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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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Barry, K. J., & Crowell-Davis, S. L. (1999). Gender differences in the social behavior of the neutered indoor-only domestic cat. Appl. Anim. Behav. Sci., 64(3), 193–211.
Abstract: The domestic cat exhibits a wide variety of social behavior. The aim of this experiment was to investigate factors which influence the affiliative and aggressive behavior of the indoor-only neutered domestic cat. Some 60 households comprised of either two males, two females or a male and female cat were observed. The cats were between 6 months and 8 years old, and were always restricted to the indoors. Each pair of housemates was observed for 10 h. There were no significant differences in affiliative or aggressive behavior based on cat gender. However, females were never observed to allorub other females. The male/male households did spend more time in close proximity. The amount of time the cats had lived together was negatively correlated with the amount of aggression observed during the study. Factors such as size of the house and weight difference between the cats did not correlate with the aggression rate. Large standard deviations and the correlations of social behavior between housemates indicated the importance of individual differences in behavior.
Keywords: Sex differences; Spatial distribution; Cat; Social; Aggression; Affiliation; Felis catus
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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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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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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
Keywords: aggression; dominance; horse; Equus caballus
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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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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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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.
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