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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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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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Leblanc, M. - A., & Duncan, P. (2007). Can studies of cognitive abilities and of life in the wild really help us to understand equine learning? Behav. Process., 76, 49–52.
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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. (1985). Time-budgets of Camargue horses III. Environmental influences. Behaviour, 92, 188–208.
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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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Mayes, E., & Duncan, P. (1986). Temporal patterns of feeding behaviour in free-ranging horses. Behav., 96, 105–129.
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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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Fleurance, G., Duncan, P., Fritz, H., Cabaret, J., Cortet, J., & Gordon, I. J. (2007). Selection of feeding sites by horses at pasture: Testing the anti-parasite theory. Appl. Anim. Behav. Sci., 108(3-4), 228–301.
Abstract: Management of grazed grasslands for production and/or conservation objectives requires a thorough understanding of the choices of feeding sites by herbivores, and of the biological processes involved. Most models of the feeding strategies of herbivores are based on the principle that optimising the intake of energy (or some nutrient) is the primary goal of foragers but other selective forces, such as parasitism, could be important. Gastrointestinal parasites (including cyathostome nematodes) have powerful effects on the fitness of herbivores and may act as a major selection pressure favouring host behaviour that reduces the risk of encountering parasites. Among large herbivores, horses have perhaps the most marked tendency to select particular feeding sites within grasslands. We test here: (1) whether horses select feeding patches with relatively low parasite densities and (2) if their choice is affected by their parasite load. We used 10 two-year old saddle-horses and three periods. In the first period, the horses were under natural parasitism which varied strongly among individuals; in the second period they were all dewormed, and in the third, a sub-set of the horses was experimentally infected with cyathostome larvae. Ninety-eight percent of the infective larvae in the pasture were found <1 m from faeces. The main determinant of the choice of feeding patch by horses was the availability of patches of different parasite risk and grass height. Controlling for availability, the horses used tall grasses (>16 cm) less than expected, whether the grass was contaminated or not, and they selected for short patches >1 m from faeces, where the risk of encountering parasites was low. These results suggest that selection of feeding sites by horses is driven by an interaction between their nutritional and anti-parasite strategies: the horses avoid the patches of tall grass which are generally of low quality and areas contaminated by parasite larvae which leads them to prefer the patches of short grass far from faeces. The parasite status of the horses at the time of the experiment had no effect on their feeding choices. However, before concluding that the challenge by cyathostomes has no effect on the selection of feeding sites in horses, it will be necessary to test whether the history of parasitism of the individuals, rather than the current status, is important.
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Edouard, N., Fleurance, G., Dumont, B., Baumont, R., & Duncan, P. (2009). Does sward height affect feeding patch choice and voluntary intake in horses? Appl. Anim. Behav. Sci., 119(3-4), 219–228.
Abstract: The numbers of horses grazing at pasture are increasing in developed countries, so a proper understanding of their feeding selectivity and of the tactics they use for extracting nutrients from swards is essential for the management of horses and grasslands. Resource acquisition in herbivores can be optimised through the modulation of their intake and patch selection, both being strongly dependent on the characteristics of swards. However, the principles by which horses adjust their grazing behaviour in response to variations in sward features are not completely understood. The aim of this study was to determine whether the behaviour of horses conforms to optimal foraging models. We hypothesized that, faced with binary choices between vegetative swards of a good and similar quality at three different heights, horses would select the taller sward, i.e. that allowing a higher reward in terms of dry matter intake rate. Three groups of three 2-year-old saddle horses were grazed on a semi-natural pasture that was managed to produce three contrasting sward heights at 6, 11 and 17 cm, in a Latin-square design. The instantaneous intake rate was determined from bite rate measured at pasture on the three sward heights, and bite mass estimated from measurements using swards offered indoors in experimental trays. Daily dry matter intake was estimated individually by total faecal collection and an estimation of digestibility from faecal nitrogen. Short-term (first 30 min) and daily preferences were assessed from the time spent grazing each sward offered in pair-wise tests at pasture. The results show that daily voluntary intake (an average of 21 g DM kg LW-1 day-1) and total grazing time (an average of 14 h day-1) were independent of sward height and of the choice of patches offered. In choice situations, the animals spent more time grazing on the taller sward, both during the first 30 min and at the daily scale. These results show that horses choose between vegetative patches of a good and similar quality according to the predictions from optimal foraging models, and select the one that they can ingest faster. Further research will now have to explore how the horses will adapt their feeding behaviour when they face a trade-off between sward height and quality.
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