Terrestrial locomotion in arachnids

In this review, we assess the current state of knowledge on terrestrial locomotion in Arachnida. Arachnids represent a single diverse (>100,000 species) clade containing well-defined subgroups (at both the order and subordinal levels) that vary morphologically around a basic body plan, yet exhibit highly disparate limb usage, running performance, and tarsal attachment mechanisms. Spiders (Araneae), scorpions (Scorpiones), and harvestmen (Opiliones) have received the most attention in the literature, while some orders have never been subject to rigorous mechanical characterization. Most well-characterized taxa move with gaits analogous to the alternating tripod gaits that characterize fast-moving Insecta - alternating tetrapods or alternating tripods (when one pair of legs is lifted from the ground for some other function). However, between taxa, there is considerable variation in the regularity of phasing between legs. Both large and small spiders appear to show a large amount of variation in the distribution of foot-ground contact, even between consecutive step-cycles of a single run. Mechanisms for attachment to vertical surfaces also vary, and may depend on tufts of adhesive hairs, fluid adhesives, silks, or a combination of these. We conclude that Arachnida, particularly with improvements in microelectronic force sensing technology, can serve as a powerful study system for understanding the kinematics, dynamics, and ecological correlates of sprawled-posture locomotion.