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R. A. Hopkins. CALIFORNIA WILDLIFE HABITAT RELATIONSHIPS SYSTEM (Vol. M174).
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Austin, N. P., & Rogers, L. J. (2012). Limb preferences and lateralization of aggression, reactivity and vigilance in feral horses, Equus caballus. Anim. Behav., 83(1), 239–247.
Abstract: Observational field studies were conducted on two remote populations of feral horses in Australia to determine whether lateralization is a characteristic of Equus caballus as a species or results from handling by humans. Group 1 had been feral for two to five generations and Group 2 for 10–20 generations. In both groups, left-side biases were present during agonistic interactions and in reactivity and vigilance. Therefore, as in other vertebrates, the right hemisphere appears to be specialized to control agonistic behaviour and responses to potential threats. The leftwards bias was stronger in measures of behaviour involving more aggression and reactivity. Preferences to place one forelimb in front of the other during grazing were also determined. No population bias of forelimb preference was found, suggesting that such limb preferences present in domestic horses may be entrained. Since stronger individual limb preferences were found in immature than in adult feral horses, limb preference may be modified by maturation or experience in the natural habitat. Stronger limb preference was associated significantly with elevated attention to the environment but only in younger feral horses. No sex differences in lateralization were found. The findings are evidence that horses show visual lateralization, as in other vertebrates, not dependent on handling by humans. Limb preference during grazing, by contrast, does appear to depend on experience.
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Cameron, E. Z., Linklater, W. L., Stafford, K. J., & Minot, E. O. (2008). Maternal investment results in better foal condition through increased play behaviour in horses. Anim. Behav., 76(5), 1511–1518.
Abstract: Play behaviour is widespread in mammals, but benefits to play have been difficult to demonstrate. Physical training is one of the many proposed hypotheses, suggesting that males and females should play differently, that increased maternal investment should lead to increases in play, and that increases in play should result in physical advantages. In a population of feral horses, Equus caballus, males and females did not differ in their play behaviour except that males initiated more of their play bouts than females. Maternal condition influenced play behaviour only in males, with sons of mothers in good condition playing more. However, when we controlled for maternal effects by comparing a son and a daughter of the same mother, daughters played more when their mother was in poor condition and sons played more when their mother was in good condition. Mothers of foals that played more lost more condition. Therefore, the difference in play behaviour could not be explained by offspring sex or maternal condition alone, but play behaviour mirrored variation in maternal investment. In addition, those individuals that played more survived better and had better body condition as yearlings despite weaning earlier. Since increased activity has been linked to enhanced musculoskeletal development in domestic horses, we suggest that play provides a link between increased maternal investment, increased body condition and future reproductive success in feral horses, and probably in other species.
Keywords: Equus caballus; feral horse; maternal investment; play
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Wolter, R., Stefanski, V., & Krueger, K. (2018). Parameters for the Analysis of Social Bonds in Horses. Animals, 8(11), 191.
Abstract: Social bond analysis is of major importance for the evaluation of social relationships in group housed horses. However, in equine behaviour literature, studies on social bond analysis are inconsistent. Mutual grooming (horses standing side by side and gently nipping, nuzzling, or rubbing each other), affiliative approaches (horses approaching each other and staying within one body length), and measurements of spatial proximity (horses standing with body contact or within two horse-lengths) are commonly used. In the present study, we assessed which of the three parameters is most suitable for social bond analysis in horses, and whether social bonds are affected by individual and group factors. We observed social behaviour and spatial proximity in 145 feral horses, five groups of Przewalski�s horses (N = 36), and six groups of feral horses (N = 109) for 15 h per group, on three days within one week. We found grooming, friendly approaches, and spatial proximity to be robust parameters, as their correlation was affected only by the animals� sex (GLMM: N = 145, SE = 0.001, t = �2.7, p = 0.008) and the group size (GLMM: N = 145, SE < 0.001, t = 4.255, p < 0.001), but not by the horse breed, the aggression ratio, the social rank, the group, the group composition, and the individuals themselves. Our results show a trend for a correspondence between all three parameters (GLMM: N = 145, SE = 0.004, t = 1.95, p = 0.053), a strong correspondence between mutual grooming and friendly approaches (GLMM: N = 145, SE = 0.021, t = 3.922, p < 0.001), and a weak correspondence between mutual grooming and spatial proximity (GLMM: N = 145, SE = 0.04, t = 1.15, p = 0.25). We therefore suggest either using a combination of the proactive behaviour counts mutual grooming and friendly approaches, or using measurements of close spatial proximity, for the analysis of social bonds in horses within a limited time frame.
