Dry season diets of sympatric ungulates in lowland Nepal: competition and facilitation in alluvial tall grasslands

Based on microhistological analyses of faecal material, we compared the early dry season diets of greater one-horned rhinoceros Rhinoceros unicornis, swamp deer Cervus duvauceli and hog deer Axis porcinus, which inhabit the same alluvial grassland habitat complex in lowland Nepal. Their diets were quite similar, both at the forage category level and within subcategories of graminoids and woody plants. Early successional tall grasses, especially Saccharum spontaneum, were the dominant food of all three species, underlining the key role of the threatened alluvial floodplains in large mammal conservation in South Asia. The two deer species ate significantly more graminoids (>66.5%) than did rhino (45.5%), and although they did not differ in proportions of graminoids, swamp deer ate significantly more late successional tall grasses (Narenga porphyrocoma and Themeda spp.) and short grasses (mainly Imperata cylindrica) than hog deer. The two deer consumed almost equal proportions of woody browse (ca. 10%), significantly less than that of rhino (33.0%). The prediction of the Jarman¿Bell hypothesis, that large-bodied herbivores are less selective and subsist on lower quality graminoids than smaller ruminants, was not supported by the data. Based on this and previous studies in the same area we propose a conceptual model where the larger megaherbivores (rhino and elephant Elephas maximus) facilitate the smaller swamp deer and hog deer during the monsoonal growing season, while the smaller and more selective deer species outcompete the larger during the dry season when food is more limited. Owing to the all-year sprouting ability of S. spontaneum, facilitation may occur also in the dry season with low numbers of megaherbivores, thus accentuating competitive exclusion at higher deer densities