Phylogenetic relationships, morphological incongruence, and geographic speciation in the fontinalaceae (Bryophyta)

Nuclear ribosomal DNA (internal transcribed spacer region) and chloroplast DNA (trnL-trnF region) were sequenced from 40 samples representing all three genera (Brachelyma, Dichelyma, and Fontinalis) and 18 species of the aquatic moss family, Fontinalaceae. Phylogenetic reconstructions recovered from separate and combined analyses were used to test the hypotheses that Fontinalis and Dichelyma are monophyletic (Brachelyma is monotypic), that groups of species within Fontinalis based on leaf morphology (keeled, concave, plane) form monophyletic groups, and that species delineation based on morphological characters within Fontinalis are congruent with nr- and cpDNA gene trees. Using Brachelyma subulata to root the tree, both Dichelyma and Fontinalis are monophyletic and patristically divergent (each united by >15 synapomorphic mutations). Groups of species within Fontinalis defined by leaf morphology are polyphyletic and it is clear that leaf morphology is labile in the genus. As defined morphologically, species of Fontinalis are nonmonophyletic for both nr- and cpDNA sequences and populations of some morphological taxa are separated in widely divergent clades. Molecular evidence suggests that at least some morphospecies are artificial, defined by convergent leaf forms. The weight of the evidence indicates that F. antipyretica is positively paraphyletic, with European populations more closely related to (i.e., share a more recent common ancestor with) European endemic species than to North American populations that are morphologically conspecific. North American populations are more closely related to North American endemic species.