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Feh, C. (1987). Etude du développement des relations sociales chez des étalons (Equus caballus) de race camargue et de leur contribution à l'organisation sociale du groupe. Master's thesis, Université d'Aix-Marseille, Marseille.
Abstract: Diplôme de recherche universitaire: neurosciences comportementales: Université d'Aix-Marseille II. Faculté des sciences de Luminy Marseille (FRA)* 117p
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Feh, C. (1988). Social behaviour and relationships of Prezewalski horses in Dutch semi-reserves. Appl. Anim. Behav. Sci., 21(1-2), 71–87.
Abstract: 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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Feh, C., & de Mazières, J. (1993). Grooming at a preferred site reduces heart rate in horses. Anim. Behav., 46(6), 1191–1194.
Abstract: Abstract. It is commonly suggested that the principal function of allogrooming is to reduce social tension between group members, but direct evidence of the physiological consequences of grooming at particular sites is lacking. By filming allogrooming sequences in a herd of Camargue horses, Equus caballus , their preferred grooming site, which lies on the lower neck, was identified. Experimental imitation of grooming at this site reduced the heart rate of the recipient while grooming on a non-preferred area did not, in both adults and foals. This preferred site lies close to a major ganglion of the autonomic nervous system.
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Feh, C., Enchbold, S., & Munchtuya, B. (1996). Preliminary assessment of the Gurvan Saikhan National Conservation Park's potential for Gobi khulan (Equus hemionus luteus). GTZ, .
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Monard, A. - M., Duncan, P., Fritz, H., & Feh, C. (1997). Variations in the birth sex ratio and neonatal mortality in a natural herd of horses. Behav. Ecol. Sociobiol., 41(4), 243–249.
Abstract: Variations in birth sex ratios and sex differences in juvenile mortality occur in a number of mammalian species, and in many cases have been linked to resource availability. Most of these biases in offspring sex ratios concern polygynous species with pronounced sexual dimorphism, and where females only are philopatric. Data on species with unusual life-history strategies, such as slight sexual dimorphism or dispersal by both sexes, are of particular interest. In this study of a natural herd of horses (Equus caballus) which experienced an eruptive cycle, and therefore a period of nutritional stress, male offspring had higher neonatal mortality rates in nutritionally poor years than in good ones, whereas “year quality” had no effect on the mortality of female offspring; year quality could therefore be used by mares as predictor of sex-specific offspring survival. We show that the environmental conditions that predicted lower survival of males were negatively related to their production: the birth sex ratio the following year was female-biased; and mares were less likely to produce a son when they had produced a son the preceding year. There was no significant effect of mother's parity, age or rank, or the timing of conception or birth on offspring sex ratios. The mechanism leading to biases in the birth sex ratio could have been the loss of male embryos by mares that did not foal. As there was no evidence for selective abortion of male foetuses in females that did foal the next year, it is not necessary to invoke maternal adjustment, though this remains a possibility. Finally, there was a suggestion that male offspring were more costly to raise than females, since mothers that reared a son in poor years tended to experience an increase in the interbirth interval between their two subsequent offspring.
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Feh, C. (1999). Alliances and reproductive success in Camargue stallions. Anim. Behav., 57(3), 705–713.
Abstract: A study of a herd of Camargue horsesEquus caballus, showed that while the majority of high-ranking stallions held single-male harems, some sons of low-ranking mares, being low ranking themselves, formed alliances that could last a lifetime. The two stallions were each other's closest associate and preferential grooming partner. Alliances were based on coalitions in which either both partners confronted an intruder synchronously or the dominant of the pair tended the female(s) while the subordinate simultaneously displayed towards the rival. Alliance partners were of similar age but were not more closely related to each other than to other stallions in the herd. Long-term paternity data revealed that subordinates sired close to a quarter of the foals born into the alliance group, and significantly more foals than low-ranking stallions in the herd adopting a `sneak'-mating strategy. The dominant appeared to benefit from the presence of his subordinate partner. Fights occurred all year round, and the subordinate stallion of each alliance pair fought outside competitors more than twice as often as the dominant. Forming short-term alliances before defending mares on their own may enhance long-term reproductive success for both partners. Other benefits to both partners include higher survivorship of their foals and increased access to proven reproductive mares. These results suggest that the relationship between alliance partners is based on mutualism, but several conditions for reciprocity seem to be fulfilled: the benefit to the dominant (assistance in fights), and the benefit to the subordinate (access to reproduction), are both costly to the other partner and delayed in time.
