Phylogenetic relationships between Bryum and supposedly closely related genera

Abstract

The phylogeny of the genus Bryum was studied using cladistic analyses under the maximum parsimony criterion of evolution of anatomical and morphological characters. Three analyses were made with 32 Bryum species plus 20 species from genera supposedly closely related to Bryum, and with Amblyodon dealbatus (Sw. ex Hedw.) Bruch & Schimp., Meesia uliginosa Hedw., and Leptostomum macrocarpum (Hedw.) Bach. Pyl., as outgroups. It is here suggested that under earlier systematic concepts the genus Bryum is paraphyletic. A clade with Bryum billarderi Schwägr., B. capillare Hedw., B. donianum Grev., B. russulum Broth. & Geh., Rhodobryum giganteum (Schwägr.) Paris, and R. keniae (Müll. Hal.) Broth. are circumscribed by spathulate stem leaves that are crowded in the stem apex, suggesting that the rosulate species of Bryum are more closely related to Rhodobryum than to the rest of Bryum. Stem leaf costae without stereids and spores that mature in the winter are synapomorphies for a clade with Anomobryum julaceum (P. Gaertn. et al.) Schimp. and Bryum argenteum (Hedw.). The tropical species B. cellulare Hook. and B. flaccum Wilson ex Mitt. appear in a clade with Plagiobryum zieri (Dicks. ex Hedw.) Lindb. and Synthetodontium pringlei Cardot. In one analysis, B. limbatum Müll. Hal., Epipterygium tozeri (Grev.) Lindb., Leptobryum pyriforme (Hedw.) Wilson, and Roellia roellii (Broth. ex Röll) H.A. Crum came out in a clade with Mniobryum atropurpureum (Wahlenb.) I. Hagen, Mnium hornum Hedw., Pohlia cruda (Hedw.) Lindb., P. longicollis (Hedw.) Lindb., and Pseudopohlia didymodontia (Mitt.) A.L.Andrews. It is here suggested that gametophytic features, such as the orientation and anatomy of the stem leaves and the appearance of vegetative propagules, are important for the internal relationships within the studied ingroup, whereas characters related to the sporophyte, especially those of the peristome, may obscure phylogenetic relationships. Most of the subgenera and the sections of Bryum, as defined by earlier authors, appear to be paraphyletic. However, due to the low stability of most clades it is suggested that analyses including anatomical, morphological, and molecular data are needed.