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Hogan, E. S., Houpt, K. A., & Sweeney, K. (1988). The effect of enclosure size on social interactions and daily activity patterns of the captive Asiatic wild horse (Equus przewalskii). Appl. Anim. Behav. Sci., 21(1-2), 147–168.
Abstract: Two herds of Przewalski horses at the Minnesota Zoological Garden were observed during 1980 in each of 2 enclosures that differed in size. The larger enclosure was a 3.4-ha pasture; the smaller enclosure was a 17 x 30-m grass-less pen. One herd was composed of a stallion, 3 adult mares and 2 foals. The other consisted of a stallion and 2 mares. All occurrences of aggression, mutual grooming and snapping were recorded, and 5-min scan-samples of the activity state of each horse were taken. The time budgets, frequency of aggression and frequency of mutual grooming differed significantly with enclosure size for both herds. More time was spent pacing and milling in the smaller enclosure, and the frequency of aggressions and of mutual grooming was also higher. Only the foals exhibited snapping; frequency of snapping did not vary with enclosure size. More time was spent feeding in the larger enclosure. Provision of hay in the smaller enclosure eliminated the differences in time spent feeding. A second study was conducted during the spring of 1984 in an intermediate-sized enclosure, 0.4 ha, a sub-division of the pasture on which the horses were kept in 1980. One herd consisted of a stallion, 2 mares and 2 yearlings; the other consisted of a stallion, 3 mares and a foal. One of the stallions and all of the mares were those studied in 1980, but that stallion and one of the mares were in different herds than they had been in 1980. The frequency of aggression was similar to that observed in 1980.
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Tilson, R. L., Sweeny, K. A., Binczik, G. A., & Reindl, N. J. (1988). Buddies and bullies: Social structure of a bachelor group of Przewalski horses. Appl. Anim. Behav. Sci., 21(1-2), 169–185.
Abstract: The Minnesota Zoological Garden established a bachelor (i.e. all-male) group of 8 Przewalski horses (Equus przewalskii) in 1984. Possible methods for mixing males emerged through preliminary introductions and review of the relevant literature. These included: (1) grouping colts (<=3 years old) after removing them from their natal bands; (2) introducing adult stallions to colts in spacious, controlled environments to reduce the risk of combat-related injury or death associated with largely unpredictable adult temperament; (3) introducing more than 1 animal at a time to established groups. Based on the frequency and orientation of agonistic interactions observed among the bachelors, a linear dominance hierarchy was discerned among the 4 adult stallions; all 4 colts were determined to be of equally low or seemingly non-ranking status. Social rank among the adults was most influenced by individual temperament, and contention for rank was limited almost exclusively to the 3 lower-ranking stallions; the alpha male participated in 73% of all agonistic interactions observed, but was identified as the recipient in less than 3% of these interactions. Intra-band associations were detected in the bachelor group. The most prevalent of these was among the 4 colts, found together in over 70% of the observations. Other affinities commonly observed were between the colts (as a sub-group) and different stallions. The dominant stallion, the colts' most frequent companion, actively discouraged the 3 lower-ranking stallions from associating with the colts. Further, the colts were subjected to significantly fewer agonistic behaviors than the non-alpha stallions (per colt=1.05+/-0.24 h-1; per non-alpha stallion=2.01+/-0.04 h-1). These findings support the hypothesis developed by other workers for ungulates that young bachelors are sheltered from aggression by association with an alpha male. With one exception, adult associations were rarely noted. Although combat-related injuries (superficial wounds and temporary lameness) were common among the bachelors, few required medical attention. No permanent disabilities or deaths have occurred since the group's inception. The existence of social organization and intra-band associations in the bachelor group support the suggestion that such groups were once a natural part of this species' social milieu. Bachelor groups are therefore recommended as an ethologically sound, space- and cost-effective means of managing “surplus” (i.e. non-harem) male Przewalski horses.
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Brennan, J., & Anderson, J. (1988). Varying responses to feeding competition in a group of rhesus monkeys (Macaca mulatta). Primates, 29(3), 353–360.
