H. Thorsten Lumbsch

Reconstructing the early evolution of Fungi using a six-gene phylogeny

The ancestors of fungi are believed to be simple aquatic forms with flagellated spores, similar to members of the extant phylum Chytridiomycota (chytrids). Current classifications assume that chytrids form an early-diverging clade within the kingdom Fungi and imply a single loss of the spore flagellum, leading to the diversification of terrestrial fungi. Here we develop phylogenetic hypotheses for Fungi using data from six gene regions and nearly 200 species. Our results indicate that there may have been at least four independent losses of the flagellum in the kingdom Fungi. These losses of swimming spores coincided with the evolution of new mechanisms of spore dispersal, such as aerial dispersal in mycelial groups and polar tube eversion in the microsporidia (unicellular forms that lack mitochondria). The enigmatic microsporidia seem to be derived from an endoparasitic chytrid ancestor similar to Rozella allomycis, on the earliest diverging branch of the fungal phylogenetic tree.

The ascomycota tree of life: A phylum-wide phylogeny clarifies the origin and evolution of fundamental reproductive and ecological traits

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.

Outline of Ascomycota: 2017

Taxonomic placement of genera have been changing rapidly as taxonomists widely use DNA sequence data in phylogenetic and evolutionary studies. It is essential to update existing databases/outlines based on recent studies, since these sources are widely used as a foundation for other research. In this outline, we merge both asexual and sexual genera into one outline. The phylum Ascomycota comprises of three subphyla viz. Pezizomycotina (including 13 classes, 124 orders and 507 families), Saccharomycotina (including one class, one order and 13 families) and Taphrinomycotina (five classes, five orders and six families). Approximately, 6600 genera have been listed under different taxonomic ranks including auxiliary (intermediate) taxonomic ranks.

Recognition of seven species in the Cryptococcus gattii/Cryptococcus neoformans species complex

Phylogenetic analysis of 11 genetic loci and results from many genotyping studies revealed significant genetic diversity with the pathogenic Cryptococcus gattii/Cryptococcus neoformans species complex. Genealogical concordance, coalescence-based, and species tree approaches supported the presence of distinct and concordant lineages within the complex. Consequently, we propose to recognize the current C. neoformans var. grubii and C. neoformans var. neoformans as separate species, and five species within C. gattii. The type strain of C. neoformans CBS132 represents a serotype AD hybrid and is replaced. The newly delimited species differ in aspects of pathogenicity, prevalence for patient groups, as well as biochemical and physiological aspects, such as susceptibility to antifungals. MALDI-TOF mass spectrometry readily distinguishes the newly recognized species.

Phylogeography of the montane caddisfly Drusus discolor: evidence for multiple refugia and periglacial survival.

We studied the genetic population structure and phylogeography of the montane caddisfly Drusus discolor across its entire range in central and southern Europe. The species is restricted to mountain regions and exhibits an insular distribution across the major mountain ranges. Mitochondrial sequence data (COI) of 254 individuals from the entire species range is analysed to reveal population genetic structure. The data show little molecular variation within populations and regions, but distinct genetic differentiation between mountain ranges. Most populations are significantly differentiated based on FST and exact tests of population differentiation and most haplotypes are unique to a single mountain range. Phylogenetic analyses reveal deep divergence between geographically isolated lineages. Combined, these results suggest that past fragmentation is the prominent process structuring the populations across Europe. We use tests of selective neutrality and mismatch distributions, to study the demographic population history of regions with haplotype overlap. The high level of genetic differentiation between mountain ranges and estimates of demographic history provide evidence for the existence of multiple glacial refugia, including several in central Europe. The study shows that these aquatic organisms reacted differently to Pleistocene cooling than many terrestrial species. They persisted in numerous refugia over multiple glacial cycles, allowing many local endemic clades to form.

Fungi evolved right on track.

Dating of fungal divergences with molecular clocks thus far has yielded highly inconsistent results. The origin of fungi was estimated at between 660 million and up to 2.15 billion y ago, and the divergence of the two major lineages of higher fungi, Ascomycota and Basidiomycota, at between 390 million y and up to 1.5 billion y ago. Assuming that these inconsistencies stem from various causes, we reassessed the systematic placement of the most important fungal fossil, Paleopyrenomycites, and recalibrated internally unconstrained, published molecular clock trees by applying uniform calibration points. As a result the origin of fungi was re-estimated at between 760 million and 1.06 billion y ago and the origin of the Ascomycota at 500–650 million y ago. These dates are much more consistent than previous estimates, even if based on the same phylogenies and molecular clock trees, and they are also much better in line with the fossil record of fungi and plants and the ecological interdependence between filamentous fungi and land plants. Our results do not provide evidence to suggest the existence of ancient protolichens as an alternative to explain the ecology of early terrestrial fungi in the absence of land plants.

Myconet Volume 14. Part One. Outline of Ascomycota—2009. Part Two. Notes on Ascomycete Systematics. Nos. 4751–5113

Abstract Part One presents the current classification that includes all accepted genera and higher taxa above the generic level in the phylum Ascomycota. It is based on the changes listed in Myconet notes 4751–5113. In the current outline, three subphyla (Pezizomycotina, Saccharomycotina, Taphrinomycotina) are accepted. Taphrinomycotina includes four classes (Neolectomycetes, Pneumocystidomycetes, Schizosaccharomyces, Taphrinomycetes). Saccharomycotina consists of one class and Pezizomycotina includes eleven classes (Arthoniomycetes, Dothideomycetes, Eurotiomycetes, Geoglossomycetes, Laboulbeniomycetes, Lecanoromycetes, Leotiomycetes, Lichinomycetes, Orbiliomycetes, Pezizomycetes, Sordariomycetes). Part Two presents 363 notes on the taxonomy and nomenclature of ascomycetes (Ascomycota) at the generic and higher levels. Numerous changes in higher levels resulted from multi-authored phylogenetic papers resulting from the AFTOL and Deep Hyphae projects, especially a publication on a revised fungal classification spearheaded by David Hibbett in 2007 and a 2009 volume of Studies in Mycology edited by Conrad Schoch and others dedicated to the phylogeny of the Dothideomycetes. The new name Alectoria gowardii Lumbsch is proposed for Gowardia arctica P. Halonen, L. Myllys, S. Velmala & H. Hyvärinen (non Alectoria arctica Elenkin & Savicz).

