Pictures of the grassland habitat around the Kabini backwaters, foraging elephant groups, and agonistic interactions;Details of focal zone sampling for grass abundance, and density of elephants in the grassland habitat;Details of within-clan and between-clan agonism: sampling, video scoring, and data analysis;Grass biomass in the grassland habitat in Kabini and in the neighbouring forest/savannah woodland habitats of Nagarahole and Bandipur National Parks, and within-zone variability in grass biomass, cover, and grass height in the grassland habitat;Comparison of agonism between females within and between clans: behaviours shown and rates.

According to the ecological model of female social relationships (EMFSR), within-group competition and between-group competition in female-bonded species are shaped by food distribution. Strong between-group contests are expected over large, monopolizable resources and high population density, but not when low-quality food is distributed across large, undefended home ranges. Within-group contests are expected to be more frequent with increasing heterogeneity among feeding sites and with group size. We tested these predictions in female Asian elephants, which show traits associated with infrequent contests –graminivory, high fission–fusion and overlapping home ranges. We examined how food distribution and competitor density affected agonistic interactions within and between female elephant clans (social groupings) in the Kabini grassland, southern India. We found stronger between-clan contest in the grassland than that known from neighbouring forests, and more frequent agonism between females between clans than within clans. Such strong between-clan contest was attributable to the grassland being a food-rich habitat patch, thus supporting EMFSR. Within-clan agonism was also frequent, but did not increase with food heterogeneity, contradicting EMFSR. Contrary to recent claims, increasing within-clan agonism with group size suggested ecological constraints on large groups despite high fission–fusion. High population density may explain such frequent contests despite graminivory and fission–fusion.