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Houpt, K.A.; Keiper, R. |
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The position of the stallion in the equine dominance hierarchy of feral and domestic ponies. |
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1982 |
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Journal of Animal Science |
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J. Anim Sci |
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54 |
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5 |
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945-950 |
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Keiper, R.R.; Sambraus, H.H. |
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Title |
The stability of equine dominance hierarchies and the effects of kinship, proximity and foaling status on hierarchy rank |
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1986 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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16 |
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2 |
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121-130 |
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Dominance hierarchies were determined in four bands of feral horses living on Assateague Island. The bands varied in size from 10 to 16 horses, and consisted of one stallion, several mares and their offspring. The animals ranged in age from less than 1 to over 18 years. Field observation of all social interactions during the summer of 1981 was used to determine dominance. 1981 hierarchies for three of the bands were compared with hierarchies determined for the same bands in 1978, and showed that hierarchies change over time. Age was significantly correlated with rank. Mares with foals did not rank any higher in the hierarchies than mares without foals. Kinship did not appear to have an effect on dominance rank either, since neither juvenile nor adult offspring ranks correlated with the ranks of their mothers. The band stallion was not the highest-ranking animal of any band, but the location of the stallion peripheral to the main body of the band, the nature of his interactions with band members, and his length of residence in the band may have contributed to his low rank. |
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683 |
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Feh, C. |
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Title |
Social behaviour and relationships of Prezewalski horses in Dutch semi-reserves |
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1988 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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21 |
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1-2 |
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71-87 |
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A short-term study was made of 2 groups of Przewalski horses, a bachelor group of 4 juvenile stallions in Ooij Polder and a harem group of 1 stallion and 4 mares. All social interactions were recorded and the nearest and farthest neighbour was noted. Correspondence analysis was used to determine what parameters determined the relationships among the horses. There was a linear hierarchy among the bachelor stallions. The dominant stallion of the group was also the oldest. The hierarchy was not linear in the harem group, and the 3-year-old stallion was subordinate to the 5-year-old mares. He was also most likely to be farthest from other horses. The mares of the same age, who had also arrived in the park at the same time, tended to be one another's nearest neighbours. The frequency of aggression is higher among Przewalski horses than among domestic horses of similar ages. Correspondence analysis revealed that head-threats and other forms of aggression accounted for more of the variance in the data than any other behaviour, but submission, play and social interactions also contributed. |
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764 |
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Keiper, R.R. |
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Social interactions of the Przewalski horse (Equus przewalskii Poliakov, 1881) herd at the Munich Zoo |
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1988 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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21 |
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1-2 |
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89-97 |
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Data were collected on 972 aggressions and 233 acts of mutual grooming in a herd of 9 Przewalski horses in the Hellabrunn Tierpark in Munich, West Germany. The herd was composed of 1 adult stallion, 5 adult mares and 3 foals. A distinct linear dominance hierarchy was present in the herd, with the stallion being the top-ranking animal. Age was significantly correlated with rank. Almost 40% of all aggressions consisted of herding actions by the stallion. Threats to bite (20% of all aggressions) and threats to kick (11.4%) were next in frequency of occurrence. Most mutual grooms (71%) involved grooming the front part of the body. Although mutual grooming may be used to appease higher-ranked animals, most grooming bouts were between related horses. Foals initiate 47.6% of all allogrooming. Mutual grooming may reduce weaning conflict between a mare and her foal or may result in female coalitions that defend against predators or aggression by the herd stallion. |
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775 |
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Keiper, R.; Receveur, H. |
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Title |
Social interactions of free-ranging Przewalski horses in semi-reserves in the Netherlands |
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1992 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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33 |
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4 |
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303-318 |
