Running on water: Three-dimensional force generation by basilisk lizards.

Water provides a unique challenge for legged locomotion because it readily yields to any applied force. Previous studies have shown that static stability during locomotion is possible only when the center of mass remains within a theoretical region of stability. Running across a highly yielding surface could move the center of mass beyond the edges of the region of stability, potentially leading to tripping or falling. Yet basilisk lizards are proficient water runners, regularly dashing across bodies of water to evade predators. We present here direct measurements of time-averaged force produced by juvenile plumed basilisk lizards (Basiliscus plumifrons) while running across water. By using digital particle image velocimetry to visualize fluid flow induced by foot movement, we show that sufficient support force is generated for a lizard to run across water and that novel strategies are also required to run across a highly yielding surface. Juvenile basilisk lizards produce greatest support and propulsive forces during the first half of the step, when the foot moves primarily vertically downwards into the water; they also produce large transverse reaction forces that change from medial (79% body weight) to lateral (37% body weight) throughout the step. These forces may act to dynamically stabilize the lizards during water running. Our results give insight into the mechanics of how basilisk lizards run across water and, on a broader scale, provide a conceptual basis for how locomotor surface properties can challenge established rules for the mechanics of legged locomotion.