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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.
Keywords: Horse behavior; Flehmen; Foal development
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Weeks, J. W., Crowell-Davis, S. L., Caudle, A. B., & Heusner, G. L. (2000). Aggression and social spacing in light horse (Equus caballus) mares and foals. Appl. Anim. Behav. Sci., 68(4), 319–337.
Abstract: Aggression and social spacing were studied in 14 light horse mares and their foals living at pasture. Focal samples were collected on each mare-foal dyad for 6 to 10.5 h from 2 months of foal age until weaning at approximately 4 months of age. Observations on foals continued until approximately 6 months of age for 7.5 to 10.5 h per foal. Every 2 min the identities of all individuals within 5 m were recorded. All occurrences of agonistic behavior, and the participants, were recorded during the focal samples. In addition, during feeding of supplemental grain, all occurrences of agonistic behavior by all subjects were recorded. Significant correlations were found between mare rank and the rank of foals both prior to and after weaning. Before weaning, the rank of the foal was significantly correlated with birth order. No significant correlation between birth order and foal rank was found for the post-weaning hierarchy. An animal's gender had no significant effect on foal rank or the choice of preferred associate. Both prior to and after weaning, foals associated preferentially with the foal of their dam's most preferred associate. In addition, significant positive correlations were found between rank of mares and foals and the rate at which they directed aggression to other herd members. (C) 2000 Elsevier Science B.V.
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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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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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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.
Keywords: Horse; Sexual behavior; Sexual preference; Vocalization
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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. |
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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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., 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.
Keywords: Animals; *Drinking Behavior; *Feeding Behavior; Female; Horses/*physiology; Male; Poaceae; Seasons; Temperature; Time Factors
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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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