Exploring behavior of an unusual megaherbivore: a spatially explicit foraging model of the hippopotamus

Herbivore foraging theories have been developed for and tested on herbivores across a range of sizes. Due to logistical constraints, however, little research has focused on foraging behavior of megaherbivores. Here we present a research approach that explores megaherbivore foraging behavior, and assesses the applicability of foraging theories developed on smaller herbivores to megafauna. With simulation models as reference points for the analysis of empirical data, we investigate foraging strategies of the common hippopotamus (Hippopotamus amphibius). Using a spatially explicit individual based foraging model, we apply traditional herbivore foraging strategies to a model hippopotamus, compare model output, and then relate these results to field data from wild hippopotami. Hippopotami appear to employ foraging strategies that respond to vegetation characteristics, such as vegetation quality, as well as spatial reference information, namely distance to a water source. Model predictions, field observations, and comparisons of the two support that hippopotami generally conform to the central place foraging construct. These analyses point to the applicability of general herbivore foraging concepts to megaherbivores, but also point to important differences between hippopotami and other herbivores. Our synergistic approach of models as reference points for empirical data highlights a useful method of behavioral analysis for hard-to-study megafauna.