Fiddler crabs accurately measure two-dimensional distance over three-dimensional terrain

Foraging fiddler crabs (Uca spp.) monitor the location of, and are able to return to, their burrows by employing path integration. This requires them to accurately measure both the directions and distances of their locomotory movements. Even though most fiddler crabs inhabit relatively flat terrain, they must cope with vertical features of their environment, such as sloping beaches, mounds and shells, which may represent significant obstacles. To determine whether fiddler crabs can successfully perform path integration among such three-dimensional obstacles, we tested their ability to measure distance while we imposed a vertical detour. By inserting a large hill in the homeward path of foraging crabs we show that fiddler crabs can cope with vertical detours: they accurately travel the correct horizontal distance, despite the fact that the shape of the hill forces them to change their gait from what would be used on flat ground. Our results demonstrate a flexible path integrator capable of measuring, and either integrating or discarding, the vertical dimension.