Molecular phylogenetics, character evolution, and suprageneric classification of Lamioideae (Lamiaceae)

This paper presents a phylogenetic analysis of Lamiaceae subfam. Lamioideae (including subfamily Pogostemonoideae) based on sequences of the trnL intron, trnL-trnF intergenic spacer, and rps16 intron of the plastid genome. It is the first analysis that includes all major lamioid and pogostemonoid genera. Monophyly of Lamioideae s.l. (i.e., including Pogostemonoideae) is strongly supported, with Cymaria Benth. as its sister group, and Pogostemonoideae, which sometimes has been recognized as a subfamily, is subsumed in Lamioideae. On the basis of the phylogenetic hypothesis, Lamioideae is divided into nine tribes. Three new tribes are established: Gomphostemmateae Scheen & Lindqvist, Phlomideae Mathiesen, and Leucadeae Scheen & Ryding. The other six tribes are: Pogostemoneae Briq., Synandreae Raf., Stachydeae Dumort., Leonureae Dumort., Lamieae Coss. & Germ., and Marrubieae Vis. The genus Betonica L. is reestablished. The results also strongly suggest that the genera Stachys L., Sideritis L., Ballota L., and Leucas R. Br. are polyphyletic or paraphyletic. The results were used to examine evolution of non-molecular characters.