How to find home backwards? Navigation during rearward homing of Cataglyphis fortis desert ants.

Cataglyphis ants are renowned for their impressive navigation skills, which have been studied in numerous experiments during forward locomotion. However, the ants' navigational performance during backward homing when dragging large food loads has not been investigated until now. During backward locomotion, the odometer has to deal with unsteady motion and irregularities in inter-leg coordination. The legs' sensory feedback during backward walking is not just a simple reversal of the forward stepping movements: compared with forward homing, ants are facing towards the opposite direction during backward dragging. Hence, the compass system has to cope with a flipped celestial view (in terms of the polarization pattern and the position of the sun) and an inverted retinotopic image of the visual panorama and landmark environment. The same is true for wind and olfactory cues. In this study we analyze for the first time backward-homing ants and evaluate their navigational performance in channel and open field experiments. Backward-homing Cataglyphis fortis desert ants show remarkable similarities in the performance of homing compared with forward-walking ants. Despite the numerous challenges emerging for the navigational system during backward walking, we show that ants perform quite well in our experiments. Direction and distance gauging was comparable to that of the forward-walking control groups. Interestingly, we found that backward-homing ants often put down the food item and performed foodless search loops around the left food item. These search loops were mainly centred around the drop-off position (and not around the nest position), and increased in length the closer the ants came to their fictive nest site.