Locomotion in the pseudoscorpion Chelifer cancroides: forward, backward and upside-down walking in an eight-legged arthropod.

While insect locomotion has been intensively studied, there are comparably few studies investigating octopedal walking behaviour, and very little is known about pseudoscorpions in particular. Therefore, we performed an extensive locomotion analysis during forward, backward and upside-down walking in the cosmopolitan pseudoscorpion Chelifer cancroides. During forward locomotion, we observed C. cancroides to freeze locomotion frequently for short time periods. These microstops were barely visible to the naked eye with a duration of 100-200 ms. Our locomotion analysis revealed that C. cancroides performs a statically stable and highly coordinated alternating tetrapod gait during forward and backward walking, with almost complete inversion of the tetrapod schemes, but no rigidly fixed leg coordination during upside-down walks with low walking speeds up to 4 body lengths per second. Highest speeds (up to 17 body lengths per second), mainly achieved by consistent leg coordination and strong phase shifts, were observed during backward locomotion (escape behaviour), whereas forward walking was characterised by lower speeds and phase shifts of ∼10% between two loosely coupled leg groups within one tetrapod. That is, during the movement of one tetrapod group, the last and the third leg are almost synchronous in their swing phases, as are the second and the first leg. A special role of the second leg pair was demonstrated, probably mainly for stability reasons and related to the large pedipalps.