Flexible locomotion in complex environments: the influence of species, speed and sensory feedback on panarthropod inter-leg coordination

Panarthropods (a clade containing arthropods, tardigrades and onychophorans) can adeptly move across a wide range of challenging terrains and their ability to do so given their relatively simple nervous systems makes them compelling study organisms. Studies of forward walking on flat terrain excitingly point to key features in inter-leg coordination patterns that seem to be 'universally' shared across panarthropods. However, when movement through more complex, naturalistic terrain is considered, variability in coordination patterns - from the intra-individual to inter-species level - becomes more apparent. This variability is likely to be due to the interplay between sensory feedback and local pattern-generating activity, and depends crucially on species, walking speed and behavioral goal. Here, I gather data from the literature of panarthropod walking coordination on both flat ground and across more complex terrain. This Review aims to emphasize the value of: (1) designing experiments with an eye towards studying organisms in natural environments; (2) thoughtfully integrating results from various experimental techniques, such as neurophysiological and biomechanical studies; and (3) ensuring that data is collected and made available from a wider range of species for future comparative analyses.