Molecular phylogeny and taxonomic revision of the Philippine endemic Villaria Rolfe (Rubiaceae)

The little known Rubiaceae genus Villaria is endemic mostly to the coastal forests of the Philippines. Traditionally, it has been placed in the tribe Gardenieae. Later it was transferred to Octotropideae sensu Robbrecht and Puff. Villaria was placed among the "primitive" genera of the tribe, which are essentially characterized by large fruits, horizontal ovules and numerous seeds. Parsimony and Bayesian analyses of the combined plastid (rps16 and trnT-F) dataset strongly support the inclusion of Villaria in Octotropideae as well as monophyly of the genus. However, our molecular results do not conform to the current informal groups of the tribe delimited by fruit size, ovule position, number of seeds and exotesta thickenings. Instead, a close relationship between Villaria and two "central genera" (Hypobathrum and Pouchetia) is revealed for the first time. This clade is sister to a group comprising "primitive" (Fernelia), "advanced" (Kraussia and Polysphaeria) and "central" (Feretia) representatives. In addition, our combined tree strongly supports a sister taxa relationship between Canephora and Paragenipa. Villaria is characterized by unilocular ovaries, parietal placentation and strictly horizontal ovules. These features are unique within the Octotropideae. We recognize a total of five Villaria species, one new species (V. leytensis) is described here, and two species (V. philippinensis and V. rolfei) are transferred into synonymy with V. odorata. Each species is fully described, and a key to the species, a distribution map and illustrations are provided.