A taxonomic and phylogenetic study of the Golovinomyces biocellatus complex (Erysiphales, Ascomycota) using asexual state morphology and rDNA sequence data

The Golovinomyces biocellatus complex consists of powdery mildew (Erysiphales) species restricted to hosts of the family Lamiaceae. Previous authors used minor morphological features of the sexual state and host range data to split the complex. The data, however, were not sufficient to define a convincing species concept. Our taxonomic study is based on molecular phylogenetic and asexual state morphology data. For morphological studies, mainly features of the asexual morph (conidiophores, conidia, germination patterns) were studied using light and scanning electron microscopy. Detailed line drawings of asexual state features are provided. For phylogenetic analyses, two markers (rDNA: ITS, LSU) of 64 specimens were applied. The phylogeny resulted in two major clades. Clade I consists of specimens with Lamiaceae hosts and three specimens of Verbena. Clade II consists of two sister groups, the first (IIa) with Salvia spp. and the second (IIb) with Lycopus europaeus (the type host G. biocellatus) and Glechoma. Clades I and IIb and two subclades of IIa with Salvia hosts are characterized by specific morphological traits (differences in conidiophore length, conidial shape, width, and germination patterns). Based on these data, we suggest to consider specimens of clades I (including specimens on Verbena) and IIb and the two subclades of IIa as distinct species, namely G. monardae, G. biocellatus, G. salviae, and G. neosalviae sp. nov. A key for the identification of species based on asexual state features is provided. The results are discussed with respect to host range, jumps, co-evolutionary aspects, and distribution patterns.