Fruit, minerals, and forest elephant trails: Do all roads lead to Rome?

Tropical forests are among the most heterogeneous environments on earth, and food resources for many animals are patchy both in time and space. In Africa's equatorial forest, permanent trails created and maintained by forest elephants are conspicuous features. Trails may be several meters wide and continue for tens of kilometers. Speculation on which resources determine the distribution of trails has identified fruit, browse, and mineral deposits as candidates. In this study, the relationships between these habitat variables and elephant trails were investigated. The size of individual trails and the density of the trail system increased dramatically with proximity to mineral deposits. Fruit tree basal area decreased with perpendicular distance from trails, while that of non-fruit trees did not. Fruit tree abundance and basal area were significantly higher on trail intersections than random sites and increased with intersection size. No relationship was found between monocotyledon browse abundance and elephant trail system characteristics. Clumped resources, which are at least partially reliable, provide a high nutritional payback, and are not rapidly depleted and can thus be visited repeatedly, appear to influence permanent trail formation by forest elephants. Permanent trails may allow naive individuals or those with imperfect knowledge to locate and acquire important resources.