Abstract: The behaviour of members of a group of rhesus monkeys was observed in experimentally created, competitive feeding situations. Socially dominant members of the group tended to start eating before lower-ranking subjects, and generally ate more. Dominants sometimes used aggression to control access to food, but overall, intermediate-ranking monkeys were involved in most agonistic episodes. Non-dominant subjects improved their feeding performance when food was presented in three piles rather than one pile, often by snatching food and consuming it away from the pile. These general patterns were less evident when realistic snake models were placed on some of the food piles. Feeding was disrupted by the presence of snakes, but notably, subordinates risked feeding in these conditions. Piles containing preferred foods and snakes were eaten from, but a low-preference food (carrot) under snakes went untouched by all subjects. The results suggest that group-members evaluate potential risks and benefits of competing for a restricted resource, and that dominance status, while an important factor, is only one element in the equation.
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Houpt, K. A., & Fraser, A. F. (1988). Przewalski horses. Appl. Anim. Behav. Sci., 21(1-2), 1–3.
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Kirkpatrick, J. F., Kasman, L. H., Lasley,, B. L., & Turner, J. W. J. (1988). Pregnancy Determination in Uncaptured Feral Horses. J Wildl Manag, 52(2), 305–308.
Abstract: The urinary excretion of estrone sulfate ($\text{E}{1}\text{S}$) by 25 free-roaming feral horses (Equus caballus) was measured by radioimmunoassay applied to extracts of urine-soaked soil. Twelve of 15 mares having $\text{E}{1}\text{S}$ concentrations >1.0 mg/mg creatinine (x = 2.64 +- 1.02 [SD]) produced foals. All 10 mares with $\text{E}{1}\text{S}$ concentrations <1.0 mg/mg creatinine (x = 0.44 +- 0.26) did not foal. Extracting urine from soil and measuring $\text{E}{1}\text{S}$ and creatinine can be used to determine pregnancy in free-roaming feral horses without the stress of capture or immobilization.
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Klimov, V. V. (1988). Spatial-ethological organization of the herd of Przewalski horses (Equus przewalskii) in Askania-Nova. Appl. Anim. Behav. Sci., 21(1-2), 99–115.
Abstract: The ethological structure of the herd of Przewalski horses includes hierarchic ranks of horses which determine their social roles in the herd. Besides the age ranks, the wild horses are characterized by the formation of harem groups, a “leading” group of females, a group of bachelor stallions, family groups, etc. The ethological structure determines the spatial one, which is the form of distribution of horses over the territory, and its assimilation and transformation into a system of informative spatial units. Under the influence of “internal” and “external” stimuli, the intragoup regulatory mechanims (social adaptations) manifest themselves, which allow the herd to function in the complicated situation of the reserve and allow humans to control the herd by using these mechanisms. There are grounds to believe that, given the balanced ethological structure of these groups, wild horses could be successfully acclimatized into natural biotopes.
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Kolter, L., & Zimmermann, W. (1988). Social behaviour of Przewalski horses (Equus p. przewalskii) in the Cologne Zoo and its consequences for management and housing. Appl. Anim. Behav. Sci., 21(1-2), 117–145.
Abstract: Between 1977 and 1986, two actual rank changes and two unsuccessful attempts occurred among the mares of the Cologne herd. The stallion was at first a low-ranking individual, but attained a dominant position during the last 3 years. At this time he started to split his group and thus to affect attachment relationships among his mares. During his absence of half a year, new bonds resulted and disappeared again some months after his return. Foals were tolerated by the sire for a long time. His behaviour to young, sub-adult mares varied with the individual. Protection of sub-adult mares by adult mares against the stallion's attacks may occur. Young mares protect and guard strange foals from their very first day. Management steps to cope with social and feeding problems consisted of enlargement and adding complexity to the enclosure, the establishment of more feeding sites, building a stable and temporary removal of the stallion.
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Boyd, L. E., Carbonaro, D. A., & Houpt, K. A. (1988). The 24-hour time budget of Przewalski horses. Appl. Anim. Behav. Sci., 21(1-2), 5–17.
