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Araba, B. D., & Crowell-Davis, S. L. (1994). Dominance relationships and aggression of foals (Equus caballus). Appl. Anim. Behav. Sci., 41(1-2), 1–25.
Abstract: Studied a herd of 15 Belgian brood-mares and 10 foals. Specific aspects of social structure studied were dominance-subordinance relationships, preferred associates, social spacing, aggression rates, the frequency of aggressions administered down the dominance hierarchy, and interactive play bouts. The rank order of the foals, both before and after weaning, was positively correlated with the rank order of their dams. There was also a significant relationship between a foal's rank and its total aggression or aggression rate per subordinate post-weaning. Higher ranking foals had higher rates of aggression. Over 80% of threats were directed down the dominance hierachy. The play-rank order of the foals, scored by the number of times foal left a play bout, was not significantly correlated with the rank order as scored by agonistic interactions. -from Authors
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Pickerel, T. M., Crowell-Davis, S. L., Caudle, A. B., & Estep, D. Q. (1993). Sexual preference of mares (Equus caballus) for individual stallions. Appl. Anim. Behav. Sci., 38(1), 1–13.
Abstract: Eight mares were tested to determine if they remained near one of two stallions longer than would be expected if association was random. Six stallions were paired in 30 combinations and each mare was tested 30 times. The mares (Equus caballus) demonstrated a definite preference for individual stallions throughout the breeding season. This preference was influenced by the estrous state of the mare. During estrus, mares' preferences for stallions were positively correlated with the rate at which a given stallion vocalized. During diestrus, mares spent significantly less time in the proximity of stallions and did not exhibit any preference for individual stallions.
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Crowell-Davis, S. L. (1985). Nursing behaviour and maternal aggression among Welsh ponies (Equus caballus). Appl Anim Behav Sci, 14(1), 11–25.
Abstract: Nursing behaviour and related aggression of mare-foal pairs was studied from birth (n = 21) to 24 weeks of age (n = 15) of the foal. Foals exhibited a decreasing length and frequency of nursing as they grew older. Mares rarely aggressed against their foals during nursing in the foal's first 4 weeks of life, but did so increasingly through Weeks 13-16, after which the rate of aggression during nursing decreased. Mares terminated nursing primarily by moving away, and were most likely to do so during the foal's first 4 weeks of life. They became gradually less likely to do so as the foal grew older. It was concluded that mares sometimes flex their hind limb on the side opposite the foal during nursing in order to conserve energy in a situation in which they would be remaining still anyway. There was no difference between colts and fillies in the frequency or duration of nursing or in the frequency with which their mothers aggressed against them or terminated nursing.
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Sharon, L., & Crowell-Davis, S. L. (2007). Sexual behavior of mares. Horm Behav, 52(1), 12–17.
Abstract: The mare is seasonally polyestrus, having an anovulatory period during the short light days of late fall and early winter, and beginning to ovulate as the days become longer during the winter. The complete estrus cycle is typically about 3 weeks, with 5 to 7 days of estrus and approximately 2 weeks of diestrus. When a mare lives within the natural social structure of the horse, i.e. a family band with several adult mares and one or more stallions, estrus is characterized by repeatedly approaching the stallion, frequent urination, deviating the tail away from the perineum, and standing still with the hind limbs spread apart. Diestrus is characterized by avoidance of an approaching stallion, and aggression toward the stallion, such as squealing, striking, and kicking, if he persists in attempting to court the diestrus mare. However, mares and stallions with long-term social relationships will often rest together, graze together and groom each other, all without sexual interactions. Hormonally, estrous behavior in the mare is initiated by estradiol that is secreted by the follicle, while estrous behavior is suppressed by progesterone, secreted by the corpus luteum. Mares are unusual among the ungulates in that they periodically exhibit estrous behavior during the anovulatory period. This is probably due to the release of estrogenic steroids secreted by the adrenal cortex. The display of sexual behavior by the mare throughout the year is thought to facilitate maintenance of the horse's social structure, in which the male remains with a group of females year round, in contrast with most ungulates in which the females and males only come together during the mating season.
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Crowell-Davis, S. L., & Houpt, K. A. (1985). Coprophagy by foals: effect of age and possible functions. Equine Vet J, 17(1), 17–19.
Abstract: In colts and fillies observed from birth to 24 weeks old, coprophagy occurred from Weeks 1 to 19. Its frequency was greatest during the first two months. Coprophagy was rarely observed in mares and stallions. Foals usually ate the faeces of their mother but were observed to eat their own and those of a stallion and another unrelated mare. Urination by the foal occurred before, during or after 26 per cent of the coprophagy incidents. It is hypothesised that foals may consume faeces in response to a maternal pheromone which signals the presence of deoxycholic acid or other acids which the foal may be deficient in and which it may require for gut immuno-competence myelination of the nervous system. Such a pheromone may also serve to accelerate growth and sexual maturation. Coprophagy may also provide nutrients and introduce normal bacterial flora to the gut.
