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Pluhacek, J., Bartos, L., & Culik, L. (2006). High-ranking mares of captive plains zebra Equus burchelli have greater reproductive success than low-ranking mares. Appl. Anim. Behav. Sci., 99(3-4), 315–329.
Abstract: Plains zebra live in harems that include one to six adult mares. Between these mares is a strong order of social hierarchy. The social rank of an equid mare is typically correlated with her age. Further, high-ranking captive plains zebra mares produce more surviving offspring than low-ranking mares. The objectives of this study were to, first, examined the factors that influence social rank of captive plains zebra mares, and second, test if high-ranking mares conceive earlier and if they have shorter inter-birth intervals than low-ranking ones. We observed three herds of captive plains zebra (a total of 18 mares) at the Dvur Kralove Zoo, Czech Republic. During the 831 h of observation, we recorded 1713 aggressive interactions (biting and offensive kicking) between the mares. These data were used to determine, for each mare, the total number of mares that dominated her in each period of social stability. The GLMM model revealed that older mares were dominated by a lower number of mares than the younger mares. We also found that the probability that a mare would conceive declined with the increasing number of dominant mares. Further, we tested the relationship between the number of dominant mares and the inter-birth interval using 29 intervals for 15 mares. These inter-birth intervals were divided into two groups. When a stallion was continuously present in the herd, the intervals lasted from one birth to the next birth (natural intervals). When a herd was without a stallion, the intervals lasted from the release of the stallion into the herd to the birth of foal (stallion-influenced intervals). The analysis revealed that the inter-birth intervals decreased with an increasing number of dominant mares and the natural intervals decreased with an increasing number of offspring successfully reared by a mare. This finding is the first one in equids and contributes to the previous findings that suggest that social status influences reproductive success.
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Kimura, R. (1998). Mutual grooming and preferred associate relationships in a band of free-ranging horses. Appl. Anim. Behav. Sci., 59(4), 265–276.
Abstract: Preferred associate (nearest neighbour) and mutual grooming relationships among mares, in an isolated family band of free-ranging horses (Equus caballus), were studied, and the structural and functional differences between these two relationships were examined. The frequent partners accompanying the mare were not the same in both these relationships (p<0.05) and mares changed their partners during the study period between 1988-1990. Individual horses of similar rank tended to remain closer together in both winter and summer. Aggressive-submissive behaviour was so infrequent during spring that rank determinations could not be made; however, in fall, although rank could be determined, rank was not correlated with nearest neighbour. Three subgroups, based on preferred associate relationships in summer, fall and winter, directly reflected the age and social rank of the mares in the group. Individual horses of higher rank tended to have many partners in winter, while individuals of lower rank had fewer. There was no significant correlation between the frequency of mutual grooming and individual rank. The mutual grooming relationship was strongly influenced by seasonal changes as the relative amount of grazing time per day increased. Thus, the frequency of mutual grooming was lowest in winter and highest in summer. The mutual grooming relationship was based on the bonds between individual horses, which were little influenced by social rank. Lower ranking individuals tended to have a greater variety of grooming partners in summer.
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Bräuer, J., Call, J., & Tomasello, M. (2004). Visual perspective taking in dogs (Canis familiaris) in the presence of barriers. Appl. Anim. Behav. Sci., 88(3-4), 299–317.
Abstract: Previous studies have shown that dogs have developed a special sensitivity to the communicative signals and attentional states of humans. The aim of the current study was to further investigate what dogs know about the visual perception of humans and themselves. In the first two experiments we investigated whether dogs were sensitive to the properties of barriers as blocking the visual access of humans. We presented dogs with a situation in which a human forbade them to take a piece of food, but the type and orientation of the barrier allowed the dog to take the food undetected in some conditions. Dogs differentiated between effective and ineffective barriers, based on their orientation or the particular features of the barriers such as size or the presence of window. In the third study we investigated whether dogs know about what they themselves have seen. We presented subjects with two boxes and placed food in one of them. In the Seen condition the location of the food was shown to the dogs while in the Unseen condition dogs were prevented from seeing the destination of the food. Before selecting one of the boxes by pressing a lever, dogs had the opportunity to seek extra information regarding the contents of the boxes, which would be particularly useful in the condition in which they had not seen where the food was hidden. Dogs rarely used the opportunity to seek information about the contents of the box before making their choice in any condition. Therefore, we found no evidence suggesting that dogs have access to what they themselves have seen, which contrasts with the positive evidence about visual perspective taking in others from the first two experiments and previous studies.
