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Arnold, W., Ruf, T., & Kuntz, R. (2006). Seasonal adjustment of energy budget in a large wild mammal, the Przewalski horse (Equus ferus przewalskii) II. Energy expenditure. J Exp Biol, 209(Pt 22), 4566–4573.
Abstract: Many large mammals show pronounced seasonal fluctuations of metabolic rate (MR). It has been argued, based on studies in ruminants, that this variation merely results from different levels of locomotor activity (LA), and heat increment of feeding (HI). However, a recent study in red deer (Cervus elaphus) identified a previously unknown mechanism in ungulates--nocturnal hypometabolism--that contributed significantly to reduced energy expenditure, mainly during late winter. The relative contribution of these different mechanisms to seasonal adjustments of MR is still unknown, however. Therefore, in the study presented here we quantified for the first time the independent contribution of thermoregulation, LA and HI to heart rate (f(H)) as a measure of MR in a free-roaming large ungulate, the Przewalski horse or Takhi (Equus ferus przewalskii Poljakow). f(H) varied periodically throughout the year with a twofold increase from a mean of 44 beats min(-1) during December and January to a spring peak of 89 beats min(-1) at the beginning of May. LA increased from 23% per day during December and January to a mean level of 53% per day during May, and declined again thereafter. Daily mean subcutaneous body temperature (T(s)) declined continuously during winter and reached a nadir at the beginning of April (annual range was 5.8 degrees C), well after the annual low of air temperature and LA. Lower T(s) during winter contributed considerably to the reduction in f(H). In addition to thermoregulation, f(H) was affected by reproduction, LA, HI and unexplained seasonal variation, presumably reflecting to some degree changes in organ mass. The observed phase relations of seasonal changes indicate that energy expenditure was not a consequence of energy uptake but is under endogenous control, preparing the organism well in advance of seasonal energetic demands.
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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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Austin, N. P., & Rogers, L. J. (2014). Lateralization of agonistic and vigilance responses in Przewalski horses (Equus przewalskii). Applied Animal Behaviour Science, 151, 43–50.
Abstract: tEye and limb preferences were scored in the closest undomesticated relative of Equuscaballus using the same methods as used previously to study laterality in feral horses.Observations were made of 33 Przewalski horses (Equus ferus przewalskii) (male N = 20,female N = 13) living under natural social conditions on a large reserve in France. Signifi-cant left-eye/side biases were found in agonistic interactions within harem bands (M ± SEbias to left 58% ± 0.01 for threats, P < 0.001; 68% ± 0.05 for attacks; P < 0.001) and in stallionfights (threats, 52% ± 0.01 left, P < 0.001; attacks, 63% ± 0.02 left, P < 0.001): as many as 80%of the horses were significantly lateralized in attack responses within harem bands. Lat-erality of vigilance was measured as lifting up the head from grazing and turning it to theleft or right side: a directional bias to the left was found (M ± SE 53% ± 0.02 left, P < 0.001).Side bias in reactivity was calculated as the percent of head lifts above the level of thewithers on the left or right side and this was also left side biased (M ± SE 73% ± 0.03 left,P < 0.001). These results indicate right-hemisphere specialization for control of aggressionand responses to novelty. The left bias in attack scores within harem bands was strongerin males than females (P = 0.024) and in immature than adult horses (P = 0.032). Immaturehorses were also more strongly lateralized than adults in vigilance scores (P = 0.022), whichmay suggest that experience reduces these side biases. Our results show that Przewalskihorses exhibit left eye preferences, as do feral horses, and do so even more strongly thanferal horses. Considering feral and Przewalski horses together, we deduce that ancestralhorses had similar lateral biases. Also similar to feral horses, the Przewalski horses showedno significant forelimb preference at the group level or in the majority of horses at theindividual level, confirming the hypothesis that previously reported limb preferences indomestic breeds are entrained or generated by breed-specific selection.
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Berger, A. (2015). Evaluation of living conditions in free running animals by chronobiological analysis of continuously recorded behavioural data. In Proceedings of the 3. International Equine Science Meeting.
Abstract: We developed a biorhythmical method to assess behaviour patterns and to evaluate living conditions of animals. All kinds of continuous and equidistant long-term recordings of behaviour are suitable for this method. As simple behavioural parameters, such as motor activity, can be conveniently recorded by telemetry from wild animals now, it is possible to investigate stressors by analysing its biorhythmic structure. It is the purpose of this report to describe the basic idea, and the procedure, and to give some examples of application measured on Przewalski horses in an Semireserve.
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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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Brinkmann, L., Gerken, M., & Riek, A. (2015). Energetic adaptations of Shetland pony mares. In Proceedings of the 3. International Equine Science Meeting.
