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Boyd, L. (1986). Behavior problems of equids in zoos. Vet Clin North Am Equine Pract, 2(3), 653–664.
Abstract: Behavior problems in zoo equids commonly result from a failure to provide for needs basic to equine nature. Equids are gregarious, and failure to provide companions may result in pacing. Wild equids spend 60 to 70 per cent of their time grazing, and failure to provide ad libitum roughage contributes to the problems of pacing, cribbing, wood chewing, and coprophagia. Mimicking the normal processes of juvenile dispersal, bachelor-herd formation, and mate acquisition reduces the likelihood of agonistic and reproductive behavior problems. Infanticide can be avoided by introducing new stallions to herds containing only nonpregnant mares and older foals.
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Boyd, L. E. (1991). The behaviour of Przewalski's horses and its importance to their management. Appl. Anim. Behav. Sci., 29(1-4), 301–318.
Abstract: Przewalski's horses (Equus przewalskii) are believed to be extinct in the wild; the current known population of 797 animals exists wholly in zoos. The Species Survival Commission of the International Union for Conservation of Nature and Natural Resources is proposing to reintroduce this endangered species into its former Mongogian habitat within the next decade. Knowledge of the behavior of harem-forming equids in general and of Przewalski's horses in particular, is of great importance to the captive propagation and eventual reintroduction of this species. Horses are rarely solitary by nature. Solitary captive animals are prone to pacing. Juvenile male feral horses (Equus caballus) form bachelor herds upon dispersal from their natal band. Zoos can set up bachelor herds as a way of managing surplus males. The older, more dominant feral horse bachelors are the first to acquire mares. Bachelors do not generally obtain females until they are 4 or 5 years of age. The first females acquired are usually 1- and 2-year-old fillies dispersing from their natal band. Because of the age differential, the stallions are generally dominant to their mares. Behavioral impotence may result if captive stallions are given a harem at too young an age, especially if the harem contains older, more dominant, females. Typical harem sizes in the wild are 3-5 mares. Captive stallions with too large a harem may become either apathetic or aggressive toward their mares. Wild horses spend 60-70% of their time foraging. Captive animals may quickly consume their limited amounts of food and develop vices out of boredom. Provision of hay ad libitum reduces the amount of pacing seen in captive animals, and virtually eliminates coprophagy.
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Boyd, L., & Houpt, K. A. (1994). Przewalski's Horse. The History and Biology of an Endangered Species. SUNY Press.
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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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Boyd, L. (1998). The 24-h time budget of a takh harem stallion (Equus ferus przewalskii) pre- and post-reintroduction. Appl. Anim. Behav. Sci., 60(4), 291–299.
Abstract: Focal animal sampling was used to determine the 24-h time budget of a takh harem stallion (Equus ferus przewalskii) during the 2 weeks prior to, and the two weeks following, reintroduction into the Hustain Nuruu Steppe Reserve, Mongolia. Both before and after release, the stallion spent approximately 47% of his time grazing, 6% standing, and 5% in recumbent rest. The biggest changes to the time budget after release were a 4-fold increase in the amount of time spent moving, and a 50% decrease in the amount of time spent resting in a standing position. During the middle of the day when the temperatures were hottest, the stallion exhibited less grazing and more standing resting behaviour than in the morning or evening hours. Recumbent rest invariably occurred in the hours before dawn.
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Boyd, L., & Bandi, N. (2002). Reintroduction of takhi, Equus ferus przewalskii, to Hustai National Park, Mongolia: time budget and synchrony of activity pre- and post-release. Appl. Anim. Behav. Sci., 78(2-4), 87–102.
Abstract: A harem of takhi (Equus ferus przewalskii) was observed during introduction to the Hustain Nuruu Steppe Reserve of Mongolia. The goals were to examine whether the harem exhibited significant behavioural synchrony, whether release had an effect on time budget, and what the implications might be regarding acclimatisation to the wild. Behaviours were scan sampled every 10 min between the hours of 06:00 and 22:00, twice before release, twice immediately after release, and twice 2 years after reintroduction. Time budgets were constructed from these data. Considerable behavioural synchrony was evidenced both before and after release. Crepuscular grazing and midday resting were typical, regardless of the date relative to release. Upon release, the amount of time spent moving doubled for all age classes. It is suggested that this increase resulted from exploration. The amount of time spent grazing and standing remained unchanged; the increased amount of time spent moving came at the expense of resting. Two years later, the horses still spent more time moving than when captive. Somewhat less time was spent grazing, although the difference was not significant. More time was spent resting in 1996 than immediately after release. These time budgets provide evidence of successful acclimatisation to the wild. Trekking between favoured sites could account for the persistent increase in time spent moving, with concomitantly less time needed to meet nutritional needs by grazing and more time available for resting. Housing captive takhi in large enclosures is evidently insufficient to permit the amount of movement exhibited by this wild harem. The time budget of the 1- and 2-year olds was more similar to that of adults than foals, indicating approaching adulthood. That 1- and 2-year olds were nursed, without loss of body condition by the dam, provided additional evidence that the takhi achieved excellent nutritional status in the wild.
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Boyd, L., & Keiper, R. (2005). Behavioural ecology of feral horses. In D. S. Mills, & McDonnell S. M. (Eds.), The domestic horse: the origins, development, and management of its behaviour. Cambridge: Cambridge University Press.
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