Lithophyllum nagaokaense sp. nov. (Corallinales, Corallinophycidae, Rhodophyta): A new rhodolith-forming non-geniculate coralline alga from Japan

Lithophyllum is a cosmopolitan coralline algal genus with 12 species currently recognized in Japan based on modern morpho-anatomical taxonomic concepts primarily characterized by having uniporate tetrasporangial and bisporangial conceptacles, the presence of secondary pit-connections between cells of adjacent filaments and a dimerous thallus construction with a single basal layer of predominantly non-palisade cells. In the present study, we describe Lithophyllum nagaokaense sp. nov. based on a combination of molecular and morpho-anatomical data of specimens from the temperate waters of Japan. The new species forms both attached epilithic thalli and free-living rhodoliths. Phylogenetic analyses of psbA, rbcL and COI markers resolved L. nagaokaense as an independent species. Morpho-anatomically, L. nagaokaense resembles Lithophyllum johansenii, originally described from Australia, but distinguished from the latter by the absence of enlarged angular cells occluding pore canals of tetrasporangial conceptacle chambers. Lithophyllum nagaokaense shared a fruticose morphology with Lithophyllum kaiseri, Lithophyllum kuroshioense, Lithophyllum neo-okamurae, Lithophyllum okamurae, and Lithophyllum pygmaeum from Japan. Of these, L. nagaokaense is the most similar to L. neo-okamurae in having attached and free-living thalli with encrusting and warty to lumpy growth forms bearing knobby protuberances that are dichotomously branched or anastomosing. The new species is distinct from other Japanese fruticose species collectively by five tetrasporangial conceptacle chamber characters and the absence of trichocytes. A taxonomic key for identifying Japanese fruticose species is provided. This is the fifth Lithophyllum species and the third rhodolith-forming species verified by DNA sequencing to be found in Japan.