Integrative Taxonomy of the Micrura alaskensis Coe, 1901 Species Complex (Nemertea: Heteronemertea), with Descriptions of a New Genus Maculaura gen. nov and Four New Species from the NE Pacific

Micrura alaskensis Coe, 1901 is a common intertidal heteronemertean known from eastern and northwest Pacific (Alaska to Ensenada, Mexico and Akkeshi, Japan, respectively). It is an emerging model system in developmental biology research. We present evidence from morphology of the adults, gametes, and sequences of cytochrome c oxidase subunit I and 16S rRNA genes that it is not one, but a complex of five, cryptic species. All five of these species co-occur at least in part of their geographic range (e.g. southern Oregon). Preliminary cross-hybridization experiments suggest that at least some of these species are reproductively isolated. The five species share characteristics of adult morphology (e.g. accessory buccal glands) and at least four are known to possess a unique larval morphotype-pilidium maculosum. We propose that these characters define a new genus, Maculaura gen. nov., which contains the following five species: Maculaura alaskensis comb. nov., Maculaura aquilonia sp. nov., Maculaura cerebrosa sp. nov., Maculaura oregonensis sp. nov., and Maculaura magna sp. nov. It is unclear which of the five species Coe originally encountered and described. We chose to retain the name alaskensis for the species that current researchers know as Micrura alaskensis, although, presently, it is only known from Washington and Oregon, and has not been collected from Alaska. Maculaura aquilonia sp. nov. is the only member of the genus we have encountered in Alaska, and we show that it also occurs in the Sea of Okhotsk, Russia.