The Faces of Fungi database: fungal names linked with morphology, phylogeny and human impacts

Taxonomic names are key links between various databases that store information on different organisms. Several global fungal nomenclural and taxonomic databases (notably Index Fungorum, Species Fungorum and MycoBank) can be sourced to find taxonomic details about fungi, while DNA sequence data can be sourced from NCBI, EBI and UNITE databases. Although the sequence data may be linked to a name, the quality of the metadata is variable and generally there is no corresponding link to images, descriptions or herbarium material. There is generally no way to establish the accuracy of the names in these genomic databases, other than whether the submission is from a reputable source. To tackle this problem, a new database (FacesofFungi), accessible at www.facesoffungi.org (FoF) has been established. This fungal database allows deposition of taxonomic data, phenotypic details and other useful data, which will enhance our current taxonomic understanding and ultimately enable mycologists to gain better and updated insights into the current fungal classification system. In addition, the database will also allow access to comprehensive metadata including descriptions of voucher and type specimens. This database is user-friendly, providing links and easy access between taxonomic ranks, with the classification system based primarily on molecular data (from the literature and via updated web-based phylogenetic trees), and to a lesser extent on morphological data when molecular data are unavailable. In FoF species are not only linked to the closest phylogenetic representatives, but also relevant data is provided, wherever available, on various applied aspects, such as ecological, industrial, quarantine and chemical uses. The data include the three main fungal groups (Ascomycota, Basidiomycota, Basal fungi) and fungus-like organisms. The FoF webpage is an output funded by the Mushroom Research Foundation which is an NGO with seven directors with mycological expertise. The webpage has 76 curators, and with the help of these specialists, FoF will provide an updated natural classification of the fungi, with illustrated accounts of species linked to molecular data. The present paper introduces the FoF database to the scientific community and briefly reviews some of the problems associated with classification and identification of the main fungal groups. The structure and use of the database is then explained. We would like to invite all mycologists to contribute to these web pages.

A molecular phylogeny and a new classification of parmelioid lichens containing Xanthoparmelia -type lichenan (Ascomycota: Lecanorales)

Generic concepts in the parmelioid lichens have been discussed intensively over the past three decades without reaching a broad consensus. We have now employed molecular data from three genes to provide a basis for a revised generic concept of the parmelioid lichens containing Xanthoparmelia-type lichenan. The phylogeny of the parmelioid lichens containing Xanthoparmelia-type lichenan was reconstructed using a combined Bayesian analysis of nuclear ITS, LSU rDNA and mitochondrial SSU rDNA sequences, and a maximum parsimony analysis was also made for comparison. 179 new partial sequences of 58 taxa were generated and 12 sequences were downloaded from GenBank. Our results indicate that the lichens containing Xanthoparmelia-type lichenan form a monophyletic group. However, the segregates proposed earlier do not form distinct clades within the group. Alternative hypotheses of monophyletic Karoowia and Neofuscelia that are not nested within Xanthoparmelia were rejected with our dataset; Karoowia is polyphyletic, and Neofuscelia is reduced to synonymy under Xanthoparmelia. Xanthomaculina convoluta also belongs to Xanthoparmelia. Since we were unable to sequence the umbilicate type species ofXanthomaculina, we refrain from synonymizing that genus with Xanthoparmelia here. The synonymy of Chondropsis and Paraparmelia under Xanthoparmelia already proposed is supported. The revised and enlarged genus Xanthoparmelia includes species that have cell walls with Xanthoparmelia-type lichenan, a palisade plectenchyma with a pored epicortex, lack pseudocyphellae, with usually simple rhizines, generally bifusiform conidia, and medullary chemical diversity. Ten new names are proposed, and 129 new combinations are made into Xanthoparmelia.

Goodbye morphology? A paradigm shift in the delimitation of species in lichenized fungi

Accurate species delimitations are crucial for ecological and conservation studies, assessments of biotic diversity, and identifying factors driving diversification. Estimates suggest that the vast majority of fungal species are currently unknown. Although many undescribed fungal taxa are expected to be indentified within understudied groups and from underexplored areas, mounting evidence suggests a substantial number of unrecognized fungal species are likely hidden within traditional phenotype-based species in lichen-forming fungi. Molecular genetics has revolutionized our ability to assess traditional species concepts and provides additional tools for robust species delimitation. In general, lichens display few taxonomically usefully characters; therefore molecular data have gained great importance in delimiting fungal species in lichen symbioses. As a result, the taxonomic value of phenotypical characters is now much better understood, and in many cases previously overlooked characters have been identified supporting molecular-based species circumscriptions. Although in some cases molecular research has verified traditional hypotheses, most studies repeatedly show that our current interpretation of morphological and chemical characters is inadequate to accurately characterize diversity. Here we report on the role of molecular data in understanding species-level diversity in lichenized fungi by reviewing current literature, focusing primarily on Ascomycota. While finding and applying the appropriate character sets and analytical tools remains one of the greatest challenges to empirical species delimitation in lichen-forming fungi, the available literature indicates that the inclusion of molecular data in species circumscription is crucial to establish robust hypotheses of species boundaries in this important group of fungi.