The Lecanoromycetes is the largest class of lichenized Fungi, and one of the
most species-rich classes in the kingdom. Here we provide a multigene phylogenetic
synthesis (using three ribosomal RNA-coding and two protein-coding genes) of the
Lecanoromycetes based on 642 newly generated and 3329 publicly available sequences
representing 1139 taxa, 317 genera, 66 families, 17 orders and five subclasses (four
currently recognized: Acarosporomycetidae, Lecanoromycetidae, Ostropomycetidae,
Umbilicariomycetidae; and one provisionarily recognized,
‘Candelariomycetidae’). Maximum likelihood phylogenetic analyses on four
multigene datasets assembled using a cumulative supermatrix approach with a progressively
higher number of species and missing data (5-gene, 5+4-gene, 5+4+3-gene and 5+4+3+2-gene
datasets) show that the current classification includes non-monophyletic taxa at various
ranks, which need to be recircumscribed and require revisionary treatments based on denser
taxon sampling and more loci. Two newly circumscribed orders (Arctomiales and Hymeneliales
in the Ostropomycetidae) and three families (Ramboldiaceae and Psilolechiaceae in the
Lecanorales, and Strangosporaceae in the Lecanoromycetes inc. sed.) are introduced. The
potential resurrection of the families Eigleraceae and Lopadiaceae is considered here to
alleviate phylogenetic and classification disparities. An overview of the photobionts
associated with the main fungal lineages in the Lecanoromycetes based on available
published records is provided. A revised schematic classification at the family level in
the phylogenetic context of widely accepted and newly revealed relationships across
Lecanoromycetes is included. The cumulative addition of taxa with an increasing amount of
missing data (i.e., a cumulative supermatrix approach, starting with taxa for which
sequences were available for all five targeted genes and ending with the addition of taxa
for which only two genes have been sequenced) revealed relatively stable relationships for
many families and orders. However, the increasing number of taxa without the addition of
more loci also resulted in an expected substantial loss of phylogenetic resolving power
and support (especially for deep phylogenetic relationships), potentially including the
misplacements of several taxa. Future phylogenetic analyses should include additional
single copy protein-coding markers in order to improve the tree of the Lecanoromycetes. As
part of this study, a new module (“Hypha”) of the freely available
Mesquite software was developed to compare and display the internodal support values
derived from this cumulative supermatrix approach.
Phylogenetic relationships within the genus Cladonia, including Cladina (Cladoniaceae, Lecanoromycetes), were reconstructed based upon simultaneous analyses of DNA sequences and morphological and chemical data. We used sequences from the internal transcribed spacer 1 (ITS1), the 5.8 rDNA gene, and the internal transcribed spacer 2 (ITS2) of the nuclear rDNA gene cluster, and partial sequences from the protein‐coding β‐tubulin gene. The analyses included 235 specimens of 168 taxa representing all currently recognized sections of Cladonia and Cladina and the outgroup genera Cladia, Pycnothelia, and Ramalea. Analyses were performed using optimization alignment with three different parameter values. The results of all analyses support the inclusion of Cladina in Cladonia. The current sectional division of Cladonia was not supported, and a new provisional classification for the genus is proposed.
The phylogenetic relationships of the genus Bryoria were examined using ITS, partial glyceraldehyde 3-phosphate dehydrogenase (GAPDH) and mitochondrial SSU rDNA sequence data in addition to 20 chemical and morphological characters. This first comprehensive molecular study to assess Bryoria phylogeny includes representatives from all the traditionally recognized four sections. Combined cladistic analyses of 88 Bryoria specimens representing at least 25 species resulted in highly resolved phylogenies. Based on the results, a new infrageneric classification for the genus is proposed. Five sections are recognized, largely corresponding to the existing classification, with the addition of section Americanae. The study shows that while most species with an erect growth-form are clearly monophyletic, current species status of many pendent taxa can be questioned.
SummaryWe studied the evolutionary history of the Parmeliaceae (Lecanoromycetes, Ascomycota), one of the largest families of lichen-forming fungi with complex and variable morphologies, also including several lichenicolous fungi. We assembled a six-locus data set including nuclear, mitochondrial and low-copy proteincoding genes from 293 operational taxonomic units (OTUs).The lichenicolous lifestyle originated independently three times in lichenized ancestors within Parmeliaceae, and a new generic name is introduced for one of these fungi. In all cases, the independent origins occurred c. 24 million yr ago. Further, we show that the Paleocene, Eocene and Oligocene were key periods when diversification of major lineages within Parmeliaceae occurred, with subsequent radiations occurring primarily during the Oligocene and Miocene.Our phylogenetic hypothesis supports the independent origin of lichenicolous fungi associated with climatic shifts at the Oligocene-Miocene boundary. Moreover, diversification bursts at different times may be crucial factors driving the diversification of Parmeliaceae. Additionally, our study provides novel insight into evolutionary relationships in this large and diverse family of lichen-forming ascomycetes.
Phylogenetic relationships within the family Parmeliaceae (lichenized ascomycetes) with emphasis on the heterogeneous group of cetrarioid lichens are reconstructed. The results are based on cladistic analyses of DNA-sequences, morphological and chemical data. Almost all currently recognized cetrarioid genera were included in the analyses together with parmelioid and alectorioid members of the presumably monophyletic family Parmeliaceae. We tried to sample taxonomic diversity of the family as widely as possible. The ITS1-5.8S-ITS2 region of the rDNA and a partial β-tubulin gene from 126 samples representing 82 species were analysed. Cetrarioid lichens were identified as a monophyletic group, supported by both ITS and β-tubulin characters. This group was reanalysed using 47 morphological, anatomical and secondary chemistry characters combined with the DNA data matrix. ITS and β-tubulin sequences provide congruent information, and a clear correlation between DNA-data and conidial shape is observed. The current taxonomy of the cetrarioid lichens is discussed and compared with the phylogenetic trees obtained here. A comprehensive study of the phylogeography of some bipolar or subcosmopolitic species with representatives from both hemispheres was performed. Cetraria aculeata is the only taxon where correlation between DNA-data and geographic origin is observed.
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