Unraveling a 70-year-old taxonomic puzzle: Redefining the genus ikedosoma (Annelida: Echiura) on the basis of morphological and molecular analyses

After a long-standing taxonomic confusion, the echiurid genus Ikedosoma Bock, 1942, endemic to Japan and surroundings, is redefined on the basis of morphological and molecular analyses of many new Japanese materials and some museum specimens. The re-examination of a syntype of I. elegans (Ikeda, 1904), the type species of the genus, first revealed that its oblique muscle layer is continuous throughout and never fasciculate between longitudinal muscle bands, unlike those described in the definitions that have prevailed for ca. 70 years, making this genus indistinguishable from Listriolobus Spengel, 1912. Two Japanese species of Ikedosoma, I. elegans and I. gogoshimense (Ikeda, 1904), which were thus redefined, had also been poorly defined in the past to the point of being nearly indistinguishable from each other, largely due to incomplete descriptions and poor collections. Molecular phylogenetic analyses using 18S and 28S ribosomal RNA, histone H3, and cytochrome c oxidase subunit I (COI) genes clearly confirmed the distinction between these two species, their monophyletic origin, and their distinction from L. sorbillans (Lampert, 1883). The genus Ikedosoma thus validated is morphologically distinguishable from Listriolobus by the absence of a rectal caecum. Ikedosoma elegans and I. gogoshimense also differ in the disposition of gonoduct pairs. The third known species, I. qingdaoense Li, Wang and Zhou, 1994, from Qingdao, North China, lacks information on oblique muscle layers, which makes even its generic affiliation uncertain.