Kinematics of burrowing by peristalsis in granular sands

Peristaltic burrowing in muds applies normal forces to burrow walls, which extend by fracture, but the kinematics and mechanics of peristaltic burrowing in sands has not been explored. The opheliid polychaete Thoracophelia mucronata uses direct peristalsis to burrow in beach sands, with kinematics consistent with the 'dual anchor system' of burrowing described for diverse organisms. In addition to expansions associated with a constrictive direct peristaltic wave, worms alternately expand the head region, which is separated by septa from the open body cavity, and expansible lateral ridges that protrude from the 10th setiger. Tracking of chaetae with fluorescent dye showed that the body wall advances while segments are thin, then stationary segments expand, applying normal forces to burrow walls. These normal forces likely compact burrow walls and serve as anchors. Perhaps more importantly, peristaltic movements minimize friction with the burrow wall, which would expand dilatant sands. Considerable slipping of worms burrowing in a lower-density sand analog suggests that this dual-anchor peristaltic burrowing may be limited to a narrow range of mechanical properties of substrata, consistent with the limited habitat of T. mucronata in a narrow swash zone on dissipative beaches.