Abstract: Recent results suggest that wild Northern herbivores exhibit signs of a hypometabolism during times of low ambient temperature and food shortage in order to reduce their energetic needs. However, there are speculations that domestic animals lost the ability to reduce energy expenditure. To examine energetic and behavioural responses 10 Shetland pony mares were exposed to different environmental conditions (summer and winter). During winter ponies were allocated into two groups receiving two different food quantities (60% and 100% of maintenance energy requirement). We measured the field metabolic rate, water turn over, body temperature, locomotor activity, lying time, resting heart rate, body mass and body condition score.
In summer, the field metabolic rate of all ponies (FMR; 63.4±15.0 MJ/day) was considerably higher compared with food restricted and control animals in winter (24.6±7.8 and 15.0±1.1 MJ/day, respectively). Furthermore, during summer, locomotor activity, resting heart rate and total water turnover were significantly elevated (P<0.001) compared with winter. Animals receiving a reduced amount of food (N=5) reduced their FMR by 26% compared with control animals (N=5) to compensate for the decreased energy supply. Furthermore, resting heart rate, body mass and body condition score were lower(29.2±2.7 beats/min, 140±22 kg and 3.0±1.0 points, respectively) than in control animals (36.8±41 beats/min, 165±31 kg, 4.4±0.7 points; P<0.05). While no difference could be found in the observed behaviour, nocturnal hypothermia was elevated in restrictively fed animals. Our results indicate that ponies adapt to different climatic conditions by changing their metabolic rate, behaviour and some physiological parameters. When exposed to energy shortage, ponies, like wild herbivores, exhibited hypometabolism and nocturnal hypothermia. Keywords: Body temperature, Energy expenditure, Food restriction, Hypometabolism, Locomotor activity, Shetland pony |
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Burla, J. - B., Ostertag, A., Schulze Westerath Niklaus, H., & Hillmann, E. (2012). Validation of the MSR145W Data Logger for Gait Determination in Horses (Equus caballus). In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: Group housed horses at a stud farm/riding stable in Belgium were observed on 17 days between 21 February and 25 April 2008, totalling 54hr25min of detailed data. The original group consisted of 8 Irish Cob mares, 1 Warmblood mare, 1 Arabian gelding and 2 Arabian mares. The group had been established in December 2007. During the course of the study 5 horses were removed from the group and 2 foals were born. 3 highly pregnant mares were housed adjacent to the group for part of the period. Horses were regularly used for lessons. Available surface area differed with the group on pasture at the end. Continuous all occurrence sampling of 10 agonistic and 2 affiliative behaviours was carried out for all group members present. Overall the group showed a frequency of 44.75 agonistic interactions per hour and 11.25 affiliative per hour. Of those agonistic interactions 46.3% were threats while 47% were less active interactions (displacement, being avoided), leaving only 6.7% more aggressive interactions ( mainly biting, some kicking and chasing). The effect on acting agonistically was not significant for age (p=0.1591) and borderline significant for density (p=0.0627). The analysis of the frequency of affiliative interactions showed there is no significant effect of age (p=0.1865) or density (p=0.7923). Agonistic and affiliative interactions were not significantly correlated (p=0.72). Affiliative behaviour a horse received showed a borderline effect (p=0.0787) on agonistic behaviour, as did the interaction between received agonistic and affiliative interactions (p=0.0725). Received agonistic interactions had a borderline negative effect (p=0.0656) on affiliative behaviour. A dominance hierarchy was calculated based on agonistic interactions using Empirical Bayes’ estimates based on Poisson regression with random effects. Agonistic behaviour expressed to other horses was significantly effected by relative rank (p=0.0243). Overall horses tended to be 3.7 times more aggressive towards lower ranking horses compared to higher ranking horses. Affiliative behaviour shown to other horses was not significantly influenced by the rank of the social partner (p=0.7915). Some individuals did show a significant effect whereby they showed more affiliative behaviour towards lower ranking individuals. This study was a small project to look at a practical situation of riding horses being kept in group housing. The frequent changes in group composition and available surface made it possible to look at agonistic and affiliative interactions in such circumstances. This is useful as instability in group composition is often used as main reason not to keep horses in social groups. The results from this study showed a borderline effect of density on agonistic behaviour. In reality it was also influenced by practical details, like a narrow paddock with only 2 hay crates on the smallest surface. Rank in a dominance hierarchy, based on agonistic behaviour, had a significant effect on the agonistic behaviour expressed towards higher or lower ranking horses. No injuries or escalating fights were observed. This study shows it is possible to keep a group of riding horses in a social context without excessive aggression.
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Burn, C. C., Dennison, T. L., & Whay, H. R. (2010). Relationships between behaviour and health in working horses, donkeys, and mules in developing countries. Appl. Anim. Behav. Sci., 126(3-4), 109–118.
