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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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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. 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. 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, R., & Silk, J. B. (1983). A method for assigning cardinal dominance ranks. Anim. Behav., 31(1), 45–58.
Abstract: Dominance hierarchies are widely described in nature. Commonly, an individual's ordinal rank is used as a measure of its position in the hierarchy, and, therefore its priority of access to resources. This use of ordinal ranks has several related drawbacks: (1) it is difficult to assess the magnitude or the significance of the difference in degree of dominance between two individuals; (2) it is difficult to evaluate the significance of differences between dominance matrices based on different behaviours or on the same behaviour at different times, and (3) it is difficult to use parametric statistical techniques to relate dominance rank to other quantities of interest. In this paper we describe a method for assigning cardinal dominance indices that does not suffer from these drawbacks. This technique is based on the Bradley-Terry model from the method of paired comparisons. We show how this model can be reinterpreted in terms of dominance interactions. and we describe a simple iterative technique for computing cardinal ranks. We then describe how to evaluate (1) whether the rank differences between individuals are significant, and (2) whether differences in the cardinal hierarchies based on different behaviours or the same behaviour at different times are significant. We then show how to generalize the method to deal with behaviours that sometimes have ambiguous outcomes, or behaviours for which the rank difference between a pair of individuals affects the rate of interaction between them.
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Breuer, K., Hemsworth, P. H., & Coleman, G. J. (2003). The effect of positive or negative handling on the behavioural and physiological responses of nonlactating heifers. Appl. Anim. Behav. Sci., 84(1), 3–22.
Abstract: This experiment investigated the effects of positive and negative tactile handling on the stress physiology and behaviour of dairy heifers. Forty-eight 5-14-month-old nonlactating Holstein-Friesian heifers were allocated to one of two handling treatments, either positive or negative tactile handling, over four time replicates. Handling was imposed twice daily, 2-5 min per session and involved moving animals individually along a 64 m outdoor route. The negatively handled heifers took longer to approach within 1 and 2 m of a stimulus person in a standard test, than their positively handled counterparts (P<0.001) and had a greater flight distance to an approaching stimulus (P<0.001). The time taken by the heifers to approach within 1 and 2 m of a familiar person was similar to that taken to approach within 1 and 2 m of an unfamiliar person in the standard test (P<0.05). There was a tendency for heifers to have a greater flight distance from the approaching unfamiliar person than from the approaching familiar person (P=0.06). The negatively handled heifers had greater (P<0.05) increases in total cortisol concentrations 5, 10 and 15 min after exposure to a human and had higher (P<0.05) free cortisol concentrations in the afternoon than the positively handled heifers. It is concluded that the nature of the human contact affects the subsequent behavioural response of heifers to humans. This behavioural response may extend to other humans through the process of stimulus generalisation, although there was some evidence of moderate discrimination. Negative handling results in an acute stress response in the presence of humans and also leads to a chronic stress response. Further research into the effect of these stress responses on milk production and welfare in fearful cows in a commercial situation is suggested.
Keywords: Dairy heifer; Fear; Handling; Stress response; Milk production; Stimulus generalisation
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Breuer, K., Sutcliffe, M. E. M., Mercer, J. T., Rance, K. A., Beattie, V. E., Sneddon, I. A., et al. (2003). The effect of breed on the development of adverse social behaviours in pigs. Appl. Anim. Behav. Sci., 84(1), 59–74.
