Interaction between path integration and visual orientation during the homing run of fiddler crabs

Foraging fiddler crabs form a strict spatial relationship between their current positions and burrows, allowing them to run directly back to their burrows when startled even without visual contacts. Path integration (PI), the underlying mechanism, is a universal navigation strategy through which animals continuously integrate directions and distances of their movements. However, we report that fiddler crabs also use visual orientation during homing runs using burrow entrances as cues, with the prioritised mechanism (i.e. PI or visual) determined by the distance (which has a threshold value) between the goal, indicated by PI, and the visual cue. When we imposed homing errors using fake entrances (visual cue) and masking their true burrows (goal of PI), we found that frightened fiddler crabs initially ran towards the true burrow following PI, then altered their behaviour depending on the distance between the fake entrance and masked true burrow: if the distance was large, they kept running until they reached the true burrow, ignoring the visual cue; however, if the distance was small, they altered the homing path and ran until they reached the fake entrance. This suggests that PI and visual mechanism in fiddler crabs are mutually mediated to achieve their homing behaviour.