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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. (1986). Spatial relations between mares and foals of the Welsh pony (Equus caballus). Anim Beh, 34(4), 1007–1015.
Abstract: Welsh pony mares and foals (Equus caballus) were usually found to be within 1 or 5 m of each other during the first week of the foal's life and gradually spent more time at greater distances as the foals became older. There was an overall levelling of the trend during the 9th-15th weeks of life of the foal, followed by a second period of change during weeks 16-24. Through weeks 21-24, mares and foals spent at least half of their time within 5 m of each other. Proximity was primarily due to foal activity except during foal recumbency. During the first 8 weeks of the foal's life, a mare remained close by when it was recumbent, either by grazing in a circle around it or by standing upright beside it. Mares and foals were most likely to be close together when they were resting upright with the other ponies in the herd and most likely to be far apart when the foal was playing. Similarities in patterns of spatial relationship between the foals of a given mare were demonstrated. There was no difference between colts and filies in the development of independence.
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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., & Carnevale, J. (1985). Feeding and drinking behavior of mares and foals with free access to pasture and water. J. Anim Sci., 60(4), 883–889.
Abstract: The feeding and drinking behavior of 11 mares and 15 foals living on pasture with free access to water was recorded during 2,340 15-min focal samples taken over 2 yr. Lactating mares on pasture spent about 70% of the day feeding. Foals began feeding on their first day of life. As they grew older, they spent progressively more time feeding, but still spent only 47 +/- 6% of the time feeding by 21 wk of age. Foals fed primarily during the early morning and evening. While grass formed the major proportion of the diet of both foals and mares, they also ate clay, humus, feces, bark, leaves and twigs. Almost all feeding by foals was done while their mothers were feeding. Movement to water sources was frequently, but not invariably, carried out by an entire herd. Frequency (P = .005) but not duration (P greater than .05) of drinking bouts by mares increased as the temperature increased. Frequency was greatest at 30 to 35 C, at which temperature mares drank once every 1.8 h. Frequency of drinking varied with the time of day (P less than .01), being rarest during the early morning (0500 to 0900 h eastern daylight time) and most frequent during the afternoon (1300 to 1700 h). Drinking by foals was very rare. The youngest age at which a foal was observed to drink was 3 wk, and 8 of 15 foals were never observed to drink before weaning.
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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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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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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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