Notes,outline and divergence times of Basidiomycota

The Basidiomycota constitutes a major phylum of the kingdom Fungi and is second in species numbers to the Ascomycota.The present work provides an overview of all validly published,currently used basidiomycete genera to date in a single document.An outline of all genera of Basidiomycota is provided,which includes 1928 currently used genera names,with 1263 synonyms,which are distributed in 241 families,68 orders,18 classes and four subphyla.We provide brief notes for each accepted genus including information on classification,number of accepted species,type species,life mode,habitat,distribution,and sequence information.Furthermore,three phylogenetic analyses with combined LSU,SSU,5.8s,rpb1,rpb2,and ef1 datasets for the subphyla Agaricomycotina,Pucciniomycotina and Ustilaginomycotina are conducted,respectively.Divergence time estimates are provided to the family level with 632 species from 62 orders,168 families and 605 genera.Our study indicates that the divergence times of the subphyla in Basidiomycota are 406-430 Mya,classes are 211-383 Mya,and orders are 99-323 Mya,which are largely consistent with previous studies.In this study,all phylogenetically supported families were dated,with the families of Agaricomycotina diverging from 27-178 Mya,Pucciniomycotina from 85-222 Mya,and Ustilaginomycotina from 79-177 Mya.Divergence times as additional criterion in ranking provide additional evidence to resolve taxonomic problems in the Basidiomycota taxonomic system,and also provide a better understanding of their phylogeny and evolution.

Species diversity of Basidiomycota

Fungi are eukaryotes that play essential roles in ecosystems.Among fungi,Basidiomycota is one of the major phyla with more than 40,000 described species.We review species diversity of Basidiomycota from five groups with different lifestyles or habitats:saprobic in grass/forest litter,wood-decaying,yeast-like,ectomycorrhizal,and plant parasitic.Case studies of Agaricus,Cantharellus,Ganoderma,Gyroporus,Russula,Tricholoma,and groups of lichenicolous yeast-like fungi,rust fungi,and smut fungi are used to determine trends in discovery of biodiversity.In each case study,the number of new species published during 2009–2020 is analysed to determine the rate of discovery.Publication rates differ between taxa and reflect different states of progress for species discovery in different genera.The results showed that lichenicolous yeast-like taxa had the highest publication rate for new species in the past two decades,and it is likely this trend will continue in the next decade.The species discovery rate of plant parasitic basidiomycetes was low in the past ten years,and remained constant in the past 50 years.We also found that the establishment of comprehensive and robust taxonomic systems based on a joint global initiative by mycologists could promote and standardize the recognition of taxa.We estimated that more than 54,000 species of Basidiomycota will be discovered by 2030,and estimate a total of 1.4–4.2 million species of Basidiomycota glob-ally.These numbers illustrate a huge gap between the described and yet unknown diversity in Basidiomycota.

The evolving species concepts used for yeasts:from phenotypes and genomes to speciation networks

Here we review how evolving species concepts have been applied to understand yeast diversity.Initially,a phenotypic species concept was utilized taking into consideration morphological aspects of colonies and cells,and growth profiles.Later the biological species concept was added,which applied data from mating experiments.Biophysical measurements of DNA similarity between isolates were an early measure that became more broadly applied with the advent of sequencing technology,leading to a sequence-based species concept using comparisons of parts of the ribosomal DNA.At present phylogenetic species concepts that employ sequence data of rDNA and other genes are universally applied in fungal taxonomy,including yeasts,because various studies revealed a relatively good correlation between the biological species concept and sequence divergence.The application of genome information is becoming increasingly common,and we strongly recommend the use of complete,rather than draft genomes to improve our understanding of species and their genome and genetic dynamics.Complete genomes allow in-depth comparisons on the evolvability of genomes and,consequently,of the species to which they belong.Hybridization seems a relatively common phenomenon and has been observed in all major fungal lineages that contain yeasts.Note that hybrids may greatly differ in their post-hybridization development.Future in-depth studies,initially using some model species or complexes may shift the traditional species concept as isolated clusters of genetically compatible isolates to a cohesive speciation network in which such clusters are interconnected by genetic processes,such as hybridization.

Notes for genera:Ascomycota

Knowledge of the relationships and thus the classification of fungi,has developed rapidly with increasingly widespread use of molecular techniques,over the past 10–15 years,and continues to accelerate.Several genera have been found to be polyphyletic,and their generic concepts have subsequently been emended.New names have thus been introduced for species which are phylogenetically distinct from the type species of particular genera.The ending of the separate naming of morphs of the same species in 2011,has also caused changes in fungal generic names.In order to facilitate access to all important changes,it was desirable to compile these in a single document.The present article provides a list of generic names of Ascomycota(approximately 6500 accepted names published to the end of 2016),including those which are lichen-forming.Notes and summaries of the changes since the last edition of‘Ainsworth&Bisby’s Dictionary of the Fungi’in 2008 are provided.The notes include the number of accepted species,classification,type species(with location of the type material),culture availability,life-styles,distribution,and selected publications that have appeared since 2008.This work is intended to provide the foundation for updating the ascomycete component of the"Without prejudice list of generic names of Fungi"published in 2013,which will be developed into a list of protected generic names.This will be subjected to the XIXth International Botanical Congress in Shenzhen in July 2017 agreeing to a modification in the rules relating to protected lists,and scrutiny by procedures determined by the Nomenclature Committee for Fungi(NCF).The previously invalidly published generic names Barriopsis,Collophora(as Collophorina),Cryomyces,Dematiopleospora,Heterospora(as Heterosporicola),Lithophila,Palmomyces(as Palmaria)and Saxomyces are validated,as are two previously invalid family names,Bartaliniaceae and Wiesneriomycetaceae.Four species of Lalaria,which were invalidly published are transferred to Taphrina and validated as new combinations.Catenomycopsis Tibell&Constant.is reduced under Chaenothecopsis Vain.,while Dichomera Cooke is reduced under Botryosphaeria Ces.&De Not.(Art.59).

Trends in yeast diversity discovery

Yeasts,usually defined as unicellular fungi,occur in various fungal lineages.Hence,they are not a taxonomic unit,but rather represent a fungal lifestyle shared by several unrelated lineages.Although the discovery of new yeast species occurs at an increasing speed,at the current rate it will likely take hundreds of years,if ever,before they will all be documented.Many parts of the earth,including many threatened habitats,remain unsampled for yeasts and many others are only superficially studied.Cold habitats,such as glaciers,are home to a specific community of cold-adapted yeasts,and,hence,there is some urgency to study such environments at locations where they might disappear soon due to anthropogenic climate change.The same is true for yeast communities in various natural forests that are impacted by deforestation and forest conversion.Many countries of the so-called Global South have not been sampled for yeasts,despite their economic promise.However,extensive research activity in Asia,especially China,has yielded many taxonomic novelties.Comparative genomics stud-ies have demonstrated the presence of yeast species with a hybrid origin,many of them isolated from clinical or industrial environments.DNA-metabarcoding studies have demonstrated the prevalence,and in some cases dominance,of yeast species in soils and marine waters worldwide,including some surprising distributions,such as the unexpected and likely common presence of Malassezia yeasts in marine habitats.

Correction to:The evolving species concepts used for yeasts:from phenotypes and genomes to speciation networks

The name of the second author was incorrectly captured in the initial online publication,and due to an error at the proofs stage,several proof corrections had been left undone.The original online article has been corrected.