Kinematics of horizontal and vertical caterpillar crawling.

Unlike horizontal crawling, vertical crawling involves two counteracting forces: torque rotating the body around its center of mass and gravity resisting forward movement. The influence of these forces on kinematics has been examined in the soft-bodied larval stage of Manduca sexta. We found that crawling and climbing are accomplished using the same movements, with both segment timing and proleg lift indistinguishable in horizontal and vertical locomotion. Minor differences were detected in stride length and in the delay between crawls, which led to a lower crawling speed in the vertical orientation. Although these differences were statistically significant, they were much smaller than the variation in kinematic parameters between animals. The ability of Manduca to crawl and climb using the same movements is best explained by Manduca's relatively small size, slow speed and strong, controlled, passive grip made possible by its proleg/crochets.