Systematics of the calcified genera of the Galaxauraceae (Nemaliales, Rhodophyta) with an emphasis on Taiwan species

The current separation of the calcified genera Actinotrichia Decaisne, Galaxaura Lamouroux, and Tricleocarpa Huisman et Borowitzka in the Galaxauraceae is largely based on type of life history (whether the gametophytes and tetrasporophytes are isomorphic or heteromorphic with respect to gross morphology or cortical-cell features) and on features of postfertilization development (such as the composition of the pericarp). We reexamined the phylogenetic relationships of these genera based on comparative rbcL sequence analysis, types of life cycle, and cystocarp development. Four distinct assemblages have been identified: an Actinotrichia clade, a Tricleocarpa clade, a Galaxaura clade (containing the type species), and a Dichotomaria clade made of a number of formerly Galaxaura species (D. obtusata [Ellis et Solander] Lamarck, D. marginata [Ellis et Solander] Lamarck, and D. diesingiana [Zanardini] Huisman, Harper and Saunders). Key differences between Dichotomaria and Galaxaura include the habit of the gametophytic and tetrasporophytic generations (isomorphic in Dichotomaria and dimorphic in Galaxaura) as well as the presence or absence of a persistent pericarp in the cystocarp (present in Dichotomaria and absent in Galaxaura). Molecular data do not support monophyly for the putatively pantropical species Galaxaura rugosa, Dichotomaria obtusata, and D. marginata, all of which we conclude are in need of taxonomic revision.