Larval Development of Two N. E. Pacific Pilidiophoran Nemerteans (Heteronemertea; Lineidae)

Unique to the phylum Nemertea, the pilidium is an unmistakable planktonic larva found in one group of nemerteans, the Pilidiophora. Inside the pilidium, the juvenile develops from a series of epidermal invaginations in the larval body, called imaginal discs. The discs grow and fuse around the larval gut over the course of weeks to months in the plankton. Once complete, the juvenile breaks free from the larval body in a catastrophic metamorphosis, and often devours the larva as its first meal. One third of nemertean species are expected to produce a pilidium, but the larvae are known for very few species; development from fertilization to metamorphosis has been described in only one species, Micrura alaskensis. Known pilidia include both planktotrophic and lecithotrophic forms, and otherwise exhibit great morphological diversity. Here, we describe the complete development in two lineiform species that are common to the northeast Pacific coast, Micrura wilsoni and Lineus sp. "red." Both species possess typical, cap-shaped planktotrophic pilidia, and the order of emergence of imaginal discs is similar to that which is described in M. alaskensis. The pilidium of Lineus sp. "red" resembles pilidia of several other species, such as Lineus flavescens, and potentially characterizes a pilidiophoran clade. M. wilsoni has relatively transparent oocytes and a pilidium with what appears to be a unique pattern of pigmentation. The adults of both species are more commonly observed in intertidal zones than their larvae are in the plankton.