Abstract: Recent studies raise serious welfare concerns regarding the estimated 93.6 million horses, donkeys and mules in developing countries. Most equids are used for work in poor communities, and are commonly afflicted with wounds, poor body condition, respiratory diseases, parasites, dental problems, and lameness. Non-physical welfare problems, such as fear of humans, are also of concern. Interventions to improve working equine welfare aim to prioritise the conditions that cause the most severe impositions on the animals' subjectively experienced welfare, but data identifying which conditions these may be, are lacking. Here we describe a stage in the validation of behavioural welfare indicators that form part of a working equine welfare assessment protocol. Over 4 years, behavioural and physical data were collected from 5481 donkeys, 4504 horses, and 858 mules across nine developing countries. Behaviours included the animals' general alertness, and their responses to four human-interaction tests, using the unfamiliar observer as the human stimulus. Avoidance behaviours correlated significantly with each other across the human-interaction tests, with 21% of animals avoiding the observer, but they showed no associations with likely anthropogenic injuries. Over 13% of equids appeared [`]apathetic': lethargic rather than alert. Measures of unresponsiveness correlated with each other across the five tests, and were associated with poor body condition, abnormal mucous membrane colour, faecal soiling, eye abnormalities, more severe wounds, and older age, depending on the equine species. This suggests that working equids in poor physical health show an unresponsive behavioural profile, consistent with sickness behaviour, exhaustion, chronic pain, or depression-like states.
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Carlstead, K., & Brown, J. L. (2005). Relationships between patterns of Fecal corticoid excretion and behavior, reproduction, and environmental factors in captive black (Diceros bicornis) and white (Ceratotherium simum) rhinoceros. Zoo Biol., 24(3), 215–232.
Abstract: Mortality is high in zoo-housed black rhinoceros (Diceros bicornis), and the reproductive rates of captive white rhinoceros (Ceratotherium simum) are unsustainably low. To determine the possible role of stress in the causation of these problems, we analyzed weekly fecal samples collected for 1 year from black (10 males and 16 females) and white (six males and 13 females) rhinoceroses at 16 zoos for corticoid metabolite concentrations. Fecal corticoid profiles were examined in relation to behavior as rated by keepers in a questionnaire, luteal phase ovarian cycles of females (Brown et al., 2001), and socioenvironmental factors. We compared individual fecal corticoid profiles by examining hormone means and variability (i.e., standard deviation (SD) and coefficient of variation (CV)). For the black rhinos, higher mean corticoid concentrations were found at zoos where rhinos were maintained in enclosures that were exposed to the public around a greater portion of the perimeter. Higher variability in corticoid excretion was correlated with higher rates of fighting between breeding partners and higher institutional mortality rates. Black rhino pairs that were kept separated exhibited lower corticoid variability and less fighting activity when they were introduced during female estrous periods compared to pairs that were kept together every day. For white rhinos, significantly lower mean corticoids were found for individuals that rated higher on “friendliness to keeper.” Higher corticoid variability was found in noncycling as compared to cycling white rhino females. Noncycling females exhibited higher rates of stereotypic pacing and lower frequencies of olfactory behaviors. Interindividual differences in mean corticoids in both species appeared to be related to responsiveness to humans, whereas corticoid variability was related to intraspecific social relationships. More importantly, high corticoid variability appeared to be an indicator of chronic or “bad” stress, because of its association with potentially deleterious consequences in each species (i.e., fighting and mortality (black rhino), and reproductive acyclicity (white rhino)). Our results provide evidence that social stressors may cause chronic stress in black and white rhinos, and that this contributes to the captive-population sustainability problems observed in each species. Zoo Biol 0:1–18, 2005. © 2005 Wiley-Liss, Inc.
Keywords: stress; adrenal activity; olfactory behavior; ovarian activity; reproduction; mortality
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Chaplin, S. J., & Gretgrix, L. (2010). Effect of housing conditions on activity and lying behaviour of horses. animal, 4(5), 792–795.
Abstract: Housing conditions for horses impose various levels of confinement, which may compromise welfare. Lying behaviour and activity can be used as welfare indicators for domestic animals and rebound behaviour suggests a build-up of motivation resulting from deprivation. The objective of this study was to determine if activity and lying behaviour of horses are affected by housing conditions and to investigate the occurrence of rebound behaviour after release from confinement. Eight horses were subjected, in pairs, to each of four experimental treatments; paddock (P), fully stabled (FS), partly stabled (PS) and yard (Y). Each horse received 6 days acclimatisation prior to the 24 h recording period. Time spent in lying and activity were electronically recorded using a tilt switch and motion sensor connected to a data logger worn on the horse's left foreleg. Time spent active during the first 5 min of release from stable to paddock in the PS treatment (days 1 and 5) and at the same time of day in the P treatment was used as a measure of rebound behaviour. Effect of housing conditions on total time spent active was highly significant (FS = 123 s, PS = 158 s, Y = 377 s, P = 779 s, P < 0.001). Housing conditions did not significantly affect total time spent lying (P = 0.646). Horses were significantly more active, compared with baseline paddock behaviour, on release from stabling on both days 1 (P = 0.006) and 5 (P = 0.025) of PS treatment. These results suggest that activity patterns of horses, but not lying behaviour, are affected by the housing conditions tested and that rebound activity occurs in horses after a period of confinement.
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