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Mader, D. R., & Price, E. O. (1980). Discrimination learning in horses: effects of breed, age and social dominance. J. Anim Sci., 50(5), 962–965.
Abstract: The discrimination learning ability of Quarter Horses and Thoroughbreds was compared by means of visual cues in a three-choice test with food as a reward. Quarter Horses learned significantly faster than Thoroughbreds, and learning progressed more rapidly for both breeds in a second discrimination task. Significant negative correlations were observed between age and rate of learning. Quarter Horses tended to be less reactive than Thoroughbreds, but individual emotional reactivity ratings and learning scores were not correlated. No correlation was found between social dominance and learning scores. Learning studies with horses may provide a better understanding of the behavioral traits that influence trainability in this species.
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VanDierendonck, M. C. (2006). General Discussion (Vol. Chapter 7). Universiteit Utrecht.
Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
Keywords: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
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VanDierendonck, M. C. (2006). Interventions in social behaviour in a herd of mares and geldings (Vol. Chapter 6). Universiteit Utrecht.
Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
Keywords: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
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VanDierendonck, M. C. (2006). Diff erences in social behaviour between late pregnant, post-partum and barren mares in a herd of Icelandic horses (Vol. Chapter 5). Ph.D. thesis, , Universiteit Utrecht.
Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
Keywords: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
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VanDierendonck, M. C. (2006). Social relationships in a group of horses without a mature stallion (Vol. Chapter 4). Universiteit Utrecht.
Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
Keywords: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.
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Krueger, K. (2007). Behaviour of horses in the “round pen technique”. Appl. Anim. Behav. Sci., 104(1-2), 162–170.
Abstract: I investigated the behavioural background of the way horses learn to follow humans in the “round pen technique” suggested by “horse whisperers” as a gentle method for initial horse training. Though the practicability of this technique has been adequately demonstrated in the past, the horses' behaviour during such training has not yet been documented in detail. In a riding arena, horses, that did not follow the trainer immediately, were chased away so that they galloped around the trainer. Galloping horses showed specific behaviour such as turning the ear to the trainer, chewing, licking, and stretching head and throat downwards. In subsequent trials horses needed to be chased for less time and finally followed immediately, even when conditions were changed or the trainer was replaced by another person. This suggests that horses learn to follow in this particular situation and also show some generalisation. However, following did not occur on a pasture even after several successful trials in the riding arena.
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Becker-Birck, M., Schmidt, A., Wulf, M., Aurich, J., von der Wense, A., Möstl, E., et al. (2013). Cortisol release, heart rate and heart rate variability, and superficial body temperature, in horses lunged either with hyperflexion of the neck or with an extended head and neck position. Journal of Animal Physiology and Animal Nutrition, 97(2), 322–330.
Abstract: Bringing the head and neck of ridden horses into a position of hyperflexion is widely used in equestrian sports. In our study, the hypothesis was tested that hyperflexion is an acute stressor for horses. Salivary cortisol concentrations, heart rate, heart rate variability (HRV) and superficial body temperature were determined in horses (n = 16) lunged on two subsequent days. The head and neck of the horse was fixed with side reins in a position allowing forward extension on day A and fixed in hyperflexion on day B. The order of treatments alternated between horses. In response to lunging, cortisol concentration increased (day A from 0.73 ± 0.06 to 1.41 ± 0.13 ng/ml, p < 0.001; day B from 0.68 ± 0.07 to 1.38 ± 0.13 ng/ml, p < 0.001) but did not differ between days A and B. Beat-to-beat (RR) interval decreased in response to lunging on both days. HRV variables standard deviation of RR interval (SDRR) and RMSSD (root mean square of successive RR differences) decreased (p < 0.001) but did not differ between days. In the cranial region of the neck, the difference between maximum and minimum temperature was increased in hyperflexion (p < 0.01). In conclusion, physiological parameters do not indicate an acute stress response to hyperflexion of the head alone in horses lunged at moderate speed and not touched with the whip. However, if hyperflexion is combined with active intervention of a rider, a stressful experience for the horse cannot be excluded.
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Kampmann, S., Hampson, B. A., & Pollitt, C. C. (2013). Population dynamics of feral horses (Equus caballus) following above-average rainfall in a semi-arid environment of Australia. Aust Vet J, 91(11), 482–487.
Abstract: Background Recent record rainfall in much of semi-arid Central Australia is the most likely reason for a feral horse population increase in excess of normal. Uncontrolled numbers of feral horses have habitat degradation and animal welfare implications. Objectives The aims of this study were to investigate the social structure of feral horses and assess their population growth rate following unseasonably high rainfall. Methods The study area was 4000 km2 of unmanaged, semi-arid country in Central Australia (latitude 24.50°S, longitude 132.10°E). Horses were identified by descriptive features from ground searches, movement-activated cameras and ‘hides’ positioned at key water holes. Wherever possible, sex and age categories were documented. Population growth rate was estimated by the number of foals divided by the number of horses older than 1 year in the observed population. Results A total of 1424 horses were identified and categorised, of which 335 were foals born within the current year. Only 123 juveniles were identified. Of the adult horses, 53.4% were male and 46.6% were female and this differed from parity (P = 0.04). Of the mares, 71.9% had a foal at foot and the population growth rate was 29.5%. Conclusions With a sustained population growth rate of 29.5%, this population of feral horses will more than double within 3 years. The high population increase will likely have a detrimental effect on native fauna and flora and the fragile, semi-arid ecosystems of Central Australia. After a period of high rainfall and plentiful resources, ‘normal’ drought conditions will return and many feral horses will starve and die as they compete for limited resources.
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Weissing, F. J. (2011). Animal behaviour: Born leaders. Nature, 474(7351), 288–289.
Abstract: Social animals face a dilemma. To reap the benefits of group living, they have to stay together. However, individuals differ in their preferences as to where to go and what to do next. If all individuals follow their own preferences, group coherence is undermined, resulting in an outcome that is unfavourable for everyone. Neglecting one's own preferences and following a leader is one way to resolve this coordination problem. But what attributes make an individual a 'leader'? A modelling study by Johnstone and Manica1 illuminates this question.
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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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