Molecular, phytochemical and morphological characterization of the liverwort genus Radula in Portugal (mainland, Madeira, Azores)

Relationships of the eight species of the liverwort genus Radula occurring in Portugal (mainland, the Madeira and Azores archipelagos), including the Macaronesian endemics R. jonesii and R. wichurae, were evaluated based on molecular, phytochemical and morphological-anatomical data. Maximum parsimony and Bayesian analyses were performed with sequences from three plastid DNA markers (trnSGGA-rps4 spacer, rps4 gene, trnLUAA intron), volatile oil compounds, as well as qualitative morphological-anatomical characters. In addition, the molecular data were subjected to maximum likelihood analysis. The eight taxa, R. aquilegia, R. carringtonii, R. complanata, R. holtii, R. jonesii, R. lindenbergiana, R. nudicaulis and R. wichurae, can be clearly distinguished from each other, either by molecular data alone or by combination of characters from all three data sets. Radula aquilegia is monophyletic according to the molecular data, but shows considerable, yet undescribed intraspecific morphological and phytochemical variability. Recognition of R. complanata and R. lindenbergiana as separate species, previously based solely on the paroecious vs. dioecious sexual condition, is moderately supported by the molecular phylogenetic analyses and strongly supported by the phytochemical data. The Radula species, narrowly distributed in Macaronesia and Atlantic Europe, probably have two different origins. For Radula holtii and R. nudicaulis, connections with Radula species from the Neotropics are indicated. The other species, among them the two Macaronesian endemics, are closely related with the R. complanata/R. lindenbergiana complex, which is widely distributed in the northern hemisphere.