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Author |
Hamilton, W.D. |
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Title |
Geometry for the selfish herd |
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Journal Article |
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1971 |
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Journal of theoretical biology |
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J. Theor. Biol. |
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31 |
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2 |
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295-311 |
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Animals; Anura; *Behavior, Animal; Breeding; Communication; Evolution; Fear; Metallurgy; *Models, Biological; Probability; Snakes; *Spatial Behavior |
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This paper presents an antithesis to the view that gregarious behaviour is evolved through benefits to the population or species. Following Galton (1871) and Williams (1964) gregarious behaviour is considered as a form of cover-seeking in which each animal tries to reduce its chance of being caught by a predator.
It is easy to see how pruning of marginal individuals can maintain centripetal instincts in already gregarious species; some evidence that marginal pruning actually occurs is summarized. Besides this, simply defined models are used to show that even in non-gregarious species selection is likely to favour individuals who stay close to others.
Although not universal or unipotent, cover-seeking is a widespread and important element in animal aggregation, as the literature shows. Neglect of the idea has probably followed from a general disbelief that evolution can be dysgenic for a species. Nevertheless, selection theory provides no support for such disbelief in the case of species with outbreeding or unsubdivided populations.
The model for two dimensions involves a complex problem in geometrical probability which has relevance also in metallurgy and communication science. Some empirical data on this, gathered from random number plots, is presented as of possible heuristic value. |
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0022-5193 |
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PMID:5104951 |
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771 |
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Jansen, T.; Forster, P.; Levine, M.A.; Oelke, H.; Hurles, M.; Renfrew, C.; Weber, J.; Olek, K. |
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Title |
Mitochondrial DNA and the origins of the domestic horse |
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Journal Article |
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2002 |
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Proceedings of the National Academy of Sciences of the United States of America |
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Proc. Natl. Acad. Sci. U.S.A. |
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99 |
Issue |
16 |
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10905-10910 |
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Animals; Animals, Domestic/classification/*genetics; Base Sequence; DNA, Complementary; *DNA, Mitochondrial; *Evolution, Molecular; Horses/classification/*genetics; Molecular Sequence Data; Phylogeny |
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The place and date of the domestication of the horse has long been a matter for debate among archaeologists. To determine whether horses were domesticated from one or several ancestral horse populations, we sequenced the mitochondrial D-loop for 318 horses from 25 oriental and European breeds, including American mustangs. Adding these sequences to previously published data, the total comes to 652, the largest currently available database. From these sequences, a phylogenetic network was constructed that showed that most of the 93 different mitochondrial (mt)DNA types grouped into 17 distinct phylogenetic clusters. Several of the clusters correspond to breeds and/or geographic areas, notably cluster A2, which is specific to Przewalski's horses, cluster C1, which is distinctive for northern European ponies, and cluster D1, which is well represented in Iberian and northwest African breeds. A consideration of the horse mtDNA mutation rate together with the archaeological timeframe for domestication requires at least 77 successfully breeding mares recruited from the wild. The extensive genetic diversity of these 77 ancestral mares leads us to conclude that several distinct horse populations were involved in the domestication of the horse. |
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Biopsytec Analytik GmbH, Marie-Curie-Strasse 1, 53359 Rheinbach, Germany. jansen@biopsytec.com |
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0027-8424 |
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PMID:12130666 |
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772 |
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Janson, C.H. |
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Social correlates of individual spatial choice in foraging groups of brown capuchin monkeys, Cebus apella |
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Journal Article |
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1990 |
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Animal Behaviour. |
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Anim. Behav. |
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40 |
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5 |
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910-921 |
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Individuals in a foraging group of wild bronwn capuchin monkeys choose different spatial positions relative to the rest of the group. Markov analysis of sequencess of individual spatial positions demonstrated significant differnces between individuals, which coul be categorized a posteriori into four homogenous subgroups. An individual's spatial position was related primarily to the amount of aggression it received from the group's dominant male, but also varied with its sex. Spatial choice varied with changes in an individual's social status, but did not vary consistently with seasonal differences in food availability. These results support the hypothesis that individuals compete for preferred spatial positions within a foraging group. |
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773 |
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Janson, C.H. |
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Title |
Ecological consequences of individual spatial choice in foraging groups of brown capuchin monkeys, Cebus apella |
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Journal Article |
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Year |
1990 |
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Animal Behaviour. |
Abbreviated Journal |
Anim. Behav. |
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40 |
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5 |
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922-934 |
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Individuals in a foraging group of brown capuchin monkeys choose different spatial positions relative to the rest of the group. An individual's choice of spatial positiion affects its foraging success and perceived predation risk (as measured by vigilance behaviour). The two most dominant group members preferred to forage where their expected forwaging success was greatest. Juveniles chose to forage where their perceived predation risk was least, not where they would achieve the highest foraging success. The positions used by non-dominant adults neither maximized foraging success nor minimized predation risk. It is likely that subordinate adults accept spatial positions with suboptimal ecological consequences to avoid the costs of frequent confrontations with the dominant members of the group over foraging sites in poreferred positions. |
