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Duncan, P., & Vigne, N. (1979). The effect of group size in horses on the rate of attacks by blood-sucking flies. Anim. Behav., 27(Part 2), 623–625.
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Boy, V., & Duncan, P. (1979). Time-budgets of Camargue horses. I. Developmental changes in the time-budgets of foals. Behaviour, 71, 187–201.
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Duncan, P. (1982). Foal killing by stallions. Appl. Animal. Ethol., 8(6), 567–570.
Abstract: Feral horses live in social systems similar to those of some species in which infant killing has been reported (e.g. lions), but such behaviour has been reported neither in horses nor in any other ungulate. The results of interviews with owners of free-ranging horses (Camargue breed) are given which show that, though rare, infant killing occurs in this breed, and that it seems to be confined to male foals. It is argued that the observed behaviour cannot simply be considered as pathological, and that close attention should be paid to the possibility that it occurs in wild and feral equids.
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Duncan, P. (1983). Determinants of the use of habitat by horses in a mediterranean wetland. J. Anim. Ecol., 52, 93–109.
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Duncan, P. (1985). Time-budgets of Camargue horses III. Environmental influences. Behaviour, 92, 188–208.
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Mayes, E., & Duncan, P. (1986). Temporal patterns of feeding behaviour in free-ranging horses. Behav., 96, 105–129.
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Duncan, P., Foose, T. J., Gordon, I. J., Gakahu, C. G., & Lloyd, M. (1990). Comparative nutrient extraction from forages by grazing bovids and equids: a test of the nutritional model of equid/bovid competition and coexistence. Oecologia, 84(3), 411–418.
Abstract: Ruminants are unevenly distributed across the range of body sizes observed in herbivorous mammals; among extant East African species they predominate, in numbers and species richness, in the medium body sizes (10-600 kg). The small and the large species are all hind-gut fermenters. Some medium-sized hind-gut fermenters, equid perissodactyls, coexist with the grazing ruminants, principally bovid artiodactyls, in grassland ecosystems. These patterns have been explained by two complementary models based on differences between the digestive physiology of ruminants and hind-gut fermenters. The Demment and Van Soest (1985) model accounts for the absence of ruminants among the small and large species, while the Bell/Janis/Foose model accounts both for the predominance of ruminants, and their co-existence with equids among the medium-sized species (Bell 1971; Janis 1976; Foose 1982). The latter model assumes that the rumen is competitively superior to the hind-gut system on medium quality forages, and that hind-gut fermenters persist because of their ability to eat more, and thus to extract more nutrients per day from high fibre, low quality forages. Data presented here demonstrate that compared to similarly sized grazing ruminants (bovids), hind-gut fermenters (equids) have higher rates of food intake which more than compensate for their lesser ability to digest plant material. As a consequence equids extract more nutrients per day than bovids not only from low quality foods, but from the whole range of forages eaten by animals of this size. Neither of the current nutritional models, nor refinements of them satisfactorily explain the preponderance of the bovids among medium-sized ungulates; alternative hypotheses are presented.
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Monard, A. - M., & Duncan, P. (1996). Consequences of natal dispersal in female horses. Anim. Behav., 52(3), 565–579.
Abstract: Social, genetic and reproductive consequences of natal dispersal were investigated in female horses,Equus caballus, living in a herd with a natural social structure. Dispersal did not as a rule reduce the level of competition the young mares faced: they did not selectively join groups with fewer resident females than the groups they left, and they did not attain higher ranks; there was also no tendency for females to disperse to groups with the fewest resident females, and they suffered more aggression from the mares in their new groups than in their natal groups. These results therefore do not support the hypothesis that a function of natal dispersal is to reduce intra-sexual competition. The young mares nevertheless dispersed non-randomly, generally joining harems with one stallion and at least two subadult females; and they preferred to move to groups with familiar females but no familiar males. As a result, most were closely related to some females of their new groups, but distantly related or unrelated to the male(s). Since after dispersal the young mares bred only with a male of their new groups, inbreeding coefficients of most (85%) of their offspring were lower than from matings between half siblings. These results are consistent with the hypothesis that a function of natal dispersal is to avoid close inbreeding. Dispersal did not appear to involve reproductive costs: the young mares suffered no delay in age at first reproduction, and the survival rates of their first foals tended to be higher if the females had emigrated, although not significantly so.
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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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Grange, S., & Duncan, P. (2006). Bottom-up and top-down processes in African ungulate communities: resources and predation acting on the relative abundance of zebra and grazing bovids. Ecography, 29(6), 899–907.
Abstract: African ungulate populations appear to be limited principally by their food resources. Within ungulate communities, plains zebras coexist with grazing bovids of similar body size, but rarely are the dominant species. Given the highly effective nutritional strategy of the equids and the resistance of zebras to drought, this is unexpected and suggests that zebra populations may commonly be limited by other mechanisms. Long-term research in the Serengeti ecosystem and in the Kruger National Park suggests that zebra could be less sensitive to food shortage, and more sensitive to predation, than grazing bovids: if this is a general principle, then, at a larger scale, resource availability should have a weaker effect on the abundance of zebra than on grazing ruminants of similar body size (wildebeest and buffalo), and zebras should be relatively more abundant in ecosystems where predators are rare or absent. We test these expectations using data on 23 near-natural ecosystems in east and southern Africa. The abundance of wildebeest is more closely related to resources than is that of zebra; buffalo are intermediate. We show that hyena densities are closely correlated with those of lions, and use the abundance of lions as an index of predation by large predators. The numerical response of lions to increases in the abundance of their prey was linear for mesoherbivores, and apparently so for the three species alone. Finally, the abundance of zebra relative to grazing bovids is lower in ecosystems with high biomasses of lions. These results indicate that zebras may commonly be more sensitive to top-down processes than grazing bovids: the mechanism(s) have not been demonstrated, but predation could play a role. If it is true, then when numbers of the large mammalian predators decline, zebra populations should increase faster than buffalo and wildebeest.
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