Three out of one: revising the species delimitation of the tree fern Gymnosphaera salletii (Cyatheaceae), with particular reference to the foliar nectary

In Indochina the species diversity of Cyatheaceae remains insufficiently known. Here we demonstrate the scaly tree fern Gymnosphaera salletii in Vietnam to be an assemblage of three distinct species based on molecular and morphological data, and report for the first time the occurrence of foliar nectaries in Gymnosphaera. The true G. salletii is morphologically unique based on the protuberant glands present on the petiole. Gymnosphaera phankelocii sp. nov. and Gymnosphaeravietnamensis sp. nov. are recognized as new species; G. phankelocii is characterized by its distant two-rowed sori and G. vietnamensis by the sub-tripinnate fronds. Our phylogenetic analyses indicated that G. phankelocii is a relatively isolated species, while G. vietnamensis is a member of the G. salletii clade along with G. austroyunnanensis, G. salletii, and G. saxicola. The two new species and members of the G. salletii clade are all local endemics, and provide good examples to understand the frequent local endemics and their evolution in scaly tree ferns. Eleven species of Gymnosphaera are now known from Indochina; we provide a diagnostic key to these species.