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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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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., & Houpt, K. A. (1985). The ontogeny of flehmen in horses. Anim. Behav., 33(3), 739–745.
Abstract: Flehmen behaviour in Welsh pony (Equus caballus) mares and foals living on pasture was observed during 807 h of focal sampling. A series of flehmens performed at one site was defined as a flehmen incident. Colts exhibited flehmen incidents and performed flehmen more frequently during an incident than did fillies or mares. Filies exhibited flehmen incidents more frequently than did mares, but did not flehmen more frequently during an incident. Colts exhibited a peak frequency of performing flehmen and of flehmen incidents during weeks 1-4 with a subsequent linear decrease in frequency up to weeks 17-20. Usually, flehmen occurred without the subject having had direct contact of the nostrils, lips, or tongue with a possible stimulant. Twenty-six per cent of the flehmen incidents occurred during or after urination by another pony. Seven per cent of the incidents occurred during or after urination by the pony showing flehmen.
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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., & Caudle, A. B. (1989). Coprophagy by foals: recognition of maternal feces. Appl. Anim. Behav. Sci., 24(3), 267–272.
Abstract: Six foals were each observed for 4 h per week during the first 6 weeks of life in an experimental situation in which they had access to feces taken from their mother and from another mare which was not pregnant or lactating. The foals sniffed at the feces equally. Two foals engaged in a total of seven bouts of coprophagy. All bouts of coprophagy involved maternal feces (χ2; P<0.01).
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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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Crowell-Davis, S. L. (1987). Self-grooming by mares and foals of the Welsh pony (Equus caballus). Appl. Anim. Behav. Sci., 17(3-4), 197–208.
Abstract: Self-grooming behaviour of 15 mare-foal pairs was studied weekly for the first 24 weeks of the foal's life. Mares self-groomed at mean rates of 1.2-2.2 times h-1 depending on the 4-week period. Foals self-groomed more often, with a peak rate of 12.3 times h-1 occurring during Week 5-8 and a subsequent decline to 6.0 times h-1 by Weeks 21-24. Self-grooming bouts of mares and foals also differed in the proportion devoted to each type of self-grooming. During a greater proportion of their total self-grooming bouts, foals scratched their head and neck with a hind limb or bit and scratched the trunk or hind limbs with the teeth. The mares spent a greater proportion of their self-grooming bouts rubbing their head or neck on an inanimate object, another pony, or their fore limb, or rolling on the ground. Particular types of individual self-grooming bouts were engaged in for different lengths of time. Foals had longer bouts of scratching the head and neck with a hind limb, scratching or biting the fore limb or hind limb with the teeth, and rubbing the head and neck on an inanimate object, while mares had longer bouts of rolling. Some of the differences in preference for particular types of self-grooming may be the result of differences in relative size of various body parts and, consequently, relative ease of a given type of self-grooming.
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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. (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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