Derivation of a polymorphic lineage of Gasteromycetes from boletoid ancestors

The phylogeny of selected gasteromycetes and hymenomycetes was inferred from partial nuclear large subunit rDNA (nuc-lsu, 28S) sequences, delimited by primers LR0R and LR5. Taxon sampling with emphasis on relationships within the Boletales further included some gasteroid groups, which obviously have evolved convergent fruiting body morphology, and therefore remained controversial in taxonomy. This study confirms the close relationship of Geastrales, Gauteriales and Phallales and the presumable derivation of Nidulariales and Tulostomatales within the euagarics clade, as widely accepted. In addition, four Hymenogaster species investigated were found to be in the euagarics clade and a relationship to the Cortinariaceae was indicated. The gasteroid fungus Zelleromyces stephensii is an example for maintaining morphological linkage by a lactiferous hyphal system to the genus Lactarius in the Russulales, and this relationship was affirmed in the sequence analysis. Several previously suggested relationships of gasteromycetes and Boletales were reproducible by analyzing nuc-lsu sequences. As a new result, Astraeus hygrometricus, the barometer earth star, is an additional representative of the Boletales. Together with Boletinellus, Phlebopus, Pisolithus, Calostoma, Gyroporus, Scleroderma, and Veligaster; Astraeus forms an unusual group comprising pileate-stipitate hymenomycetes and polymorphic gasteromycetes. This group is a major lineage within the Boletales and we propose the new suborder Sclerodermatineae, including the six families Boletinellaceae fam. nov. (Boletinellus and Phlebopus), Gyroporaceae (Singer) fam. nov. (Gyroporus), Pisolithaceae (Pisolithus), Astraeaceae (Astraeus), Calostomataceae (Calostoma), and the typus subordinis Sclerodermataceae (Scleroderma and Veligaster). Morphological and ecological characters, and pigment synthesis support the delimitation of the Sclerodermatineae, and indicate the radiation of different lineages in the Boletales originating from fungi with primitive tubular hymenophores. We regard such boletes with gyroid-boletinoid hymenophores, like Boletinellus, Gyrodon, Gyroporus, Paragyrodon and Phlebopus as key taxa in the evolution of Paxillineae, Sclerodermatineae and Boletineae.