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Goodwin, D., Davidson, H. P. B., & Harris, P. (2007). A note on behaviour of stabled horses with foraging devices in mangers and buckets. Appl. Anim. Behav. Sci., 105(1-3), 238–243.
Abstract: Processed feed for stabled horses is usually presented in buckets or mangers, and is easily and rapidly consumed. Foraging devices based on the Edinburgh foodball can be used to provide part of the ration. Current designs are all placed on the floor, raising concerns regarding ingestion of foreign materials along with the dispensed food. Alternative devices were evaluated, when presented within suitable, clean containers to prolong food-handling times but avoid such issues. In four Latin square designed replicated trials we investigated behaviour of 12 stabled horses with three foraging devices. These were separately presented for 5 min, varied in sensory complexity (round, square, polyhedral) and contained 500 g high fibre pellets. In Trials 1 and 2 six geldings were presented with devices in buckets then mangers. All individuals foraged successfully from at least one device and behaviour was compared. However, all individuals exhibited some frustration while using the devices (either pawing or biting them). Horses frequently removed the devices from the buckets in Trial 1 terminating these sessions. In Trial 2 mean device foraging duration was ranked polyhedral > round > square. Mean pawing rate in Trial 2 was calculated for horses (frequency of pawing per individual/summed duration manipulation and foraging) and was highest with square (0.11, npawers = 6). In Trial 3 six stabled mares were presented with the same foraging devices in mangers. Mean foraging duration with devices again ranked polyhedral > round > square. Mean pawing rate was highest with round device (0.08, npawers = 4). Trial 4 investigated behaviour of six horses when devices initially containing five high fibre pellets became empty. Mean foraging duration with devices ranked round > polyhedral > square. Mean pawing rate was highest with square device (0.11, npawers = 4). All horses foraged successfully from at least one foraging device in buckets and mangers. Devices met initial objectives but the unpredictability of reward suggests a source of frustration and warrants further investigation.
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King, S. R. B. (2002). Home range and habitat use of free-ranging Przewalski horses at Hustai National Park, Mongolia. Appl. Anim. Behav. Sci., 78(2-4), 103–113.
Abstract: Przewalski horses (Equus ferus przewalskii), also known as takhi, were first re-introduced to the wild in Hustai National Park, Mongolia, in 1994. Since then the number of free harems increased to a maximum of seven; there are currently six (October 2000). The size of the home range of each of the harems changed among years and among seasons. The horses tended to settle in a home range close to where they were released although they explored the surrounding area. The use of the habitat within each home range changed through the day, with the horses grazing in the valleys during the morning and evening, and moving to higher places to stand rest and use as a refuge from heat and flies during the middle of the day. Range establishment and area, as well as habitat use are discussed.
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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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McGrogan, C., Hutchison, M. D., & King, J. E. (2008). Dimensions of horse personality based on owner and trainer supplied personality traits. Appl. Anim. Behav. Sci., 113(1), 206–214.
Abstract: Ratings on 36 personality descriptive adjectives were made on 100 companion horses by owners, trainers, and stable managers. The descriptors had been previously provided by a subset of 30 raters based on their own experience with horses. Principal components analysis revealed three statistically significant factors accounting for 59% of the variance. The first was a large, complex factor resembling the human Agreeableness factor. This factor contained four clusters of items reflecting obedience, kindness, sociality, and non-aggressiveness. The second factor resembled human Extraversion, although it also contained items reflecting intellect and curiosity. The third factor reflected Neuroticism or Emotionality. Factors similar to Agreeableness, Extraversion, and Emotionality are frequently observed in other taxa. The diverse composition of the Agreeableness factor indicating covariation among obedience, kindness, sociality, and non-aggressiveness may be a result of the intense artificial selection for all four components during the 6000 years of horse domestication.
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Simpson, B. S. (2002). Neonatal foal handling. Appl. Anim. Behav. Sci., 78(2-4), 303–317.