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refbase @ user @ |
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774 |
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Author |
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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Journal Article |
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Year |
1988 |
Publication |
Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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21 |
Issue |
1-2 |
Pages |
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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Christensen, J.W.; Zharkikh, T.; Ladewig, J.; Yasinetskaya, N. |
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Title |
Social behaviour in stallion groups (Equus przewalskii and Equus caballus) kept under natural and domestic conditions |
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Journal Article |
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2002 |
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Applied Animal Behaviour Science |
Abbreviated Journal |
Appl. Anim. Behav. Sci. |
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76 |
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1 |
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11-20 |
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Domestic horse; Przewalski horse; Stallion group; Social behaviour; Equus caballus; Equus przewalskii |
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The aim of this study was to investigate social behaviour in differently reared stallions in their respective environments; one group of stallions was reared under typical domestic conditions whereas the other group was reared and lives under natural conditions. The domestic group consisted of 19, 2-year-old stallions (Equus caballus), which were all weaned at 4 months of age and experienced either individual or group housing facilities before being pastured with the other similarly aged stallions. The natural living and mixed age group of Przewalski stallions (E. przewalskii) consisted of 13 stallions, most of which were juveniles (n=11, <=4 years; n=2, >9 years). The domestic group was studied in a 4-ha enclosure at the Danish Institute of Agricultural Sciences and the Przewalski group under free-ranging conditions in a 75-ha enclosure in the Askania Nova Biosphere Reserve, Ukraine. Behavioural data was collected during 168 h of direct observation. The occurrence of 14 types of social interactions was recorded and group spacing behaviour was studied using nearest neighbour recordings. In spite of very different environments, reflecting domestic and natural rearing conditions, many similarities in behaviour was found. Play and play fight behaviour was very similar in the two stallion groups. Quantitative differences were found in social grooming since Przewalski stallions groomed more frequently (P=0.004), and in investigative behaviours, since domestic stallions showed more nasal (P=0.005) and body sniffing (P<0.001), whereas Przewalski stallions directed more sniffing towards the genital region (P<0.001). These differences may, however, be attributed to environmental factors and in the period of time the stallions were together prior to the study period. Quantitative differences appeared in some agonistic behaviours (kick threat, P<0.001; and kick, P<0.001), but data do not support earlier findings of Przewalski horses being significantly more aggressive than domestic horses. In general, Przewalski stallions engaged in more social interactions, and they showed less group spacing, i.e. maintained a significantly shorter distance between neighbours (P<0.001). The study indicates that also domestic horses, which have been reared under typical domestic conditions and allowed a period on pasture, show social behaviour, which is very similar to that shown by their non-domestic relatives. |
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refbase @ user @ |
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776 |
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Bahloul, K.; Pereladova, O.B.; Soldatova, N.; Fisenko, G.; Sidorenko, E.; Sempere, A.J. |
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Social organization and dispersion of introduced kulans (Equus hemionus kulan) and Przewalski horses (Equus przewalski) in the Bukhara Reserve, Uzbekistan |
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Journal Article |
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2001 |
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Journal of Arid Environments |
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J. Arid. Environ. |
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47 |
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3 |
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309-323 |
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Przewalski horses; kulans; Central Asia; home range; behaviour |
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Asiatic wild asses and Przewalski horses initially inhabited steppe, semi-desert and desert areas, but Przewalski horses became extinct in the wild, and kulans disappeared at the beginning of the 20th century, except for a small population in Turkmenistan. The Bukhara Breeding Centre (Uzbekistan) was created in 1976 for reintroduction and conservation of wild ungulate species. In 1977-1978, five kulans (two males and three females), from Barsa-Kelmes island on the Aral sea, were introduced into the reserve. The group increased to 25-30 animals in 1989-1990, when eight Przewalski horses from Moscow and St Petersburg zoos were introduced. We analysed the home ranges, preferred habitats and social interactions of these closely related species during 1995-1998 by seasonal and group composition. Horses and asses formed a reproductive group and a secondary non-reproductive group. The home range of the secondary group was larger than the reproductive group and seemed to be less dependent from the watering places. Przewalski horses were less adapted to semi-desert conditions (both water and vegetation needs) than kulan. |
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777 |
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Keiper, R.; Receveur, H. |
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Social interactions of free-ranging Przewalski horses in semi-reserves in the Netherlands |
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1992 |
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Applied Animal Behaviour Science |
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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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Journal Article |
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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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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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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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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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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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