Gates, S. (1979). A Study of home ranges of free ranging Exmoore ponies. Mamm. Rev., 9, :3–18.
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Ginsberg, J. R. (1988). Social organisation and mating strategies of an arid adapted equid: The Grevy`s zebra. Ph.D. thesis, Princeton University, Princeton.
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Sayigh, L. S., Tyack, P. L., Wells, R. S., Solow, A. R., Scott, M. D., & Irvine, A. B. (1999). Individual recognition in wild bottlenose dolphins: a field test using playback experiments. Anim. Behav., 57(1), 41–50.
Abstract: We conducted playback experiments with wild bottlenose dolphins, Tursiops truncatus, to determine whether there is sufficient information in their individually distinctive signature whistles for individual recognition. We conducted experiments with members of a resident community of dolphins in waters near Sarasota, Florida, during temporary capture-release projects. We used a paired playback design, wherein the same two whistle sequences were predicted to evoke opposite responses from two different target animals. This design controlled for any unknown cues that may have been present in the playback stimuli. We predicted that mothers would respond more strongly to the whistles of their own independent offspring than to the whistles of a familiar, similar-aged nonoffspring. Similarly, we predicted that independent offspring would respond more strongly to the whistles of their own mother than to the whistles of a familiar, similar-aged female. Target animals were significantly (P<0.02) more likely to respond to the predicted stimuli, with responses measured by the number of head turns towards the playback speaker. In bottlenose dolphin societies, stable, individual-specific relationships are intermixed with fluid patterns of association between individuals. In primate species that live in similar 'fission-fusion' type societies, individual recognition is commonplace. Thus, when taken in the context of what is known about the social structure and behaviour of bottlenose dolphins, these playback experiments suggest that signature whistles are used for individual recognition. Copyright 1999 The Association for the Study of Animal Behaviour.
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Hamilton, W. D. (1971). Geometry for the selfish herd. J. Theor. Biol., 31(2), 295–311.
Abstract: 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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Jansen, T., Forster, P., Levine, M. A., Oelke, H., Hurles, M., Renfrew, C., et al. (2002). Mitochondrial DNA and the origins of the domestic horse. Proc. Natl. Acad. Sci. U.S.A., 99(16), 10905–10910.
Abstract: 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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Janson, C. H. (1990). Social correlates of individual spatial choice in foraging groups of brown capuchin monkeys, Cebus apella. Anim. Behav., 40(5), 910–921.
Abstract: 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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Janson, C. H. (1990). Ecological consequences of individual spatial choice in foraging groups of brown capuchin monkeys, Cebus apella. Anim. Behav., 40(5), 922–934.
Abstract: 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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Keiper, R. R. (1988). Social interactions of the Przewalski horse (Equus przewalskii Poliakov, 1881) herd at the Munich Zoo. Appl. Anim. Behav. Sci., 21(1-2), 89–97.
Abstract: 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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Christensen, J. W., Zharkikh, T., Ladewig, J., & Yasinetskaya, N. (2002). Social behaviour in stallion groups (Equus przewalskii and Equus caballus) kept under natural and domestic conditions. Appl. Anim. Behav. Sci., 76(1), 11–20.
Abstract: 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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Bahloul, K., Pereladova, O. B., Soldatova, N., Fisenko, G., Sidorenko, E., & Sempere, A. J. (2001). Social organization and dispersion of introduced kulans (Equus hemionus kulan) and Przewalski horses (Equus przewalski) in the Bukhara Reserve, Uzbekistan. J. Arid. Environ., 47(3), 309–323.
Abstract: 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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