Abstract: Recent interest has focused on the advantage of intensively handling young foals as a means of producing more tractable horses, accustomed to humans and receptive to training. To date, the effect of this intensive handling, dubbed “imprint training” in the popular literature, has not been tested. The present study compares seven foals handled intensively as neonates with eight untreated controls. The handling protocol started from 2-8 h after birth and continued daily for a total of 5 days. The protocol consisted of a series of stimuli and experiences that were each repeated until the foal no longer resisted or reacted negatively. Subsequently, foals were tested before weaning, at 4 months of age. Results indicated that handled foals (HF) ranked higher than control foals (CF) in subjective ratings of calmness (*P<0.0117) and friendliness (*P<0.0001) and in several specific handling tasks (venipuncture *P<0.0220; placing in stock *P<0.0128). Although, in approach tests all foals but one allowed approach of a person to 4 m, significantly more HF approached the person than CF (P<0.0080). In stimulus tests, foals were presented specific stimuli to which they had been tested as neonates. Two of eight CF were too unruly and dangerous to test. Of foals that could be tested, CF required significantly more time to hook-up a heart rate monitor (**P<0.0055). Split-plot analysis indicated that HF had lower heart rates to initial left-sided stimuli, presented first, than CF (*P<0.0421). In response to right-sided stimuli, heart rate scores of CF were not significantly different from HF (P<0.2259), suggesting reduced reactivity over time due to a learning effect. Behavioral responses to specific stimuli did not differ between CF and HF, suggesting that neonatal handling has a general rather than specific effect on subsequent behavior. Cortisol concentrations were measured before and after testing and the difference calculated. All foals had higher post-testing levels than pre-testing levels. There was a significant difference between HF and CF, indicating greater reactivity among the CF (*P<0.050). In general, the results indicated that foals handled as neonates were more tractable and less reactive. Specific neonatal handling tasks, such as sticking a finger up the foal's nose or patting the bottom of the foot, seemed to have no beneficial effect on related tasks such as passing a nasogastric tube or tapping with a farrier's hammer at 4 months of age. Mechanisms for the observed effect of neonatal handling require further investigation.
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Nagy, K., Bodó, G., Bárdos, G., Bánszky, N., & Kabai, P. (2010). Differences in temperament traits between crib-biting and control horses. Appl. Anim. Behav. Sci., 122(1), 41–47.
Abstract: Recent studies have suggested that crib-biting in horses is associated with diminished capacity of learning or coping with stress. Such findings raise the question whether trainability, which is fundamentally important in practice, could also be affected by stereotypic behaviour. Trainability of a horse is difficult to assess in simple tests, however, it is reliably estimated by experienced riders. To assess trainability and other characteristics related to that, a questionnaire survey was conducted with the owners of 50 crib-biting and 50 control horses. Where possible, control horses were selected from the same establishment as crib-biters. Groups did not differ significantly regarding age, breed, gender, training level or usage. Principal component analysis revealed three main factors which can be labelled as [`]Anxiety', [`]Affability' and [`]Trainability'. The [`]Anxiety' factor consisted of the items [`]Nervousness', [`]Excitability', [`]Panic', [`]Inconsistent emotionality', [`]Vigilance', [`]Skittishness', and [`]Timidity'. [`]Affability' consisted of [`]Friendliness toward people', [`]Cooperation', [`]Docility' and [`]Friendliness toward horses'. [`]Trainability' involved [`]Concentration', [`]Trainability', [`]Memory', and [`]Perseverance'. Temperament traits were not affected by age, gender, breed or training level, but the usage of the horse and the presence of crib-biting behaviour had significant effects. Competition horses had lower level of [`]Anxiety' (p = 0.032) and higher level of [`]Trainability' (p = 0.068) than leisure horses. Crib-biting horses had significantly lower level of [`]Anxiety' than control horses (p < 0.001), while [`]Trainability' and [`]Affability' did not differ between groups (p = 0.823 and p = 0.543, respectively). Competition horses are more often exposed to novel environment and to frightening stimuli (e.g. colourful obstacles) than leisure horses and therefore might have also become more habituated to these types of stimuli. Coping with novel situation may be enhanced by defusing nervous behaviour by the more experienced riders of competition. Previous studies indicated crib-biting horses to be less reactive when challenged as compared to control horses. We suggest that the virtual calmness and lower nervousness of the crib-biting horses might be due to the passive coping style of these animals. [`]Affability' of horses might be more related to housing and management conditions than to crib-biting. Contrary to expectations, scores on [`]Trainability' had not coincided with the impaired learning of crib-biting horses reported in laboratory tests. However, previous behavioural tests on equine learning rarely had a direct relevance to the training abilities of the horses. Our results do not support crib-biting stereotypy to affect performance in training, which is a complex learning process involving cooperation and docility in the social environment.
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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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