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Khalil, A. M., & Kaseda, Y. (1998). Early experience affects developmental behaviour and timing of harem formation in Misaki horses. Appl. Anim. Behav. Sci., 59(4), 253–263.
Abstract: A study was made of the behavior of young male Misaki feral horses in the developmental stage, by observing nine of them once a week from January 1988 to December 1996. The relationship between behavior before separation and in the developmental stage was also investigated. This stage begins just after young males separate from their natal band or mothers, and it continues until they start to form harems. The duration of the developmental stage in the study ranged from 0.6 to 3.9 years, depending on the age of the young males at the time of separation. Young males associated with three types of social groups at the beginning of the developmental stage, according to their social groups before separation. These were bachelor groups, harem groups and wandering female groups. During this period, males joined the three groups, mixed sex groups and sometimes were solitary. It was considered that these associations provided a good opportunity for males to acquire different behavioral patterns and experiences before they entered the next stage. Depending on the groups with which they associated, young males that spent more time with bachelor groups had the longest average developmental stage. They associated with harem groups more often during the breeding season and more frequently with other groups or were solitary during the non-breeding season. This may be a transition period because by the end of this stage all males had spent time in solitude before forming their own harem bands.
Keywords: Feral horse; Young male; Social behavior; Developmental stage
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Khalil, A. M., & Kaseda, Y. (1997). Behavioral patterns and proximate reason of young male separation in Misaki feral horses. Appl. Anim. Behav. Sci., 54(4), 281–289.
Abstract: The present investigation was undertaken to study the proximate reasons why and the behavioral patterns of young male Misaki feral horses when they left their natal band or mothers. We observed a total of ten young males twice a month from January 1988 to December 1995. Almost all young males left their natal band or mothers at between 1 and 4 years of age. We found that, during the separation process, all the young males from first parity dams returned several times after the initial separation, indicating a strong attachment between primiparous mares and their male offspring. The other five separated only once without rejoining. Our observations showed five variable behavior patterns of young males at separation time, depending on the consort relation between their mothers and harem stallion and the reason for separation at that time. Eight young males separated in the non-breeding season at average 2.1 years and the other two separated in the breeding season at average 3 years and the average difference was not significant. These results revealed that 80% of the young males separated voluntarily when the natural resources become poor whereas 20% separated when their siblings were born.
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Kaseda, Y., & Khalil, A. M. (1996). Harem size and reproductive success of stallions in Misaki feral horses. Appl. Anim. Behav. Sci., 47(3-4), 163–173.
Abstract: Over a 16-year period (1979-1994), long-term investigations were carried out on 14 Misaki feral stallions to analyze changes in harem size and the reproductive success. Harem size changed with the age of the stallions. Most stallions formed harem groups with four to five mares at the age of 4-6 and then the number of mares increased rapidly to the maximum at the age of 6-9 years. Thereafter, harem size decreased gradually to a minimum with advancing age. The harem size of 60 stable harem groups ranged from 1 to 9, and the average varied from a minimum mean of 1.8 in 1988 to a maximum mean of 5.3 in 1982. Mean harem size increased as adult sex ratio increased and a significant and positive correlation was found between them. One hundred and ninety-eight sire-foal pairs were determined by a paternity test with blood types and consort relations between stallions and mares during the study period. Out of 99 foals which were born in the stable harem groups, the true sires of 84 foals (85%) were the harem stallions in which the foals were born but the remaining 15 foals (15%) were sired by other harem stallions. Two out of three stallions which were studied throughout their lifetime produced 24 and 25 foals in 10 and 11 years of their reproductive lifespan, respectively. Another one produced only five foals in 6 years. The number of foals sired by the harem stallions was less than two over harem size 7 and some of the foals born in the harem were sired by other harem stallions. These results suggest that if a particular stallion monopolizes too many mares, he could not sire so many offspring because he could not always prevent his rival stallions from mating with his mares in wild or feral circumstances.