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Feh, C. (2001). Alliances between stallions are more than just multimale groups: reply to Linklater & Cameron (2000). Anim. Behav., 61, F27–F30.
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Feh, C., Munkhtuya, B., Enkhbold, S., & Sukhbaatar, T. (2001). Ecology and social structure of the Gobi khulan Equus hemionus subsp. in the Gobi B National Park, Mongolia. Biol. Conserva., 101(1), 51–61.
Abstract: The status of the Gobi khulan Equus hemionus subsp. is recorded as “insufficiently known” in the Species Survival Commission's Equid Action Plan. Recent counts confirm that Mongolia holds the most important population of the whole species. Since 1953, the animals have benefited from a protected status, but this is now challenged. A 5-year study in the B part of the Gobi National Park on one subpopulation showed that it has remained stable over the past 15 years with an adequate mean reproductive rate of 15% and a 50% survival rate over the first year. Age/sex related mortality and prey analysis indicate that wolf predation probably has some impact on the population, in particular for 4-6-year-olds of both sexes at the start of reproduction. Desert and mountain steppes are the khulan's year-round preferred habitat, but `oases', play an important role at the beginning of lactation. Anthropogenic factors affect both home range and habitat use through direct intervention or permanent occupation of the scarce water sources. Khulans of this subpopulation, unlike other Asian and African wild asses, form year-round stable, non-territorial families. These families and all-male groups join together into “bands” in winter, and herds of several hundred animals, where reproductive rate is highest, form throughout the year. The existence of such herds may thus be critical for the breeding success of the population. Our study provides the first detailed quantitative data for this subspecies, which will help to monitor changes in the future.
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Feh, C. (2005). Relationships and Communication in Socially Natural Horse Herds. In D. S. Mills, & S. M. McDonnell (Eds.), The domestic horse : the origins, development, and management of its behaviour. Cambridge: Cambridge University Press 2005.
Abstract: Horses are quite unique. In most mammals, sexes segregate and maintain bonds only during the breeding season (Clutton-Brock, 1989). Some canids, a few rodents and primate species such as gorillas, hamadryas baboons and red howler monkeys are the exception, where the same males stay with the same females all year round and over many breeding seasons. Typically, both sexes disperse at puberty in these species. In horses, it was clearly shown that the causes for female dispersal were incest avoidance and not intra-specific competition (Monard, 1996). As a rule, this is confirmed for mammal species where tenure length by males exceeds the age at first reproduction in females (Clutton-Brock, 1989). When horses are allowed to choose their mating partner freely, the inbreeding coefficient of the offspring is lower than expected should they mate randomly (Duncan et al, 1984).
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Feh, C., & Munkhtuya, B. (2008). Male infanticide and paternity analyses in a socially natural herd of Przewalski`s horses: Sexual selection? Behav. Process., 78(3), 335–339.
Abstract: The sexual selection hypothesis explains infanticide by males in many mammals. In our 11-year study, we investigated this hypothesis in a herd of Przewalski's horses where we had witnessed infanticidal attacks. Infanticide was highly conditional and not simply linked to takeovers. Attacks occurred in only five of 39 cases following a takeover, and DNA paternity revealed that, although infanticidal stallions were not the genetic fathers in four cases out of five, stallions present at birth did not significantly attempt to kill unrelated foals. Infanticide did not reduce birth intervals; only in one case out of five was the infanticidal stallion, the father of the next foal; mothers whose foals were attacked subsequently avoided associating with infanticidal stallions. Therefore, evidence for the sexual selection hypothesis was weak. The “human disturbance” hypothesis received some support, as only zoo bred stallions which grew up in unnatural social groups attacked foals of mares which were pregnant during takeovers.
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