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We 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 protein-coding 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., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

Accurate specimen identification is challenging in groups with subtle or scarce taxonomically diagnostic characters, and the use of DNA barcodes can provide an effective means for consistent identification. Here, we investigate the utility of DNA barcode identification of species in a cosmopolitan genus of lichen-forming fungi, Parmelia (Parmeliaceae). Two hundred and two internal transcribed spacer (ITS) sequences generated from specimens collected from all continents, including Antarctica, were analysed, and DNA barcodes of 14 species of Parmelia s.s. are reported. Almost all species show a barcode gap. Overall, intraspecific divergence values were lower than the threshold previously established for Parmeliaceae. However, the mean and range were elevated by deep barcode divergences in three species, indicating the likely occurrence of overlooked specieslevel lineages. Here, we provide a DNA barcode reference library with well-identified specimens sampled worldwide and sequences from most of the type material to enable easy and fast accurate sample identification and to assist in uncovering overlooked species in Parmelia s.s. Further, our results confirm the efficiency of the ITS region in the identification of species of Parmelia s.s. [ABSTRACT FROM AUTHOR]

The lichen-forming fungal genus Flavoparmelia includes species with distinct distribution patterns, including subcos-mopolitan, restricted, and disjunct species. We used a dataset of nuclear ITS and LSU ribosomal DNA including 51 specimens to understand the influence of historical events on the current distribution patterns in the genus. We employed Bayesian, maxi-mum likelihood and maximum parsimony approaches for phylogenetic analyses, a likelihood-based approach to ancestral area reconstruction, and a Bayesian approach to estimate divergence times of major lineages within the genus. We identified two major clades in the genus, one of them separating into two subclades and one of those into four groups. Several of the groups and clades have restricted geographical ranges in the Southern Hemisphere, but two groups include species with wider distribution areas. Our analyses suggest that the genus originated in southern South America during the Eocene-Oligocene transition and that the diversification of the Australasian groups occurred recently. The subcosmopolitan distribution of species is explained by long-distance dispersal, while vicariance probably played a major role in the origin of the genus. Several currently accepted species were found to be non-monophyletic, indicating that the species delimitation in the genus requires further studies. [ABSTRACT FROM AUTHOR]

* Premise of the study: In spite of the recent advances in generic and species circumscriptions and in recognizing species diversity in lichen-forming fungi, the timing of speciation and the factors that promote diversification in lichens remain largely unexplored. We used brown parmelioids as a model to assess the timing of divergence and explore the impact of geological and climatic events on lineage divergence and diversification in lichenized fungi. Additionally, to clarify the phylogenetic position of the species currently placed in Melanelia disjuncta group, we evaluated the taxonomic status and phylogenetic relationships within Parmeliaceae. * Methods: Phylogenetic relationships and divergence time estimates were inferred from a four-loci data set. Alternative hypotheses were tested using Shimodaira-Hasegawa and expected likelihood weights tests. * Key results: The M. disjuncta group forms a strongly supported, monophyletic lineage independent from Melanelia s.s. The M. disjuncta clade arose ca. 23.1 million years ago (Ma). Our results suggest that most of the lineages within the clade diversified during the Miocene (17.6 to 11.2 Ma). The split of other brown parmelioids, such as Emodomelanelia-Melanelixia occurred ca. 41.70 Ma, and the radiation of Melanelixia began during the Eocene-Oligocene transition (ca. 33.75 Ma). * Conclusions: Montanelia is described here as a new genus to accommodate species of the Melaneliadisjuncta group. Further, the study indicates that the current species delimitation within the newly described genus requires revision. We provide evidence of lineage divergence of Montanelia at the Oligocene-Miocene boundary. Our results indicate that the diversification during Miocene would have happened during major mountain uplifts. [ABSTRACT FROM AUTHOR]

We used a genetic distance approach in conjunction with molecular phylogeny to establish species boundaries and detect cryptic lineages in the Parmotrema reticulatum – P. pseudoreticulatum complex. The phylogeny of specimens from a broad geographic distribution was reconstructed from the internal transcribed spacer region. Pairwise genetic distances were calculated and compared to an intraspecific range defined for the parmelioid lichens to circumscribe species-level groups. Our results showed that P. reticulatum and P. pseudoreticulatum are polyphyletic, being comprised of at least seven well-supported lineages. In contrast, the genetic distance approach revealed ten cryptic lineages within the P. reticulatum – P. pseudoreticulatum complex. Neither morphology nor geography was conclusive in attempting to corroborate these genetic lineages. However FST indices suggest significant genetic differentiation between these lineages. Our results suggest that the morphology-based circumscriptions underestimated species in Parmotrema and that, in some cases, genetic distances may be used as an additional tool to determine species boundaries in morphologically cryptic species complexes. The most significant contribution of the present study is the application of a fast and accurate method to identify problematic groups and candidate species using the ITS locus with a genetic distances approach. [ABSTRACT FROM AUTHOR]

This study focuses on European Usnea species with sorediate shrubby thalli, with the aim to evaluate the morphological and chemical separation of species in the light of molecular data. Twenty-two Usnea species, including widely distributed taxa such as U. diplotypus, U. fulvoreagens, U. glabrescens, U. lapponica, U. subfloridana, U. substerilis and U. wasmuthii, were included in the study using Bayesian and maximum parsimony analyses of nuclear ITS and beta-tubulin sequences. The analyses showed that: 1) most taxa that are morphologically well delimited are also distinct by means of molecular characters, 2) shrubby taxa in the section Usnea that are difficult to determine by traditional characters form a group of closely related but still genetically distinct entities, except U. diplotypus and U. substerilis which appear to be polyphyletic. The branch lengths differed largely between two parts of the ITS tree (sections Usnea and Ceratinae). Usnea intermedia is proposed as the sexually reproducing counterpart for the sorediate U. lapponica. Additionally, some new chemotypes of Usnea species were determined. [ABSTRACT FROM AUTHOR]