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Ellard, M. - E., & Crowell-Davis, S. L. (1989). Evaluating equine dominance in draft mares. Appl. Anim. Behav. Sci., 24(1), 55–75.
Abstract: The social hierarchy of a herd of 12 draft mares was assessed using agonism in the field, paired-feeding tests and a group-feeding test. Results from the paired-feeding test correlated significantly, but imperfectly, with those from the field. Differential motivation among subjects for the feed and disruption of ambiguous relationships among mares reduced the reliability of the paired-feeding test as a measure of social dominance. Results from the group-feeding test did not correlate significantly with the field hierarchy and only a few mares ever ate from the bucket. Height, weight and age each correlated significantly with rank; a mare's tendency to remain alone did not. Total aggressive scores during the paired-feeding test correlated with rank. However, a high-ranking mare was no more aggressive to each of her subordinates than was a low-ranking mare. Rather, all mares aggressed more against individuals close in rank to themselves and with preferred field associates. In the field, mares associated most with other mares of similar rank.
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Hall, C., Crowell-Davis, S. L., & Warren, R. J. (1993). Maternal and developmental behavior of the feral horses of Cumberland Island, Georgia. Appl. Anim. Behav. Sci., 37(1), 85.
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Smith-Funk, E. D., & Crowell-Davis, S. L. (1992). Maternal behavior of draft mares (Equus caballus) with mule foals (Equus asinus x Equus caballus). Appl. Anim. Behav. Sci., 33(2-3), 93–119.
Abstract: Draft mares and their mule foals were observed from the day of birth to Week 17 of each foal's life. The rate of nursing was recorded and the duration of nursing activity to the nearest second. The rate at which foals engaged in nursing activity varied at each age. The duration of nursing bouts varied slightly as the foals matured. Aggression was recorded during both nursing and non-nursing activity for both the mares and foals. Maternal aggression was highest during nursing activity, especially during the pre-nurse nuzzling period. Maternal aggression increased as the foals matured. Mother-directed foal aggression was primarily in response to maternal aggression. Spatial relationships between each focal dyad were recorded when the foals were upright, not nursing and when they were recumbent. Spatial relationships differed based on the foal's state. The activity in which the mare engaged while her foal was recumbent was recorded. The movements of the mares were also recorded during foal recumbency. Mares approached or maintained their distance from their recumbent foal more than they left their recumbent foal in all weeks of the study, except Week 2.
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Crowell-Davis, S. L., Houpt, K. A., & Carini, C. M. (1986). Mutual grooming and nearest-neighbor relationships among foals of Equus caballus. Appl. Anim. Behav. Sci., 15(2), 113–123.
Abstract: A 3-year study was carried out on the developmental behavior of foals from birth to 24 weeks of age and the behavior of mares living with foals. Mutual-grooming partners of foals were primarily other foals. The peak frequency of mutual grooming occurred during Weeks 9-12, when fillies mutual-groomed 1.6 times h-1 and colts mutual-groomed 0.9 times h-1. Fillies mutual-groomed more frequently than colts (P < 0.025). Fillies mutual-groomed randomly with colts and other fillies (P < 0.05), whereas colts mutual-groomed almost exclusively with fillies (P = 0.03). At all ages studied, if a foal's nearest neighbor was not its mother, it was more likely to be another foal than would be expected if the foal was associating randomly with non-mother ponies. Fillies were more likely than expected to have a filly rather than a colt as their nearest neighbor (P = 0.01). Thus, during their first few months of life, the foals studied exhibited patterns of behavior which were consistent with the development of the usual social milieu of unmanaged adults, in which several mares form a cohesive herd with one or more stallions associating with them.
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Crowell-Davis, S. L., Houpt, K. A., & Kane, L. (1987). Play development in Welsh pony (Equus caballus) foals. Appl. Anim. Behav. Sci., 18(2), 119–131.
Abstract: The structure of the play of colts and fillies living on pasture was studied from birth (n = 15) for up to 24 weeks. Foal play was categorized as running and bucking alone, running and bucking in a group, interactive (contact or combat) play, play with an object, and play at an adult. The rate of play decreased with increasing age and ambient temperature. Fillies and colts played with equal frequency, but engaged in some different types of play at different rates. There was no difference between colts and fillies in the proportion of play bouts of running and bucking in a group or playing with an object. Fillies engaged in running and bucking alone more than colts. Colts engaged in interactive play and play at an adult more than fillies. While there was no significant difference between colts and fillies in the duration of either type of running and bucking play, the interactive play bouts of colts were significantly longer than those of fillies. Both mares and stallions were tolerant of foal play which involved use of their body as a play object, including mounting play. Both fillies and colts engaged in mounting play. Foals used various natural objects found in the pasture for repeated bouts of play with inanimate objects, a behaviour which may explain, from a developmental perspective, the occasional use of “tools” in adult equids. The sex differences in type of play were consistent with the social structure of unmanaged adults in which males must compete with each other in order to associate with females.
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