Abstract
BACKGROUND: The nemertean pilidium is one of the most notable planktotrophic larval types among marine invertebrates. The juvenile forms inside the larva from a series of isolated rudiments, called the imaginal discs. The development culminates in catastrophic metamorphosis, in which the larval body is consumed by the juvenile worm. Although the pilidium was first described in 1847, and is commonly found among marine plankton, there is not a single complete description of its development. The few published studies of pilidial development are based on observations of typically unidentified larvae opportunistically collected from plankton at various developmental stages. RESULTS: The development of Micrura alaskensis, a common Northwest Pacific coast intertidal nemertean, is described from fertilization to metamorphosis. A staging scheme is proposed based on characteristic developmental milestones. Three pairs of imaginal discs develop as invaginations of larval epidermis. The cephalic discs invaginate from the larval epidermis above the ciliated band, while the cerebral organ discs and the trunk discs invaginate below the ciliated band. All paired imaginal disc invaginations are closely associated with different portions of the larval ciliated band. In addition, two unpaired rudiments contribute to the juvenile - the proboscis rudiment and the dorsal rudiment, which do not develop as invaginations. A pair of thick-walled esophageal pouches previously thought to represent nephridial rudiments give rise to the juvenile foregut. Branched rudiments of protonephridia, and their efferent ducts are also described. Larval and juvenile serotonergic nervous systems are briefly described. Development of the juvenile is completed by 5-8 weeks at 11-15 degrees C. During the rapid metamorphosis the juvenile emerges from and devours the larva. CONCLUSIONS: This study is the first description of pilidial development from fertilization to metamorphosis in a single species. It is illustrated with photomicrographs of live larvae, diagrams, confocal images, and videos. The findings are discussed in the context of previously published accounts of pilidial development, with which they disagree on several accounts. The results described here indicate a different number, origin and fate of various juvenile rudiments. The proposed staging scheme will be useful in subsequent studies of pilidial development.