oral, investigation of substances; 2) The stallions' rate of Flehmen varied with the estrous cycles of the mares; 3) The rate of Flehmen response did not show a variation with time of day; and 4) The Flehmen response was most frequently followed by marking behaviors rather than courtship behaviors. The results suggest that the Flehmen response is not an immediate component of sexual behavior, e.g., courtship of the stallion but may be involved in the overall monitoring of the mare's estrous cycle. Therefore the Flehmen response may contribute to the chemosensory priming of the stallion for reproduction. In Experiment 2 stallions were presented with urine or feces of mares in various stages of the reproductive cycle as well as with their own or other males' urine or feces. The occurrence of sniffing and Flehmen was used to determine the discriminatory ability of the stallions. Stallions can differentiate the sex of a horse on the basis of its feces alone, but cannot differentiate on the basis of urine. This ability may explain the function of fecal marking behavior of stallions.
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Houpt, K. A. (1976). Animal behavior as a subject for veterinary students. Cornell Vet, 66(1), 73–81.
Abstract: Knowledge of animal behavior is an important asset for the veterinarian; therefore a course in veterinary animal behavior is offered at the New York State College of Veterinary Medicine as an elective. The course emphasizes the behavior of those species of most interest to the practicing veterinarian: cats, dogs, horses, cows, pigs and sheep. Dominance heirarchies, animal communication, aggressive behavior, sexual behavior and maternal behavior are discussed. Play, learning, diurnal cycles of activity and sleep, and controls of ingestive behavior are also considered. Exotic and zoo animal behaviors are also presented by experts in these fields. The critical periods of canine development are related to the optimum management of puppies. The behavior of feral dogs and horses is described. The role of the veterinarian in preventing cruelty to animals and recognition of pain in animals is emphasized. Whenever possible behavior is observed in the laboratory or on film.
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Khalil, A. M., Murakami, N., & Kaseda, Y. (1998). Relationship between plasma testosterone concentrations and age, breeding season and harem size in Misaki feral horses. J Vet Med Sci, 60(5), 643–645.
Abstract: Jugular vein blood samples were collected from 23 young and sexual mature feral stallions to examine the relationship between plasma testosterone concentration and age, breeding season or harem size. Testosterone concentration increased with the age of the stallions until they formed their own harems, at about 4 to 6 years old. Seasonal variations in testosterone concentrations were observed, and found to be significantly higher (P<0.001) throughout the breeding season than non-breeding season, from 3 years of age. Testosterone levels were correlated with harem size for individual stallions. It can be inferred from these results that there is a relationship between plasma testosterone concentration and age, breeding season and harem size.
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Turner, J. W. J., & Kirkpatrick, J. F. (1982). Androgens, behaviour and fertility control in feral stallions. J Reprod Fertil Suppl, 32, 79–87.
Abstract: This field study of feral stallions in Montana and Idaho examines and correlates the seasonal pattern of plasma androgens and specific sociosexual behaviour and reports the effect of a long-acting androgenic steroid on this behaviour and on fertility. Plasma testosterone was measured by competitive protein binding assay in samples obtained by jugular venepuncture from captured animals. In samples taken from 34 sexually mature stallions in 6 different months during the year, a definite seasonal pattern in testosterone was present, with a peak in May (3.04 +/- 0.63 ng/ml) and a nadir in December (1.55 +/- 0.34 ng/ml). Values were less than 2.0 ng/ml in non-breeding months and greater than 2.4 ng/ml in breeding months. Behavioural endpoints measured were (1) stallion scent marking in response to elimination by mares (elimination marking), (2) mounting and (3) copulation. The frequencies of each of these endpoints followed closely the seasonal pattern seen for plasma androgens. In the fertility study microcapsulated testosterone propionate (microTP) was administered i.m. to 10 harem stud stallions 3 months before the 1980 breeding season. In these stallions and in 10 control harem studs, the above behavioural endpoints were examined in the 1980 and 1981 breeding seasons, and foal counts were made in 1981. There were no direct inhibitory or stimulatory effects of microTP treatment on any of the behavioural endpoints in either year. In 1981 foals were produced in 87.5% of the control bands and 28.4% of the microTP-treated bands. These results indicate that microencapsulated testosterone propionate can provide effective fertility control in feral horses without causing significant alterations in sociosexual behaviour.
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Kirkpatrick, J. F., Vail, R., Devous, S., Schwend, S., Baker, C. B., & Wiesner, L. (1976). Diurnal variation of plasma testosterone in wild stallions. Biol Reprod, 15(1), 98–101.
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de Waal, F. B. M. (2005). A century of getting to know the chimpanzee. Nature, 437(7055), 56–59.
Abstract: A century of research on chimpanzees, both in their natural habitat and in captivity, has brought these apes socially, emotionally and mentally much closer to us. Parallels and homologues between chimpanzee and human behaviour range from tool-technology and cultural learning to power politics and intercommunity warfare. Few behavioural domains have remained untouched by this increased knowledge, which has dramatically challenged the way we view ourselves. The sequencing of the chimpanzee genome will no doubt bring more surprises and insights. Humans do occupy a special place among the primates, but this place increasingly has to be defined against a backdrop of substantial similarity.
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Parish, A. R., & De Waal, F. B. (2000). The other “closest living relative”. How bonobos (Pan paniscus) challenge traditional assumptions about females, dominance, intra- and intersexual interactions, and hominid evolution. Ann N Y Acad Sci, 907, 97–113.
Abstract: Chimpanzee (Pan troglodytes) societies are typically characterized as physically aggressive, male-bonded and male-dominated. Their close relatives, the bonobos (Pan paniscus), differ in startling and significant ways. For instance, female bonobos bond with one another, form coalitions, and dominate males. A pattern of reluctance to consider, let alone acknowledge, female dominance in bonobos exists, however. Because both species are equally “man's” closest relative, the bonobo social system complicates models of human evolution that have historically been based upon referents that are male and chimpanzee-like. The bonobo evidence suggests that models of human evolution must be reformulated such that they also accommodate: real and meaningful female bonds; the possibility of systematic female dominance over males; female mating strategies which encompass extra-group paternities; hunting and meat distribution by females; the importance of the sharing of plant foods; affinitive inter-community interactions; males that do not stalk and attack and are not territorial; and flexible social relationships in which philopatry does not necessarily predict bonding pattern.
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de Waal, F. B. (1996). Macaque social culture: development and perpetuation of affiliative networks. J Comp Psychol, 110(2), 147–154.
Abstract: Maternal affiliative relations may be transmitted to offspring, similar to the way in which maternal rank determines offspring rank. The development of 23 captive female rhesus monkeys (Macaca mulatta) was followed from the day of birth until adulthood. A multivariate analysis compared relations among age peers with affiliative relations, kinship, and rank distance among mothers. Maternal relations were an excellent predictor of affiliative relations among daughters, explaining up to 64% of the variance. Much of this predictability was due to the effect of kinship. However, after this variable had been controlled, significant predictability persisted. For relations of female subjects with male peers, on the other hand, maternal relations had no significant predictive value beyond the effect of kinship. One possible explanation of these results is that young rhesus females copy maternal social preferences through a process of cultural learning.
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de Waal, F. B. (1995). Bonobo sex and society. Sci Am, 272(3), 82–88.
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