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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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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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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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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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Weeks, J. W., Crowell-Davis, S. L., & Heusner, G. (2002). Preliminary study of the development of the Flehmen response in Equus caballus. Appl. Anim. Behav. Sci., 78(2-4), 329–335.
Abstract: The flehmen response is commonly seen in most ungulates as well as in several other species (e.g. felids). The behavior is most often thought to be part of the sexual behavioral repertoire of males. One reigning hypothesis suggests that this behavior allows the male to determine the estrous state of a female through the chemosensory functions of the vomeronasal organ. However, females and young of both sexes also exhibit this behavior. Horse foals most frequently show the flehmen response during their first month of life with colts showing the behavior more often than fillies. This study tested the flehmen response on male and female foals throughout their pre-pubertal period. Foals were separately presented estrous and non-estrous urine weekly during the first month of life and then monthly until they were approximately 7 months of age. No significant differences were found between male and female foals for the following variables: latency to flehmen, duration of flehmen, frequency of flehmen and sniffs.
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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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Barber, J. A., & Crowell-Davis, S. L. (1994). Maternal behavior of Belgian (Equus caballus) mares. Appl. Anim. Behav. Sci., 41(3-4), 161–189.
Abstract: The relationship between ten Belgian mares and their offspring was studied from the first day of foal life to 17 weeks of age. Mares and foals spent more time at greater distances from each other as foals matured. Mares exhibited the recumbency response, being in closer proximity to their foals when foals were recumbent than when they were upright. Foals initiated the majority of nursing bouts. Frequency and duration of nursing bouts and percentage of time resting recumbently declined as foals matured. Foals also terminated the their foals, and they were most likely to do so in the first month of foal life. Maternal initiation of nursing. There was usually no discernible foal response to maternal aggression. Little difference between maternal behavior directed towards colts and fillies was found for all aspects of the study. Maternal behavior in the Belgian draft horse was similar to that reported for other equid breeds.
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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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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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Crowell-Davis, S. L. (1994). Daytime rest behavior of the Welsh pony (Equus caballus) mare and foal. Appl. Anim. Behav. Sci., 40(3-4), 197–210.
Abstract: Upright and recumbent rest of 15 Welsh pony foals and their mothers was studied over a 2 year period. During their first week of life, the foals spent 32% of the time in recumbent rest. Subsequently, the percentage of time spent in recumbent rest decreased, but was still greater than for the foal's mother by Week 21, when the foals spent 6.5% of their time in recumbent rest. Adults spent little time in recumbent rest. Foals rested upright only 3.5% of the time during their first week of life. Mares rested upright more than foals did to Week 13, at which time peak values for time spent in upright rest occurred for both mares (32.5%) and foals (23%). Subsequently, mares and foals spent equal, but decreasing, amounts of time resting upright. The total time spent resting by the foals decreased gradually, and was characterized by a transition from recumbent rest to upright rest. Foals were more likely to be resting, either recumbent or upright, if their mother was resting upright. During the late spring, summer, and early autumn, mares and foals were most likely to be resting upright between 09:00 and 17:00 h.
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