Redescription of Cerebratulus marginatus auct. (Nemertea: Pilidiophora) from Hokkaido, Japan, as a new species

Abstract

The heteronemertean Cerebratulus orochi sp. nov. is described based on material collected intertidally at a muddy beach in Akkeshi, northern Japan. For the last 80 years, the species has been confused with Cerebratulus marginatus Renier, 1804; the latter was originally described from the Adriatic and once believed to occur in many places in the northern hemisphere including Japan. Cerebratulus orochi sp. nov. is morphologically different from all the congeners including C. marginatus by the following combination of characters: several layers of diagonal-muscle meshwork coated with connective tissue, proximo-distally distributed in cross section from the distal portion of the body-wall outer longitudinal muscle layer to the cutis-gland zone throughout the anterior portion of the body from the precerebral to the foregut regions; the cephalic vascular system consisting of lateral and mid-dorsal vessels; and the sub-rhynchocoelic vessel possessing a pair of antero-lateral diverticula before the former forks posteriorly into a pair of lower lateral vessels in the post-cerebral, pre-oral region. Previous records of C. marginatus from Japanese waters are no longer considered to be substantiated. Multi-locus phylogenetic analyses based on the mitochondrial 16S rRNA and cytochrome c oxidase subunit I (COI), as well as the nuclear 18S rRNA, 28S rRNA, and histone H3 genes among heteronemerteans comprising the “Cerebratulus clade” indicated that C. orochi sp. nov. was closely related to C. cf. marginatus from the US Pacific coast. A MegaBLAST search at the NCBI website with the 16S rRNA gene sequence from C. orochi sp. nov. followed by a couple of species delimitation analyses suggests that larvae of the species are also distributed in Vostok Bay, Far East Russia.