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Social interactions were recorded in two bands of free-ranging Przewalski horses living on large (greater than 30 ha) pastures in the Netherlands. The average number of aggressive interactions per hour was 8.86 at Lelystad and 10.36 at Noorderheide. The most common aggressive interactions were lower intensity, lower cost displacements (17.2% of all aggressive acts at Lelystad, 13.2% at Noorderheide), threats to bite (42.3% and 40.7%, respectively) and threats to kick (15.4% and 23.9%, respectively). Analysis of aggression revealed that a clear, linear dominance hierarchy was present in each band. For each band there was a positive and highly significant correlation between the age of a horse and its rank in the hierarchy. In each band, the stallion was not the highest ranked horse. Non-agonistic behaviors exceeded the number of agonistic interactions (1253 vs. 558 for Lelystad; 1257 vs. 995 at Noorderheide). There was a negative correlation between the rank of a horse in the dominance hierarchy and the number of non-agonistic behaviors displayed. The group displaying the highest number of non-agonistic interactions were foals (48.9% of total non-agonistic behaviors at Lelystad; 51.1% at Noorderheide). The non-agonistic/agonistic ratio was greater than 1 for yearlings and the band stallion, as was also the case for foals, but was less than 1 for males. |
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778 |
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Hogan, E.S.; Houpt, K.A.; Sweeney, K. |
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The effect of enclosure size on social interactions and daily activity patterns of the captive Asiatic wild horse (Equus przewalskii) |
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1988 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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21 |
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1-2 |
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147-168 |
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Two herds of Przewalski horses at the Minnesota Zoological Garden were observed during 1980 in each of 2 enclosures that differed in size. The larger enclosure was a 3.4-ha pasture; the smaller enclosure was a 17 x 30-m grass-less pen. One herd was composed of a stallion, 3 adult mares and 2 foals. The other consisted of a stallion and 2 mares. All occurrences of aggression, mutual grooming and snapping were recorded, and 5-min scan-samples of the activity state of each horse were taken. The time budgets, frequency of aggression and frequency of mutual grooming differed significantly with enclosure size for both herds. More time was spent pacing and milling in the smaller enclosure, and the frequency of aggressions and of mutual grooming was also higher. Only the foals exhibited snapping; frequency of snapping did not vary with enclosure size. More time was spent feeding in the larger enclosure. Provision of hay in the smaller enclosure eliminated the differences in time spent feeding. A second study was conducted during the spring of 1984 in an intermediate-sized enclosure, 0.4 ha, a sub-division of the pasture on which the horses were kept in 1980. One herd consisted of a stallion, 2 mares and 2 yearlings; the other consisted of a stallion, 3 mares and a foal. One of the stallions and all of the mares were those studied in 1980, but that stallion and one of the mares were in different herds than they had been in 1980. The frequency of aggression was similar to that observed in 1980. |
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780 |
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Tilson, R.L.; Sweeny, K.A.; Binczik, G.A.; Reindl, N.J. |
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Buddies and bullies: Social structure of a bachelor group of Przewalski horses |
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Journal Article |
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1988 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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21 |
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1-2 |
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169-185 |
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The Minnesota Zoological Garden established a bachelor (i.e. all-male) group of 8 Przewalski horses (Equus przewalskii) in 1984. Possible methods for mixing males emerged through preliminary introductions and review of the relevant literature. These included: (1) grouping colts (<=3 years old) after removing them from their natal bands; (2) introducing adult stallions to colts in spacious, controlled environments to reduce the risk of combat-related injury or death associated with largely unpredictable adult temperament; (3) introducing more than 1 animal at a time to established groups. Based on the frequency and orientation of agonistic interactions observed among the bachelors, a linear dominance hierarchy was discerned among the 4 adult stallions; all 4 colts were determined to be of equally low or seemingly non-ranking status. Social rank among the adults was most influenced by individual temperament, and contention for rank was limited almost exclusively to the 3 lower-ranking stallions; the alpha male participated in 73% of all agonistic interactions observed, but was identified as the recipient in less than 3% of these interactions. Intra-band associations were detected in the bachelor group. The most prevalent of these was among the 4 colts, found together in over 70% of the observations. Other affinities commonly observed were between the colts (as a sub-group) and different stallions. The dominant stallion, the colts' most frequent companion, actively discouraged the 3 lower-ranking stallions from associating with the colts. Further, the colts were subjected to significantly fewer agonistic behaviors than the non-alpha stallions (per colt=1.05+/-0.24 h-1; per non-alpha stallion=2.01+/-0.04 h-1). These findings support the hypothesis developed by other workers for ungulates that young bachelors are sheltered from aggression by association with an alpha male. With one exception, adult associations were rarely noted. Although combat-related injuries (superficial wounds and temporary lameness) were common among the bachelors, few required medical attention. No permanent disabilities or deaths have occurred since the group's inception. The existence of social organization and intra-band associations in the bachelor group support the suggestion that such groups were once a natural part of this species' social milieu. Bachelor groups are therefore recommended as an ethologically sound, space- and cost-effective means of managing “surplus” (i.e. non-harem) male Przewalski horses. |
