Origin and Evolution of the Lichenized Ascomycete Order Lichinales: Monophyly and Systematic Relationships Inferred from Ascus, Fruiting Body and SSU rDNA Evolution

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Abstract
The Lichinales are a group of lichenized ascomycetes that almost exclusively possess cyanobacteria as their primary photobiont and are hitherto separated from the Lecanorales, the major group of lichenized ascomycetes, by thallus structure, ascoma ontogeny, ascus structure and ascus function. The relationship of the two families Peltulaceae and Lichinaceae, both placed within the Lichinales, with the Heppiaceae, placed within the Lecanorales, was investigated, as well as a possible sister group relationship of the Lichinales to the Lecanorales. Phylogenetic analyses included non‐molecular data as well as 18S rDNA sequence data. The monophyly of the Lichinales including the family Heppiaceae and a sister group relationship of Lichinales and Lecanorales, based on the shared presence of lecanoralean asci, are proposed in a morphological hypothesis. Parsimony and distance analyses of 18S rDNA sequence data strongly support the monophyly of the Lichinales, including all three families. Therefore, the presence of rostrate, lecanoralean asci in Peltula and part of the Lichinaceae suggests that this ascus type is an autapomorphy of the monophyletic Lichinales. Furthermore, the occurrence of prototunicate asci in the Heppiaceae and most of the Lichinaceae is autapomorphic and was gained independently by reduction of the rostrate ascus. The 18S rDNA analysis did not reject the non‐molecular hypothesis of a sister group relationship of the Lichinales and the Lecanorales as based on ascus characters. The alternative placement of the Lichinales as the sister group of all inoperculate euascomycetes excluding the Sordariomycetes and most of the Leotiales in the gene tree received unsufficient bootstrap support and no support from any non‐molecular data and consequently was rejected.