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Phylogenetic analyses using mtSSU and nuITS sequences of Buellia violaceofusca (previously placed in Lecanoromycetes), a sterile, sorediate lichen having a trebouxioid photobiont, surprisingly prove that the species is conspecific with Lecanographa amylacea (Arthoniomycetes), a fertile, esorediate species with a trentepohlioid photobiont. These results suggest that L. amylacea and B. violaceofusca are photomorphs of the same mycobiont species, which, depending on the photobiont type, changes the morphology and the reproduction strategy. This is the first example of a lichenized fungus that can select between Trebouxia (Trebouxiophyceae) and trentepohlioid (Ulvophyceae) photobionts. Trebouxia photobionts from the sorediate morphotype belong to at least three different phylogenetic clades, and the results suggest that Lecanographa amylacea can capture the photobiont of other lichens such as Chrysothrix candelaris to form the sorediate morphotype. Phylogenetic analyses based on rbcL DNA data suggest that the trentepohlioid photobiont of L. amylacea is closely related to Trentepohlia isolated from fruticose lichens. The flexibility in the photobiont choice enables L. amylacea to use a larger range of tree hosts. This strategy helps the lichen to withstand changes of environmental conditions, to widen its distribution range and to increase its population size, which is particularly important for the survival of this rare species.

Taeniolella is a genus of asexual ascomycetes with saprophytic, endophytic, and lichenicolous life styles. A phylogeny of representative species is presented, with a focus on lichenicolous taxa. We obtained mtSSU and nuLSU sequence data from culture isolates of Taeniolella and from freshly collected specimens of other taxa. Taeniolella is recovered as strongly polyphyletic. The type species, Taeniolella exilis, is placed in Kirschsteiniotheliaceae within Dothideomycetes. Other saprophytic/endophytic Taeniolella species previously assigned to Sordariomycetes based on sequences were found to represent either contaminants or species that cannot be assigned to Taeniolella for morphological reasons. Lichenicolous species are restricted to Asterotexiales (Dothideomycetes) where the sequenced species of Taeniolella do not form a monophyletic group, but are related to species of Buelliella s. lat., Karschia, Labrocarpon, Melaspilea s. lat., and Stictographa. Molecular data are, however, not sufficient to reallocate the lichenicolous Taeniolella species to other genera so far. Anamorph-teleomorph relationships between these taxa and lichenicolous Taeniolella are discussed but could not be demonstrated. The type of Buelliella is placed in Asterotexiales, and the genus recovered as polyphyletic. Three new lichenicolous Taeniolella species are described, Taeniolella hawksworthiana, Taeniolella pyrenulae and Taeniolella toruloides. Taeniolella rudis is transferred to Sterigmatobotrys., (Copyright © 2016 British Mycological Society. Published by Elsevier Ltd. All rights reserved.)

Lichenicolous fungi are a specialized group of taxa which inhabit lichens and develop diverse degrees of specificity and parasitic behaviour towards their hosts. They are recognized only by their phenotypic symptoms and sexual or asexual spore-producing structures on the lichen thalli. Only recently, molecular data and culture dependent approaches have helped in uncovering the species diversity and in verifying the phylogenetic position and anamorph-teleomorph relationships of some taxa. Here, we studied the phylogenetic placement of representative taxa of two lichenicolous genera, the coelomycete Lichenodiplis and the ascomycete Muellerella. We obtained molecular data for three nuclear and mitochondrial loci (28S, 18S, and 16S), both from fresh collected specimens and culture isolates. Our multilocus phylogeny places Lichenodiplis and Muellerella samples in one monophyletic, fully supported clade, sister to Epibryon (Epibryaceae) in Chaetothyriales (Eurotiomycetes). Morphological analyses of axenically cultured fungi show the formation of conidiomata and conidiospores in both Lichenodiplis and Muellerella isolates. We suggest that the species Lichenodiplis lecanorae and Muellerella atricola represent, respectively, the anamorphic and teleomorphic stages of the same fungus and discuss their relationships with the other fungal families in Chaetothyriomycetidae., (Copyright © 2015 The Authors. Published by Elsevier Ltd.. All rights reserved.)

The generic segregates of the widespread fruticose genus Ramalina (mostly based on empirical data on morphology, cortex anatomy and secondary metabolites) are studied using maximum parsimony, maximum likelihood and Bayesian analyses of nuclear LSU and ITS sequences. The species examined include the three species aggregates within Niebla from the western coasts of North America, all species except one assumed to belong to the same genus from Macaronesia and the Mediterranean basin, the type species of Dievernia and Ramalina, and representatives of the genus Fistulariella. The genus Niebla is strongly supported when restricted to species from the New World, and all species referred to it from Macaronesia and the Mediterranean basin belong to Ramalina (R. bourgeana, R. crispatula, R. cupularis, R. hamulosa, R. portosantana, R. rosacea, R. subwebbiana and R. webbii). No support is found for the genera Dievernia and Fistulariella. The internal topology of the large genus Ramalina is unresolved and needs further studies., (Copyright © 2010 The British Mycological Society. Published by Elsevier Ltd. All rights reserved.)

