Should elephants graze or browse? The nutritional and functional consequences of dietary variation in a mixed-feeding megaherbivore

Abstract

Unlike specialist browsers and grazers, the diets of mixed-feeding megaherbivores like elephants are complex and broad, comprising numerous plant species of variable nutritional quality. Here, we revisit an unresolved debate on whether browse is more nutritious than grasses for elephants, as browse is thought to have greater crude protein content. We first analyzed carbon isotopes in 102 fecal samples of Asian elephants to quantify the contribution of browsing and grazing to their diet in Nagarahole National Park, southern India. We show that elephants are predominantly browsers in the grass-scarce Nagarahole forests, in contrast with predominant grazing reported in the nearby grass-abundant savannas of the Nilgiri Biosphere Reserve. We then compared the forage quality of high-browsing and high-grazing diets by analyzing these samples for two proxies of crude protein content (CP): nitrogen content (N%) and Carbon-to-Nitrogen ratio (C:N). Interestingly, high-browsing diets did not have higher N% or lower C:N (proxies for high CP) than low-browsing (high-grazing) diets. To explore the generality of this finding, we analyzed nutritional differences (i.e., CP and fibre content values) between browse and grass across 141 plant species consumed by Asian elephants. We show that woody browse and non-legume browse, which are major components of elephant browse, do not have appreciably higher CP than grasses. Our findings suggest that browsing and grazing broadly have similar nutritional value for such bulk-feeding mixed feeders. Finally, based on the observed habitat-wide variation in browsing, we provide a new framework for assessing how elephants shape woody vegetation across the forests and savannas of Asia, with important implications for conservation and carbon cycling.