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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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Sundaresan, S. R., Fischhoff, I. R., Dushoff, J., & Rubenstein, D. I. (2007). Network metrics reveal differences in social organization between two fission-fusion species, Grevy's zebra and onager. Oecologia, 151(1), 140–149.
Abstract: For species in which group membership frequently changes, it has been a challenge to characterize variation in individual interactions and social structure. Quantifying this variation is necessary to test hypotheses about ecological determinants of social patterns and to make predictions about how group dynamics affect the development of cooperative relationships and transmission processes. Network models have recently become popular for analyzing individual contacts within a population context. We use network metrics to compare populations of Grevy's zebra (Equus grevyi) and onagers (Equus hemionus khur). These closely related equids, previously described as having the same social system, inhabit environments differing in the distribution of food, water, and predators. Grevy's zebra and onagers are one example of many sets of coarsely similar fission-fusion species and populations, observed elsewhere in other ungulates, primates, and cetaceans. Our analysis of the population association networks reveals contrasts consistent with their distinctive environments. Grevy's zebra individuals are more selective in their association choices. Grevy's zebra form stable cliques, while onager associations are more fluid. We find evidence that females associate assortatively by reproductive state in Grevy's zebra but not in onagers. The current approach demonstrates the utility of network metrics for identifying fine-grained variation among individuals and populations in association patterns. From our analysis, we can make testable predictions about behavioral mechanisms underlying social structure and its effects on transmission processes.
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Gauvin, S., & Giraldeau, L. - A. (2004). Nutmeg mannikins ( Lonchura punctulata) reduce their feeding rates in response to simulated competition. Oecologia, 139(1), 150–156.
Abstract: Group feeding animals experience a number of competitive foraging costs that may result in a lowered feeding rate. It is important to distinguish between reductions in feeding rates that are caused by reduced food availability and physical interactions among foragers from those caused by the mere presence of foraging companions that may be self-imposed in order to obtain some benefit of group membership. Starlings ( Sturnus vulgaris) reduce their feeding rates when in the company of simulated competitors located in an adjacent cage that cannot affect the food availability or interact with the forager. In the present study, we investigate whether the presence of simulated competitors in another species of passerine, nutmeg mannikins ( Lonchura punctulata), can result in self-imposed reductions in feeding rates. When feeding in the company of simulated competitors, mannikins spent more non-foraging time near them, fed more slowly, reduced travel times between patches, reduced their scanning time and pecked more slowly. These results provide evidence that simulated competitors induce a reduction in pecking rate: behavioural interference. These self-imposed responses to competitors may have resulted from attempts to remain close to the non-feeding companions. Such self-imposed reductions in feeding rates may be a widespread yet generally unrecognised foraging cost to group feeding individuals.
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