A hotspot of lichen diversity and lichenological research in the Alps: the Paneveggio-Pale di San Martino Natural Park (Italy).

A checklist of 916 lichenised taxa is reported from the Paneveggio-Pale di San Martino Natural Park and its surroundings (Trentino-Alto Adige, N Italy), based on 7351 records from: (a) 72 literature sources, (b) eight public and private herbaria and (c) field observations by some of the authors. The study area appears as a hotspot of lichen diversity, hosting 30.1% of the lichen biota of the Alps in a territory that has 0.064% of their total surface area. This is mainly due to its high climatical, geological and orographic heterogeneity, but also to the long history of lichenological exploration, that started in the 19th century with Ferdinand Arnold and is still ongoing. The present work highlights the importance of detailed species inventories to support knowledge of biodiversity patterns, taxonomy and ecology and to properly address conservation issues. Fuscideamollisvar.caesioalbescens, Hydropunctariascabra, Protoparmeliabadiavar.cinereobadia and Variosporapaulii are new to Italy, 18 other taxa are new to Trentino-Alto Adige.

Refining the picture: new records to the lichen biota of Italy.

Based on the analysis of both historical and recent collections, this paper reports an annotated list of taxa which are new to the lichen biota of Italy or of its administrative regions. Specimens were identified using a dissecting and a compound microscope; routine chemical spot tests and standardized thin-layer chromatography (TLC or HPTLC). The list includes 225 records of 153 taxa. Twenty taxa are new to Italy, the others are new to one or more administrative regions, with 15 second records and 5 third records for Italy. Some of the species belong to recently-described taxa, others are poorly known, sterile or ephemeral lichens which were largely overlooked in Italy. Several species are actually rare, either because of the rarity of their habitats (e.g. old-growth forests), or because in Italy they are at the margins of their bioclimatic distribution. The picture of the lichen biota of Italy has now new pixels, but its grain is still coarse. Further analysis of historical collections, increased efforts in the exploration of some areas, and the taxonomic revision of critical groups are still necessary to provide more complete distributional data for new biogeographic hypotheses, taxonomic and ecological research, and biodiversity conservation.

The lichens of the Majella National Park (Central Italy): an annotated checklist.

The botanical exploration of the Majella National Park has a long tradition dating back to the eighteenth century. However, the lichen biota of this area is still poorly investigated. To provide a baseline for future investigations, in this annotated checklist, we summarised all available information on the occurrence of lichens in the Majella National Park, retrieved from previous literature, herbarium material and original data produced by recent research. The checklist includes 342 infrageneric taxa. However, seven taxa are considered as dubious, thus setting the number of accepted taxa at 335, i.e. 45.8% of those currently known to occur in the Abruzzo Region. This checklist provides a baseline of the lichens known to occur in the Majella National Park, highlighting the potential of this area as a hotspot of lichen biodiversity, especially from a biogeographical point of view as indicated by the occurrence of several arctic-alpine species that form disjunct populations in the summit area of the massif.

The lichens of the Alps - an annotated checklist.

This is the first attempt to provide an overview of the lichen diversity of the Alps, one of the biogegraphically most important and emblematic mountain systems worldwide. The checklist includes all lichenised species, plus a set of non- or doubtfully lichenised taxa frequently treated by lichenologists, excluding non-lichenised lichenicolous fungi. Largely based on recent national or regional checklists, it provides a list of all infrageneric taxa (with synonyms) hitherto reported from the Alps, with data on their distribution in eight countries (Austria, France, Germany, Liechtenstein, Monaco, Italy, Slovenia, Switzerland) and in 42 Operational Geographic Units, mostly corresponding to administrative subdivisions within the countries. Data on the main substrates and on the altitudinal distribution are also provided. A short note points to the main ecological requirements of each taxon and/or to open taxonomic problems. Particularly poorly known taxa are flagged and often provided with a short description, to attract the attention of specialists. The total number of infrageneric taxa is 3,163, including 117 non- or doubtfully lichenised taxa. The richness of the lichen biota fairly well corresponds with the percent of the Alpine area occupied by each country: Austria (2,337 taxa), Italy (2,169), France (2,028), Switzerland (1,835), Germany (1,168), Slovenia (890) and Lichtenstein (152), no lichen having ever been reported from Monaco. The number of poorly known taxa is quite high (604, 19.1% of the total), which indicates that, in spite of the Alps being one of the lichenologically most studied mountain systems worldwide, much work is still needed to reach a satisfactory picture of their real lichen diversity. Thirteen new combinations are proposed in the genera Agonimia, Aspicilia, Bagliettoa, Bellemerea, Carbonea, Lepra, Miriquidica, Polysporina, Protothelenella, Pseudosagedia and Thelidium.

Arthonia parietinaria - A common but frequently misunderstood lichenicolous fungus on species of the Xanthoria parietina-group.