Abstract: A herd of 8 Przewalski horses were observed on pasture in summer. Fifteen-minute focal animal samples were used to determine the time budget of the horses during the periods 00.00-04.00, 04.00-08.00, 08.00-12.00, 12.00-16.00, 16.00-20.00 and 20.00-24.00 h EDT. The behavioral states recorded were feeding (grazing and eating grain), nursing, drinking, standing, stand-resting, self-grooming, mutual grooming, locomoting, playing, and lying laterally and sternally. The average number of behavioral states occurring per hour, and the defecation, urination, aggression and vocalization rates were also determined. Overall, the horses spent 46.4 +/- 5.9% of their time feeding, 1.3 +/- 0.1% nursing, 0.5 +/- 0.1% drinking, 20.6 +/- 5.4% standing, 15.7 +/- 3.2% stand-resting, 1.7 +/- 0.2% self-grooming, 2.2 +/- 0.7% mutual grooming, 7.4 +/- 1.0% locomoting, 1.2 +/- 0.3% playing, 1.2 +/- 0.5% lying laterally and 4.1 +/- 3.0% lying sternally. The horses averaged 45.2 +/- 5.8 behavioral states per hour, and 0.2 +/- 0.0 defecations, 0.3 +/- 0.0 urinations, 1.5 +/- 0.3 aggressions and 0.7 +/- 0.1 vocalizations per hour. The horses spent the greatest amount of time foraging between 20.00 and 04.00 h, when the temperatures were lower. They spent 68.2 +/- 2.2% of their time between 20.00 and 24.00 h feeding, but only 31.2 +/- 2.1% of their time feeding between 08.00 and 12.00 h. Recumbent rest was most common between 00.00 and 04.00 h. As temperatures rose during the daylight hours, the horses spent more time drinking and standing, rather than grazing. Stand-resting was the most common form of rest during the day. The horses exhibited the greatest number of activities per hour from 08.00 to 20.00 h. While standing in close proximity to one another during these hours, the horses exhibited the highest number of aggressions per hour (1.9-2.4).
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Boyd, L. E. (1988). Ontogeny of behavior in Przewalski horses. Appl. Anim. Behav. Sci., 21(1-2), 41–69.
Abstract: Twelve colts and 12 fillies were observed during their first 2 years of life. Data on the foal's nearest neighbor, distance to dam and stallion, and time budget were compiled by age. The birth of one foal was witnessed. During their first month of life, Przewalski foals were dependent on the dam. She provided most of their nourishment and foals spent 54% of their time within 1 m of her. The biggest change in behavior of foals occurred between Months 1 and 2. The amount of time spent resting and nursing declined, while the amount of time spent foraging increased sharply. Foals began to leave their mothers and interact with peers by 3 weeks of age, and at 2 months they were interacting with older herd members. By 5 months of age, the amount of time spent in most behaviors was identical to that of adults, except that vocalization rates and involvement in aggression were lower than for adults. Juveniles spent less time stand-resting than adults throughout their first year, but more time in recumbent rest. Foals spent far less time with their sire than with their dam. However, an orphaned foal spent more time with his sire than did mothered foals, indicating that the sire assumed part of the role of the missing dam.
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Berger, J., & Cunningham, C. (1988). Size-Related Effects on Search Times in North American Grassland Female Ungulates. Ecology, 69(1), 177–183.
Abstract: Feeding and searching (= vigilance) rates arise as a result of many interrelated factors including trophic level, diet, reproductive condition, sex, habitat, body mass, and potential predation pressure. Because of unique ecological conditions in which the confounding influences of all but two of these variables could be minimized, we examined the hypothesis that body mass alone accounts for interspecific differences in search times, and tested it with females of four sympatric native North American ungulates (Bison bison, Antilocapra americana, Ovis canadensis, and Odocoileus hemionus). When the effects of group size were controlled, smaller bodied species were more vigilant (per unit body mass) than larger ones. However, search times (ST) also scaled to body mass, and between 81 and 97% of the ST variance was explained by either exponential or power functions. To remove the potential bias that predators exert different influences on species of varying size, search times of bison in areas with and without their major predator, wolves (Canis lupus), were contrasted; search times did not differ between sites. Our results highlight the importance of designing field research that controls for confounding variables prior to attempting to scale behavioral processes to ecological events. See full-text article at JSTOR
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