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.

A six-gene phylogenetic overview of Basidiomycota and allied phyla with estimated divergence times of higher taxa and a phyloproteomics perspective

In this paper,we provide a phylogenetic overview of Basidiomycota and related phyla in relation to ten years of DNA based phylogenetic studies since the AFTOL publications in 2007.We selected 529 species to address phylogenetic relationships of higher-level taxa using a maximumlikelihood framework and sequence data from six genes traditionally used in fungal molecular systematics(nrLSU,nrSSU,5.8S,tef1-a,rpb1 and rpb2).These species represent 18 classes,62 orders,183 families,and 392 genera from the phyla Basidiomycota(including the newly recognized subphylum Wallemiomycotina)and Entorrhizomycota,and 13 species representing 13 classes of Ascomycota as outgroup taxa.We also conducted a molecular dating analysis based on these six genes for 116 species representing 17 classes and 54 orders of Basidiomycota and Entorrhizomycota.Finally we performed a phyloproteomics analysis from 109 Basidiomycota species and 6 outgroup taxa using amino-acid sequences retrieved from 396 orthologous genes.Recognition of higher taxa follows the criteria in Zhao et al(Fungal Divers 78:239–292,2016):(i)taxa must be monophyletic and statistically well-supported in molecular dating analyses,(ii)their respective stem ages should be roughly equivalent,and(iii)stem ages of higher taxa must be older than those of lower level taxa.The time-tree indicates that the mean of stem ages of Basidiomycota and Entorrhizomycota are ca.530 Ma;subphyla of Basidiomycota are 406–490 Ma;most classes are 358–393 Ma for those of Agaricomycotina and 245–356 Ma for those of Pucciniomycotina and Ustilaginomycotina;most orders of those subphyla split 120–290 Ma.Monophyly of most higherlevel taxa of Basidiomycota are generally supported,especially those taxa introduced in the recent ten years:phylum Entorrhizomycota,classes Malasseziomycetes,Moniliellomycetes,Spiculogloeomycetes,Tritirachiomycetes and orders Amylocorticiales,Golubeviales,Holtermanniales,Jaapiales,Lepidostromatales,Robbauerales,Stereopsidales and Trichosporonales.However,the younger divergence times of Leucosporidiales(Microbotryomycetes)indicate that its order status is not supported,thus we propose combining it under Microbotryales.On the other hand,the families Buckleyzymaceae and Sakaguchiaceae(Cystobasidiomycetes)are raised to Buckleyzymales and Sakaguchiales due to their older divergence times.Cystofilobasidiales(Tremellomycetes)has an older divergence time and should be amended to a higher rank.We however,do not introduce it as new class here for Cystofilobasidiales,as DNA sequences from these taxa are not from their respective types and thus await further studies.Divergence times for Exobasidiomycetes,Cantharellales,Gomphales and Hysterangiales were obtained based on limited species sequences in molecular dating study.More comprehensive phylogenetic studies on those four taxa are needed in the future because our ML analysis based on wider sampling,shows they are not monophyletic groups.In general,the six-gene phylogenies are in agreement with the phyloproteomics tree except for the placements of Wallemiomycotina,orders Amylocorticiales,Auriculariales,Cantharellales,Geastrales,Sebacinales and Trechisporales from Agaricomycetes.These conflicting placements in the six-gene phylogeny vs the phyloproteomics tree are discussed.This leads to future perspectives for assessing gene orthology and problems in deciphering taxon ranks using divergence times.

Towards standardizing taxonomic ranks using divergence times-a case study for reconstruction of the Agaricus taxonomic system

The recognition of taxonomic ranks in the Linnean classification system is largely arbitrary.Some authors have proposed the use of divergence time as a universally standardized criterion.Agaricus(Agaricaceae,Agaricales)is a mushroom genus that contains many species of high commercial value.Recent studies using ITS sequence data discovered 11 new phylogenetic lineages within the genus,however their taxonomic ranks were uncertain due to the lack of criteria to define them within traditional taxonomy.In this study,we analyzed ITS sequence data from 745 collections(nearly 600 being newly generated)including 86 from type specimens of previously recognized subgenera and sections.Many monophyletic groups were recognized,but most basal relationships were unresolved.One hundred and fourteen representatives of the identified ITS clades were selected in order to produce a multi-gene phylogeny based on combined LSU,tef-1α,and rpb2 sequence data.Divergence times within the multi-gene phylogeny were estimated using BEAST v1.8.Based on phylogenetic relationships and with respect to morphology,we propose a revised taxonomic system for Agaricus that considers divergence time as a standardized criterion for establishing taxonomic ranks.We propose to segregate Agaricus into five subgenera and 20 sections.Subgenus Pseudochitonia is substantially emended;circumscription of the subgenera Agaricus and Flavoagaricus is restricted to taxa of sections Agaricus and Arvenses,respectively;and two new subgenera(Minores and Spissicaules)are introduced.Within Pseudochitonia,sections Bivelares,Brunneopicti,Chitonioides,Nigrobrunnescentes,Sanguinolenti and Xanthodermatei are maintained,but the latter two are reduced because we raise subsection Bohusia to sectional rank and a clade within section Xanthodermatei is formally introduced as section Hondenses;and sections Rubricosi,Crassispori,Flocculenti,and Amoeni are introduced.Section Laeticolores is placed in the subgenus Minores and sections Rarolentes and Subrutilescentes are placed in the subgenus Spissicaules.Twenty-two new species belonging to various sections are described.This work exemplifies that ITS data,while useful at lower taxonomic levels(i.e.,detection of species and species groups),are of limited value for inferring deeper phylogenetic relationships.Finally,we suggest that the establishment of a standardized taxonomic system based on divergence times could result in a more objective,and biologically more meaningful,taxonomic ranking of fungi.

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.

Molecular phylogeny, morphology, pigment chemistry and ecology in Hygrophoraceae (Agaricales)

Molecular phylogenies using 1–4 gene regions and information on ecology,morphology and pigment chemistry were used in a partial revision of the agaric family Hygrophoraceae.The phylogenetically supported genera we recognize here in the Hygrophoraceae based on these and previous analyses are:Acantholichen,Ampulloclitocybe,Arrhenia,Cantharellula,Cantharocybe,Chromosera,Chrysomphalina,Cora,Corella,Cuphophyllus,Cyphellostereum,Dictyonema,Eonema,Gliophorus,Haasiella,Humidicutis,Hygroaster,Hygrocybe,Hygrophorus,Lichenomphalia,Neohygrocybe,Porpolomopsis and Pseudoarmillariella.A new genus that is sister to Chromosera is described as Gloioxanthomyces.Revisions were made at the ranks of subfamily,tribe,genus,subgenus,section and subsection.We present three new subfamilies,eight tribes(five new),eight subgenera(one new,one new combination and one stat.nov.),26 sections(five new and three new combinations and two stat.nov.)and 14 subsections(two new,two stat.nov.).Species of Chromosera,Gliophorus,Humidicutis,and Neohygrocybe are often treated within the genus Hygrocybe;we therefore provide valid names in both classification systems.We used a minimalist approach in transferring genera and creating new names and combinations.Consequently,we retain in the Hygrophoraceae the basal cuphophylloid grade comprising the genera Cuphophyllus,Ampulloclitocybe andCantharocybe,despite weak phylogenetic support.We include Aeruginospora and Semiomphalina in Hygrophoraceae based on morphology though molecular data are lacking.The lower hygrophoroid clade is basal to Hygrophoraceae s.s.,comprising the genera Aphroditeola,Macrotyphula,Phyllotopsis,Pleurocybella,Sarcomyxa,Tricholomopsis and Typhula.