We present a 6-gene, 420-species maximum-likelihood phylogeny of Ascomycota, the largest phylum of Fungi. This analysis is the most taxonomically complete to date with species sampled from all 15 currently circumscribed classes. A number of superclass-level nodes that have previously evaded resolution and were unnamed in classifications of the Fungi are resolved for the first time. Based on the 6-gene phylogeny we conducted a phylogenetic informativeness analysis of all 6 genes and a series of ancestral character state reconstructions that focused on morphology of sporocarps, ascus dehiscence, and evolution of nutritional modes and ecologies. A gene-by-gene assessment of phylogenetic informativeness yielded higher levels of informativeness for protein genes (RPB1, RPB2, and TEF1) as compared with the ribosomal genes, which have been the standard bearer in fungal systematics. Our reconstruction of sporocarp characters is consistent with 2 origins for multicellular sexual reproductive structures in Ascomycota, once in the common ancestor of Pezizomycotina and once in the common ancestor of Neolectomycetes. This first report of dual origins of ascomycete sporocarps highlights the complicated nature of assessing homology of morphological traits across Fungi. Furthermore, ancestral reconstruction supports an open sporocarp with an exposed hymenium (apothecium) as the primitive morphology for Pezizomycotina with multiple derivations of the partially (perithecia) or completely enclosed (cleistothecia) sporocarps. Ascus dehiscence is most informative at the class level within Pezizomycotina with most superclass nodes reconstructed equivocally. Character-state reconstructions support a terrestrial, saprobic ecology as ancestral. In contrast to previous studies, these analyses support multiple origins of lichenization events with the loss of lichenization as less frequent and limited to terminal, closely related species.

A multi-locus phylogenetic study of the order Arthoniales is presented here using the nuclear ribosomal large subunit (nuLSU), the second largest subunit of RNA polymerase II (RPB2) and the mitochondrial ribosomal small subunit (mtSSU). These genes were sequenced from 43 specimens or culture isolates representing 33 species from this order, 16 of which were from the second largest genus, Opegrapha. With the inclusion of sequences from GenBank, ten genera and 35 species are included in this study, representing about 18% of the genera and ca 3% of the species of this order. Our study revealed the homoplastic nature of morphological characters traditionally used to circumscribe genera within the Arthoniales, such as exciple carbonization and ascomatal structure. The genus Opegrapha appears polyphyletic, species of that genus being nested in all the major clades identified within Arthoniales. The transfer of O. atra and O. calcarea to the genus Arthonia will allow this genus and family Arthoniaceae to be recognized as monophyletic. The genus Enterographa was also found to be polyphyletic. Therefore, the following new combinations are needed: Arthonia calcarea (basionym: O. calcarea), and O. anguinella (basionym: Stigmatidium anguinellum); and the use of the names A. atra and Enterographa zonata are proposed here. The simultaneous use of a mitochondrial gene and two nuclear genes led to the detection of what seems to be a case of introgression of a mitochondrion from one species to another (mitochondrion capture; cytoplasmic gene flow) resulting from hybridization.

The Lecanoromycetes includes most of the lichen-forming fungal species (> 13500) and is therefore one of the most diverse class of all Fungi in terms of phenotypic complexity. We report phylogenetic relationships within the Lecanoromycetes resulting from Bayesian and maximum likelihood analyses with complementary posterior probabilities and bootstrap support values based on three combined multilocus datasets using a supermatrix approach. Nine of 10 orders and 43 of 64 families currently recognized in Eriksson's classification of the Lecanoromycetes (Outline of Ascomycota--2006 Myconet 12:1-82) were represented in this sampling. Our analyses strongly support the Acarosporomycetidae and Ostropomycetidae as monophyletic, whereas the delimitation of the largest subclass, the Lecanoromycetidae, remains uncertain. Independent of future delimitation of the Lecanoromycetidae, the Rhizocarpaceae and Umbilicariaceae should be elevated to the ordinal level. This study shows that recent classifications include several nonmonophyletic taxa at different ranks that need to be recircumscribed. Our phylogenies confirm that ascus morphology cannot be applied consistently to shape the classification of lichen-forming fungi. The increasing amount of missing data associated with the progressive addition of taxa resulted in some cases in the expected loss of support, but we also observed an improvement in statistical support for many internodes. We conclude that a phylogenetic synthesis for a chosen taxonomic group should include a comprehensive assessment of phylogenetic confidence based on multiple estimates using different methods and on a progressive taxon sampling with an increasing number of taxa, even if it involves an increasing amount of missing data.