Two asexual reproductive strategies of the common lichen Pseudevernia furfuracea are described. Although the species propagates mainly by isidia, some specimens also show the development of soralia. Morphological, chemical and molecular analyses were performed on three such sorediate specimens from the Canary Islands, Morocco and Turkey. Maximum parsimony, maximum likelihood and Bayesian analyses indicate that: a) sorediate samples represent only a morphological variant of the reproductive mode and b) the separation of taxa (at species level or below) on the basis of their containing either olivetoric acid or physodic and oxyphysodic acids is not appropriate. In addition, a phylogenetic reconstruction of the genus Pseudevernia is presented for the first time. The tree shows two sister monophyletic clades, one containing American species (P. intensa, P. cladonia, P. consocians), and the second encompassing the P. furfuracea samples (including sorediate specimens). The biological and taxonomic significance of soralia in sorediate samples is discussed. [ABSTRACT FROM AUTHOR]

Abstract: The species delimitation in fungi is currently in flux. A growing body of evidence shows that the morphology-based species circumscription underestimates the number of existing species. The large and ever growing number of DNA sequence data of fungi makes it possible to use these to identify potential cases of hidden species, which then need to be studied with extensive taxon samplings. We used Parmeliaceae, one of the largest families of lichenized fungi as a model. Intra- and interspecific distances derived from maximum-likelihood phylogenetic trees inferred from 491 nuclear ITS rDNA sequences were examined for five major clades of parmelioid lichens. The intra- and interspecific distances were well separated in most cases allowing the calculation of a threshold, with exceptions of highly deviating distances in a few cases. These situations are shown to be taxa in which the current delimitation needs revision. Thus the analysis of the distance distributions is shown to be a powerful tool for identifying species complexes. [Copyright &y& Elsevier]

Biogeographical studies of lichens used to be complicated because of the large distribution ranges of many species. Molecular systematics has revitalized lichen biogeography by improving species delimitation and providing better information about species range limitations. This study focuses on the major clade of tropical parmelioid lichens, which share a chemical feature, the presence of isolichenan in the cell wall, and a morphological feature, microscopic pores in the uppermost layer. Our previous phylogenetic studies revealed that the largest genus in this clade, Hypotrachyna, is polyphyletic with a clade mainly distributed in South and East Asia clustering distant from the core of the genus. To divide the Hypotrachyna clade into monophyletic groups and to reevaluate morphological and chemical characters in a phylogenetic context, we sampled ITS, nuclear large subunit (nuLSU) and mitochondrial small subunit (mtSSU) rDNA sequences from 77 species. We are erecting the new genus Remototrachyna for a core group of 15 former Hypotrachyna species. The segregation of Remototrachyna from Hypotrachyna receives support from morphological and chemical data, as well from maximum parsimony, maximum likelihood, and Bayesian phylogenetic analyses of the DNA. We used a likelihood approach to study the geographic range evolution of Remototrachyna and Bulbothrix, which are sister groups. This analysis suggests that the ancestral range of Remototrachyna was restricted to India and that subsequent long-distance dispersal is responsible for the pantropical occurrence of two species of Remototrachyna. [ABSTRACT FROM AUTHOR]

Morphological and phylogenetic relationships of the worldwide Mediterranean lichen forming fungus, Parmelina quercina, have been studied. Specimens from western Europe, western North America and southern Australia were analysed using molecular data (nuITS rDNA, nuLSU rDNA and mtSSU rDNA) and selected morphological features (upper cortex maculae, scanning electron microscopy examination of the epicortex, ascospores and conidia shape and size, and amphithecial retrorse rhizines). The results conclusively reveal that: (1) there is not one single species but four separate species in the Mediterranean or sub Mediterranean areas of the world. Parmelina quercina and Parmelina carporrhizans (Euroasiatic species), Parmelina coleae sp. nov. (North America) and Parmelina elixia sp. nov. (Australia); (2) largely debated P. carporrhizans is not a synonym of P. quercina but supported as a valid species circumscribed to Macaronesic relict sites; (3) the geographical isolation of the Australian population is correlated with a large genetic distance; (4) morphological characters (ascospores and conidial variability and thallus epicortex) correlate with the phylogenetic hypothesis; (5) the new or revalidated species within Parmelina quercina are not cryptic species but morphologically recognizable taxa. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91, 455–467. [ABSTRACT FROM AUTHOR]

Abstract: Sections of apothecia were used to study the internal morphology of ascospores in the largest monophyletic clade within Parmeliaceae composed of Xanthoparmelia and related genera. The results were compared with fertile representative species of most other parmelioid clades. All the Xanthoparmelia species had spores with a single smooth vacuole, which was peanut-shaped, with different degrees of constriction in the equatorial plane. This differs from the ellipsoid vacuole of other parmelioids. In the Xanthoparmelia clade, sexual reproduction seems much more common than in other parmelioids. Thus, we suggest that the presence of this unique spore morphology might contribute to the evolutionary success of this monophyletic group. Further, the discovery of this useful ascospore character demonstrates that detailed ascospore morphological studies significantly enhance molecular phylogenetic analyses. Ascospore features may be more taxonomically significant in Parmeliaceae than hitherto considered. [Copyright &y& Elsevier]