Quadrupedal Walking with the Skin: The Ambulatory Flaps in "Walking" Cuttlefish (Paintpot Cuttlefish, Metasepia tullbergi).

AbstractThe locomotion strategy of cephalopods is an important factor that influences their ability to exploit various oceanic environments. Particularly, Metasepia cuttlefish have a unique locomotion strategy; they prefer slow walking (ambling) on the seafloor over swimming. For this locomotion, they use their ventral arms as forelimbs and ambulatory flaps as hindlimbs. This locomotion is similar to the gait of quadruped vertebrates, where the forelimbs and hindlimbs on the left and right move alternately. The original description and some textbooks have considered these flaps to be muscular; however, this has not been proven. Here, we report the histological morphology of the ambulatory flaps of Metasepia tullbergi and their ambling locomotion. Histological observations indicated that the ambulatory flaps had a papillae structure comprising papillae musculature (dermal erector or retractor muscles) and connective tissue in the skin. Behavioral observations indicated that the ambulatory flaps changed their shape during ambling, which could explain the existence of the skin papillae. Our results suggest that ambulatory flaps are skin papillae, which can change shape by using their papillae musculature and connective tissue. This is a unique feature of Metasepia species that use the skin papillae for locomotion.