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Baker, A.E.M.; Crawford, B.H. |
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Observational learning in horses |
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1986 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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15 |
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1 |
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7-13 |
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This experiment was designed to determine if a horse could learn the location of grain by watching another horse find grain in one of two feed buckets. Both experimental and control groups contained 9 quarter horses consisting of five 2-year-old mares, two 2-year-old geldings, and two 3-year-old geldings. Two mature geldings were used as “demonstrators”. An “experimental” was a horse that could watch three times daily another horse, the “demonstrator”, choose between and eat grain from a black or white bucket, only one of which contained grain. A “control” was a horse that could watch a demonstrator in the same arena for 3 min daily when both feed buckets were removed. When the demonstrator was removed on each of 15 successive days, the experimental or control horse was given five trials to determine if it could find the feed bucket with grain. No significant difference between experimentals and controls occurred for both first and total correct choices and for time to reach the feed bucket with grain. We conclude that no observational learning occurred. This experiment was also used to determine if the identity of horses that learned rapidly by trial and error could be predicted by the time it took to reach the feed bucket with grain. Data from the last three trials of experimentals and controls were combined. Significantly less time to find feed was needed by horses with more than the median number of correct choices. Both number of correct choices and time needed to contact a feed bucket summed over the first 5 days accurately predicted the same data summed over the last 10 days. We conclude that horses that learn rapidly by trial and error make correct choices rapidly, and that these horses can by identified after 5 days of testing. |
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Heird, J.C.; Lokey, C.E.; Cogan, D.C. |
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Repeatability and comparison of two maze tests to measure learning ability in horses |
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1986 |
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Applied Animal Behaviour Science |
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Appl. Anim. Behav. Sci. |
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16 |
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2 |
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103-119 |
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Sixteen Quarter Horses were randomly divided into two groups after sorting by age and sex. After a 10-day preconditioning period, each animal was scored for emotionality and trainability. Each group then completed a series of learning tasks in a modified T-maze for 20 consecutive days. Group P/D was initially tested on a simple place-learning task, while Group D/P was trained in a visual discrimination task. The groups were tested alternately on the two tasks with 10-day extinction periods between each task. Upon reaching a criterion of 11 of 12 correct responses (the last 8 responses consecutive), a horse was retired for the day. If this criterion was not attained, the horse completed 20 trials. Learning occurred at a faster rate on the discrimination tasks compared to the gradual learning curves observed on place tasks. Animals learned more rapidly and reached higher levels of performance as the series of tasks progressed. Trainability and emotionality scores tended to predict the final level of learning achieved. Correlations of performance ranks within emotionality and training groups were higher between tasks of the same type than between the different tasks. Rank correlations between odd and even days on each task indicated that the within-group rankings were more consistent on the discrimination task than on the place task. |
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C. K. Hemelrijk, |
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An individual-orientated model of the emergence of despotic and egalitarian societies |
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1999 |
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Proceedings of the Royal Society B: Biological Sciences |
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Proc. Roy. Soc. Lond. B Biol. Sci. |
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266 |
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1417 |
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361-361 |
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Single behavioural differences between egalitarian and despotic animal societies are often assumed to reflect specific adaptations. However, in the present paper, I will show in an individual-orientated model, how many behavioural traits of egalitarian and despotic virtual societies arise as emergent characteristics. The artificial entities live in a homogeneous world and only aggregate, and upon meeting one another and may perform dominance interactions in which the effects of winning and losing are self-reinforcing. The behaviour of these entities is studied in a similar way to that of real animals. It will be shown that by varying the intensity of aggression only, one may switch from egalitarian to despotic virtual societies. Differences between the two types of society appear to correspond closely to those between despotic and egalitarian macaque species in the real world. In addition, artificial despotic societies show a clearer spatial centrality of dominants and, counter-intuitively, more rank overlap between the sexes than the egalitarian ones. Because of the correspondence with patterns in real animals, the model makes it worthwhile comparing despotic and egalitarian species for socio-spatial structure and rank overlap too. Furthermore, it presents us with parsimonious hypotheses which can be tested in real animals for patterns of aggression, spatial structure and the distribution of social positive and sexual behaviour. |
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