Universal features in panarthropod inter-limb coordination during forward walking

Abstract

Terrestrial animals must often negotiate heterogeneous, varying environments. Accordingly, their locomotive strategies must adapt to a wide range of terrain, as well as to a range of speeds in order to accomplish different behavioral goals. Studies in \textit{Drosophila} have found that inter-leg coordination patterns (ICPs) vary smoothly with walking speed, rather than switching between distinct gaits as in vertebrates (e.g., horses transitioning between trotting and galloping). Such a continuum of stepping patterns implies that separate neural controllers are not necessary for each observed ICP. Furthermore, the spectrum of \textit{Drosophila} stepping patterns includes all canonical coordination patterns observed during forward walking in insects. This raises the exciting possibility that the controller in \textit{Drosophila} is common to all insects, and perhaps more generally to panarthropod walkers. Here, we survey and collate data on leg kinematics and inter-leg coordination relationships during forward walking in a range of arthropod species, as well as include data from a recent behavioral investigation into the tardigrade \textit{Hypsibius exemplaris}. Using this comparative dataset, we point to several functional and morphological features that are shared amongst panarthropods. The goal of the framework presented in this review is to emphasize the importance of comparative functional and morphological analyses in understanding the origins and diversification of walking in Panarthropoda.