Arthonia parietinaria is described as new to science. Host of the type and at the same time the only confirmed host species is the foliose macrolichen Xanthoria parietina. Sequence data of nucLSU rRNA genes reveal a close relationship to Arthonia molendoi. A. parietinaria is recorded for many countries in Europe, western Asia, and northern Africa.

Community Analyses Uncover High Diversity of Lichenicolous Fungi in Alpine Habitats.

Lichens are frequently colonized by specialized, lichenicolous fungi. Symptomatic lichenicolous fungi usually display typical phenotypes and reproductive structures on the lichen hosts. The classification based on these structures revealed different host specificity patterns. Other fungi occur asymptomatically in the lichen thalli and are much less known. We aimed at studying the diversity of lichen-associated fungi in specific, lichen-rich communities on rocks in the Alps. We tested whether lichenicolous fungi developing symptomatically on their known hosts also occur asymptomatically in other thalli of the same or of different host species. We collected lichen thalli according to a uniform sampling design comprising individuals adjacent to thalli that showed symptoms of lichenicolous fungal infections. The total fungal communities in the selected lichen thalli were further studied by single-strand conformation polymorphism (SSCP) fingerprinting analyses and sequencing of internal transcribed spacer (ITS) fragments. The systematic, stratified sampling strategy helped to recover 17 previously undocumented lichenicolous fungi and almost exhaustively the species diversity of symptomatic lichenicolous fungi in the studied region. The results from SSCP and the sequencing analyses did not reveal asymptomatic occurrence of normally symptomatic lichenicolous fungi in thalli of both the same and different lichen host species. The fungal diversity did not correlate with the species diversity of the symptomatic lichenicolous fungus-lichen host associations. The complex fingerprint patterns recovered here for fungal communities, in associations of well-delimited lichen thalli, suggest lichen symbiosis as suitable subjects for fungal metacommunity studies.

Lichenized and lichenicolous fungi from the Albanian Alps (Kosovo, Montenegro).

396 taxa (381 species) of lichenized and 45 species of lichenicolous fungi from the upper montane, subalpine and alpine belts of the Albanian Alps (= Prokletije Mountain Range, Bjeshkët e Nemuna) are presented. 92 lichenized and 26 lichenicolous fungi are new to Montenegro, 165 lichenized and 24 lichenicolous fungi are new to Kosovo, and 25 lichenized fungi (23 species) are new for the Balkan Peninsula.

A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317 genera and 66 families.

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.

The sterile microfilamentous lichenized fungi Cystocoleus ebeneus and Racodium rupestre are relatives of plant pathogens and clinically important dothidealean fungi.

The phylogenetic positions of the always-sterile microfilamentous lichens Cystocoleus ebeneus and Racodium rupestre were studied in a phylogenetic framework using sequence data of 5' nuSSU, nuLSU, and mtSSU rDNA. The analysis reveals that both genera are ascomycetes and belong to Dothideomycetidae: they are not close to lichenized members within the subclass, but rather belong to Capnodiales. The macroscopically scarcely distinguishable C. ebeneus and R. rupestre do not form a monophyletic group. The well-supported clade of R. rupestre is basal to the one in which C. ebeneus is close to Mycosphaerellaceae. This study provides another example of ascomycetes with very different life-styles and ecologies being closely related.

A five-gene phylogeny of Pezizomycotina.

Pezizomycotina is the largest subphylum of Ascomycota and includes the vast majority of filamentous, ascoma-producing species. Here we report the results from weighted parsimony, maximum likelihood and Bayesian phylogenetic analyses of five nuclear loci (SSU rDNA, LSU rDNA, RPB1, RPB2 and EF-lalpha) from 191 taxa. Nine of the 10 Pezizomycotina classes currently recognized were represented in the sampling. These data strongly supported the monophyly of Pezizomycotina, Arthoniomycetes, Eurotiomycetes, Orbiliomycetes and Sordariomycetes. Pezizomycetes and Dothideomycetes also were resolved as monophyletic but not strongly supported by the data. Lecanoromycetes was resolved as paraphyletic in parsimony analyses but monophyletic in maximum likelihood and Bayesian analyses. Leotiomycetes was polyphyletic due to exclusion of Geoglossaceae. The two most basal classes of Pezizomycotina were Orbiliomycetes and Pezizomycetes, both of which comprise species that produce apothecial ascomata. The seven remaining classes formed a monophyletic group that corresponds to Leotiomyceta. Within Leotiomyceta, the supraclass clades of Leotiomycetes s.s. plus Sordariomycetes and Arthoniomycetes plus Dothideomycetes were resolved with moderate support.

New insights into classification and evolution of the Lecanoromycetes (Pezizomycotina, Ascomycota) from phylogenetic analyses of three ribosomal RNA- and two protein-coding genes.

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.