How the pilidium larva feeds

Introduction: The nemertean pilidium is a long-lived feeding larva unique to the life cycle of a single monophyletic group, the Pilidiophora, which is characterized by this innovation. That the pilidium feeds on small planktonic unicells seems clear; how it does so is unknown and not readily inferred, because it shares little morphological similarity with other planktotrophic larvae. Results: Using high-speed video of trapped lab-reared pilidia of Micrura alaskensis, we documented a multi-stage feeding mechanism. First, the external ciliation of the pilidium creates a swimming and feeding current which carries suspended prey past the primary ciliated band spanning the posterior margins of the larval body. Next, the larva detects prey that pass within reach, then conducts rapid and coordinated deformations of the larval body to re-direct passing cells and surrounding water into a vestibular space between the lappets, isolated from external currents but not quite inside the larva. Once a prey cell is thus captured, internal ciliary bands arranged within this vestibule prevent prey escape. Finally, captured cells are transported by currents within a buccal funnel toward the stomach entrance. Remarkably, we observed that the prey of choice - various cultured cryptomonads - attempt to escape their fate. Conclusions: The feeding mechanism deployed by the pilidium larva coordinates local control of cilia-driven water transport with sensorimotor behavior, in a manner clearly distinct from any other well-studied larval feeding mechanisms. We hypothesize that the pilidium's feeding strategy may be adapted to counter escape responses such as those deployed by cryptomonads, and speculate that similar needs may underlie convergences among disparate planktotrophic larval forms.