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Lichenized and non-lichenized filamentous ascomycetes produce a great variety of polyketide secondary metabolites. Some polyketide synthase (PKS) genes from non-lichenized fungi have been characterized, but the function of PKS genes from lichenized species remains unknown. Phylogenetic analysis of keto synthase (KS) domains allows prediction of the presence or absence of particular domains in the PKS gene. In the current study we screened genomic DNA from lichenized fungi for the presence of non-reducing and 6-methylsalicylic acid synthase (6-MSAS)-type PKS genes. We developed new degenerate primers in the acyl transferase (AT) region to amplify a PKS fragment spanning most of the KS region, the entire linker between KS and AT, and half of the AT region. Phylogenetic analysis shows that lichenized taxa possess PKS genes of the 6-MSAS-type. The extended alignment confirms overall phylogenetic relationships between fungal non-reducing, 6-MSAS-type and bacterial type I PKS genes.

We analysed ketoacyl synthase domains of type I polyketide synthase (PKS) gene fragments of 163 lichenized and 51 non-lichenized fungi in a Bayesian phylogenetic framework. Lichenized taxa from several unrelated taxonomic groups, some of which produce identical secondary metabolites, were included. We found 12 clades of non-reducing PKS genes, which represent monophyletic PKS paralogues. PAML and SELECTON analyses indicated that purifying selection is the prevailing selective force in the evolution of the keto synthase domain of these paralogues. We detected no unambiguous correlation between PKS clades and the distribution of lichen substances. Together with the strong evidence for purifying selection, the wide distribution of certain paralogues in ascomycetes suggested early gene duplication events in the evolutionary history of this gene family in the Ascomycota.

The phylogeny of Agyriaceae was investigated using MP and Bayesian approaches based on a combined dataset of nuLSU rDNA, mtSSU rDNA, and RPB1 sequences of 78 ascomycetes. The type genus of the family is shown to be a strongly supported sister to Coccotremataceae+Pertusariaceae, whereas the remaining species currently classified in Agyriaceae have a well-supported sister-group relationship with Baeomycetales. Monophyly of Agyriaceae is significantly rejected using two independent alternative topology tests. The micromorphology in Agyriaceae s. lat., Coccotremataceae, and Pertusariaceae is restudied. It is confirmed that the ascus type of Agyrium agrees with that of other taxa currently placed in Agyriaceae, and hence the ascus type is interpreted as homoplasious and phylogenetically misleading in this group of fungi. Consequently, we suggest that Trapeliaceae be resurrected for the lichenized taxa currently classified in Agyriaceae, and that this family be placed in Baeomycetales. The ordinal classification in Ostropomycetidae, especially the circumscription of Pertusariales and its distinction from Agyriales, requires additional studies.

Results of molecular studies regarding the phylogenetic placement of the order Ostropales and related taxa within Lecanoromycetes were thus far inconclusive. Some analyses placed the order as sister to the rest of Lecanoromycetes, while others inferred a position nested within Lecanoromycetes. We assembled a data set of 101 species including sequences from nuLSU rDNA, mtSSU rDNA, and the nuclear protein-coding RPB1 for each species to examine the cause of incongruencies in previously published phylogenies. MP, minimum evolution, and Bayesian analyses were performed using the combined three-region data set and the single-gene data sets. The position of Ostropales nested in Lecanoromycetes is confirmed in all single-gene and concatenated analyses, and a placement as sister to the rest of Lecanoromycetes is significantly rejected using two independent methods of alternative topology testing. Acarosporales and related taxa (Acarosporaceae group) are basal in Lecanoromycetes. However, if the these basal taxa are excluded from the analyses, Ostropales appear to be sister to the rest of Lecanoromycetes, suggesting different ingroup rooting as the cause for deviating topologies in previously published phylogenies.

The phylogenetic position of Trypetheliaceae was studied using partial sequences of the mtSSU and nuLSU rDNA of 100 and 110 ascomycetes, respectively, including 48 newly obtained sequences. Our analysis confirms Trypetheliaceae as monophyletic and places the family in Dothideomycetes. Pyrenulaceae, which were previously classified with Trypetheliaceae in Pyrenulales or Melanommatales, are supported as belonging to Chaetothyriomycetes. Monophyly of Pyrenulales, including Trypetheliaceae is rejected using three independent test methods. Monophyly of Arthopyreniaceae plus Trypetheliaceae, the two families including lichen-forming fungi in Dothideomycetes, is also rejected, as well as a placement of Trypetheliaceae in Pleosporales (incl. Melanommatales).

We examined the phylogenetic position of Omphalina foliacea and its putative closest relative, the lichen-forming basidiomycete Lichenomphalia lobata. Both species are restricted to specific habitats in the high andine regions of tropical America. We generated nuclear ITS and LSU rDNA sequences of two collections of each species and analysed the data together with sequences of omphalinoid basidiomycetes retrieved from GenBank using a Bayesian and a maximum parsimony approach. Our analyses show that both, O. foliacea and L. lobata are lichenized basidiomycetes. L. lohata is related to other species of the genus Lichenomphalia, whereas O. foliacea is not closely related to other Omphalina or Lichenomphalia species, but probably belongs in the hymenochaetoid clade. The analysis of a nuclear LSU data set of a broader sampling of hymenochaetoid basidiomycetes supported the placement of O. foliacea in this clade, but did not reveal any close relative.

The phylogeny of Lecanoromycetes (Ascomycota, Fungi) is investigated utilizing parsimony and Bayesian Markov Chain Monte Carlo analyses, of combined nLSU rDNA and mtSSU rDNA sequence datasets. The results suggest that Acarosporaceae, Candelariaceae, Phlyctis and Pycnora are not members of the monophyletic Lecanorales, and that Timdalia and Pleopsidium are members of a monophyletic Acarosporaceae. Pycnora, Candelariaceae and Acarosporaceae form a monophyletic group. Umbilicariaceae, Hypocenomyce scalaris, H. friesii, Ophioparmaceae, Boreoplaca, Elixia and Fuscidea form either a basal paraphyletic assemblage in Lecanoromycetes, or a monophyletic group which is the sister-group to Lecanorales and the rest of Lecanoromycetes (excluding Acarosporaceae). The Acarosporaceae forms a group with Pycnora and Candelariaceae, which may be outside the Lecanoromycetes. Chaetothyriales, Verrucariales, Eurotiales, Lichinales and Mycocaliciales form a monophyletic group, but with low support. We briefly discuss incongruence between datasets from different genetic markers, comparing the differences between the separate parsimony analyses, where the ILD test indicated a very significant incongruence. The phylogenetic significance of ascus-types that have influenced most recent Ascomycota classifications heavily is also discussed, and we finally point out risks with formalizing classifications too early.