Phylogenetic re-evaluation of Thielavia with the introduction of a new family Podosporaceae

The genus Thielavia is morphologically defined by having non-ostiolate ascomata with a thin peridium composed of textura epidermoidea, and smooth, single-celled, pigmented ascospores with one germ pore. Thielavia is typified with Th. basicola that grows in close association with a hyphomycete which was traditionally identified as Thielaviopsis basicola. Besides Th. basicola exhibiting the mycoparasitic nature, the majority of the described Thielavia species are from soil, and some have economic and ecological importance. Unfortunately, no living type material of Th. basicola exists, hindering a proper understanding of the classification of Thielavia. Therefore, Thielavia basicola was neotypified by material of a mycoparasite presenting the same ecology and morphology as described in the original description. We subsequently performed a multi-gene phylogenetic analyses (rpb2, tub2, ITS and LSU) to resolve the phylogenetic relationships of the species currently recognised in Thielavia. Our results demonstrate that Thielavia is highly polyphyletic, being related to three family-level lineages in two orders. The redefined genus Thielavia is restricted to its type species, Th. basicola, which belongs to the Ceratostomataceae (Melanosporales) and its host is demonstrated to be Berkeleyomyces rouxiae, one of the two species in the "Thielaviopsis basicola" species complex. The new family Podosporaceae is sister to the Chaetomiaceae in the Sordariales and accommodates the re-defined genera Podospora, Trangularia and Cladorrhinum, with the last genus including two former Thielavia species (Th. hyalocarpa and Th. intermedia). This family also includes the genetic model species Podospora anserina, which was combined in Triangularia (as Triangularia anserina). The remaining Thielavia species fall in ten unrelated clades in the Chaetomiaceae, leading to the proposal of nine new genera (Carteria, Chrysanthotrichum, Condenascus, Hyalosphaerella, Microthielavia, Parathielavia, Pseudothielavia, Stolonocarpus and Thermothielavioides). The genus Canariomyces is transferred from Microascaceae (Microascales) to Chaetomiaceae based on its type species Can. notabilis. Canariomyces is closely related to the human-pathogenic genus Madurella, and includes three thielavia-like species and one novel species. Three monotypic genera with a chaetomium-like morph (Brachychaeta, Chrysocorona and Floropilus) are introduced to better resolve the Chaetomiaceae and the thielavia-like species in the family. Chrysocorona lucknowensis and Brachychaeta variospora are closely related to Acrophialophora and three newly introduced genera containing thielavia-like species; Floropilus chiversii is closely related to the industrially important and thermophilic species Thermothielavioides terrestris (syn. Th. terrestris). This study shows that the thielavia-like morph is a homoplastic form that originates from several separate evolutionary events. Furthermore, our results provide new insights into the taxonomy of Sordariales and the polyphyletic Lasiosphaeriaceae.