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Moehlman, P. D. (1998). Behavioral patterns and communication in feral asses (Equus africanus). Appl. Anim. Behav. Sci., 60(2-3), 125–169.
Abstract: The behavior of feral populations of the African wild ass (Equus africanus) were studied in the Northern Panamint Range of Death Valley National Monument for 20 months from 1970 to 1973 [Moehlman, P.D., 1974. Behavior and ecology of feral asses (Equus asinus). PhD dissertation, University of Wisconsin, Madison, 251 pp.; Moehlman, P.D., 1979. Behavior and ecology of feral asses (Equus asinus). Natl. Geogr. Soc. Res. Reports, 1970: 405-411]. Maintenance behavior is described and behavior sequences that were used in social interactions are quantified by sex and age class. Agonistic, sexual, and greeting behavior patterns are described and analyzed in conjunction with the responses they elicited. Mutual grooming mainly occurred between adult males, and between females and their offspring. Five types of vocalizations were distinguished: brays, grunts, growls, snorts, and whuffles. A second population was studied for 1 month on Ossabaw Island, GA (Moehlman, 1979). This population had more permanent social groups and had a higher rate of mutual grooming and foal social play.
Keywords: Equids; Feral asses; Behavior patterns; Facial expressions; Postures; Locomotion
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Moehlman, P. D. (1998). Feral asses (Equus africanus): intraspecific variation in social organization in arid and mesic habitats. Appl. Anim. Behav. Sci., 60(2-3), 171–195.
Abstract: Feral asses have been studied in the arid habitats of the southwestern United States [Moehlman, P.D., 1974. Behavior and ecology of feral asses (Equus asinus). PhD dissertation, University of Wisconsin, Madison, 251 pp.; Moehlman, P.D., 1979. Behavior and ecology of feral asses (Equus asinus). Nat. Geogr. Soc. Res. Reports 1970, 405-411.; Woodward, S.L., 1979. The social system of feral asses (Equus asinus). Z. Tierpsychol. 49, 304-316] and in the mesic habitat of Ossabaw Island, Georgia [Moehlman, P.D., 1979, ibid; McCort, W.D., 1980. The feral asses (Equus asinus) of Ossabaw Island, Georgia. PhD dissertation, Pennsylvania State University, University Park, 219 pp]. The feral ass populations in these two locales exhibited intraspecific variation in polygynous mating systems and social organization which were consistent with the ecological classification of mating systems of Emlen and Oring (1977) [Emlen, S.T., Oring, S.W., 1977. Ecology, sexual selection, and the evolution of mating systems. Science 197 (4300), 215-223]. Feral asses in the arid environment have a `resource defense' polygynous mating system, and those in the mesic habitat exhibit `female (harem) defense' polygyny. The intraspecific variation observed in feral asses encompasses the interspecific variation observed in the family Equidae.
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Moehlman, P. D., Fowler, L. E., & Roe, J. H. (1998). Feral asses (Equus africanus) of Volcano Alcedo, Galapagos: behavioral ecology, spatial distribution, and social organization. Appl. Anim. Behav. Sci., 60(2-3), 197–210.
Abstract: Feral asses were studied on Volcano Alcedo, Galapagos Islands, Ecuador, during the wet season of 1980. On the volcano rim during March/April, two stable groups were observed to have a `female (harem) defense' polygynous mating system [Emlen, S.T., Oring, S.W., 1977. Ecology, sexual selection, and the evolution of mating systems. Science 197 (4300), pp. 215-223] and social behavior patterns and feeding ecology similar to feral asses living in a habitat where forage and climate are similar, e.g., Ossabaw Island, Georgia [Moehlman, P.D., 1979. Behavior and ecology of feral asses (Equus asinus). Nat. Geogr. Soc. Res. Rep., 1970, pp. 405-411; Moehlman, P.D., 1997. Feral asses (Equus africanus): intraspecific variation in social organization in arid and mesic habitats. J. Appl. Anim. Behav. Sci., this issue; McCort, W.D., 1980. The feral asses (Equus asinus) of Ossabaw Island, Georgia., PhD Dissertation, Pennsylvania State University, University Park, 219 pp.].
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