Abstract: Tail-biting and other harmful social behaviours are a common problem on pig farms. The aims of the current experiment were (1) to investigate the genetic component of harmful social behaviours such as tail-biting by assessing breed differences, and (2) to further investigate the reliability and predictability of a test, [`]the tail-chew test', previously identified as potentially being capable of predicting a pig's predisposition for tail- and ear-biting. The behaviour of three pig breeds (Large White (LW), Landrace (LR), Duroc (DR)), with 100 pigs per breed, was observed in a [`]tail-chew test', and by observing the performance of harmful social behaviour directed to pen mates in flat deck pens after weaning. The tail-chew test, carried out on two consecutive days pre-weaning, involved observing the behaviour of individual pigs towards two suspended ropes. Pigs were weaned at 28 days and the occurrence of harmful social behaviour was recorded 4 weeks later over 2 consecutive days (1 h per day) using a group [`]period occurrence' scanning method. Breed had a significant effect on rope-directed behaviour in the tail-chew test and on harmful social behaviour. DR pigs interacted with the ropes in the tail-chew test more often (median 23.0 vs 19.0 and 17.5 times in 20 min, P<0.001) and for longer (31.0 vs 20.0 and 23.2 s, P<0.001) than LR and LW pigs, respectively. Although not significantly different from LW, DR pigs tended to direct more total harmful social behaviour towards pen mates than the other breeds. In particular, DR were observed in more total pig-directed biting of pen mates (median 9.0 vs 6.0 and 7.0, P<0.01) than LR and LW, and tended to nose pen mates more often than the other breeds (13.0 vs 11.0 and 10.0, P=0.06). LR pigs bit the ears of pen mates less often than LW and DR (4.0 vs 5.0 and 6.0, P<0.001). Belly-nosing activity was low, with a median of 0 for all breeds, but LR belly-nosed pen mates more often than Durocs (interquartile ranges 0-2.0 vs 0-1.0 and 0-1.0, P<0.01). The behaviour observed in the tail-chew test on day 1 correlated significantly with that observed on day 2 of the test (e.g. frequency of rope-directed behaviour rs=0.380,P<0.01). There were significant but weak correlations between rope-directed behaviour and the performance of some harmful social behaviours. The significant breed differences indicate some genetic contribution to expression of harmful social behaviours. However, the tail-chew test was found to be of limited ability to predict tail- and ear-biting under commercial conditions.
Keywords: Pig; Breed; Harmful social behaviour; Ear-biting; Tail-biting; Tail-chew test
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Briard, L., Deneubourg, J. - L., & Petit, O. (2017). How stallions influence the dynamic of collective movements in two groups of domestic horses, from departure to arrival. Behav. Process., 142, 56–63.
Abstract: Abstract The role of leader in polygynous species has been solely attributed to the male for some time, but recent studies shown decision making to be distributed within the group. However, the specific reproductive strategy and behavioural repertoire of males in polygynous species such as horses may mean that these individuals still have the potential to play a specific role during decision-making. To investigate this subject, we thoroughly studied the behaviour of two domestic stallions during collective movements of their group. We found that they initiated rarely and sometimes failed to recruit the entire group. When departing as followers, they did not accelerate the joining process. Both stallions preferentially occupied the rear position and exhibited numerous monitoring behaviours. Herding behaviours were performed by only one stallion and mostly occurred outside movement context. Finally, we removed this herding stallion from its group to evaluate how the group dynamic changed. As a result, half of the collective movements were five times slower and mares were more dispersed in comparison when the stallion was in the group. Overall, our results suggest that, the two stallions maintained their role of group monitors from departure to arrival. Their influence on the movement dynamic was indirect and did not play a specific role in the process of decision making.
Keywords: consensus; herding; polygyny; personal leadership; shared decision
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Brooks, C. J., & Harris, S. (2008). Directed movement and orientation across a large natural landscape by zebras, Equus burchelli antiquorum. Anim. Behav., 76(2), 277–285.
Abstract: We investigated how plains zebras moved across a large natural landscape by analysing the movement paths of nine zebra mares foraging out from spatially confined waterholes during the dry season in the Makgadikgadi Pans National Park, Botswana. Since it was essential to investigate directed movement over a range of spatial scales to determine the correct movement behaviour and strategy, we used Nams's scaling test for oriented movement. Zebras followed directed movement paths in the lower to medium spatial scales (10 m–3.7 km) and above their visual, and possibly olfactory, range. The spatial scale of directed movement suggests that zebras had a well-defined spatial awareness and cognitive ability. Seven zebras used directed movement paths, but the remaining two followed paths not significantly different to a correlated random walk (CRW). At large spatial scales (>3 km) no distinct movement pattern could be identified and paths could not be distinguished from a CRW. Foraging strategy affected the extent of directed movement: zebras with a confined dispersion of grazing patches around the central place directed their movements over a longer distance. Zebras may extend the distance at which they can direct their movement after improving their knowledge of the local environment.
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