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.

Fungal diversity notes 929-1035:taxonomic and phylogenetic contributions on genera and species of fungi

This article is the ninth in the series of Fungal Diversity Notes,where 107 taxa distributed in three phyla,nine classes,31 orders and 57 families are described and illustrated.Taxa described in the present study include 12 new genera,74 new species,three new combinations,two reference specimens,a re-circumscription of the epitype,and 15 records of sexualasexual morph connections,new hosts and new geographical distributions.Twelve new genera comprise Brunneofusispora,Brunneomurispora,Liua,Lonicericola,Neoeutypella,Paratrimmatostroma,Parazalerion,Proliferophorum,Pseudoastrosphaeriellopsis,Septomelanconiella,Velebitea and Vicosamyces.Seventy-four new species are Agaricus memnonius,A.langensis,Aleurodiscus patagonicus,Amanita flavoalba,A.subtropicana,Amphisphaeria mangrovei,Baorangia major,Bartalinia kunmingensis,Brunneofusispora sinensis,Brunneomurispora lonicerae,Capronia camelliaeyunnanensis,Clavulina thindii,Coniochaeta simbalensis,Conlarium thailandense,Coprinus trigonosporus,Liua muriformis,Cyphellophora filicis,Cytospora ulmicola,Dacrymyces invisibilis,Dictyocheirospora metroxylonis,Distoseptispora thysanolaenae,Emericellopsis koreana,Galiicola baoshanensis,Hygrocybe lucida,Hypoxylon teeravasati,Hyweljonesia indica,Keissleriella caraganae,Lactarius olivaceopallidus,Lactifluus midnapurensis,Lembosia brigadeirensis,Leptosphaeria urticae,Lonicericola hyaloseptispora,Lophiotrema mucilaginosis,Marasmiellus bicoloripes,Marasmius indojasminodorus,Micropeltis phetchaburiensis,Mucor orantomantidis,Murilentithecium lonicerae,Neobambusicola brunnea,Neoeutypella baoshanensis,Neoroussoella heveae,Neosetophoma lonicerae,Ophiobolus malleolus,Parabambusicola thysanolaenae,Paratrimmatostroma kunmingensis,Parazalerion indica,Penicillium dokdoense,Peroneutypa mangrovei,Phaeosphaeria cycadis,Phanerochaete australosanguinea,Plectosphaerella kunmingensis,Plenodomus artemisiae,P.lijiangensis,Proliferophorum thailandicum,Pseudoastrosphaeriellopsis kaveriana,Pseudohelicomyces menglunicus,Pseudoplagiostoma mangiferae,Robillarda mangiferae,Roussoella elaeicola,Russula choptae,R.uttarakhandia,Septomelanconiella thailandica,Spencermartinsia acericola,Sphaerellopsis isthmospora,Thozetella lithocarpi,Trechispora echinospora,Tremellochaete atlantica,Trichoderma koreanum,T.pinicola,T.rugulosum,Velebitea chrysotexta,Vicosamyces venturisporus,Wojnowiciella kunmingensis and Zopfiella indica.Three new combinations are Baorangia rufomaculata,Lanmaoa pallidorosea and Wojnowiciella rosicola.The reference specimens of Canalisporium kenyense and Tamsiniella labiosa are designated.The epitype of Sarcopeziza sicula is re-circumscribed based on cyto-and histochemical analyses.The sexual-asexual morph connection of Plenodomus sinensis is reported from ferns and Cirsium for the first time.In addition,the new host records and country records are Amanita altipes,A.melleialba,Amarenomyces dactylidis,Chaetosphaeria panamensis,Coniella vitis,Coprinopsis kubickae,Dothiorella sarmentorum,Leptobacillium leptobactrum var.calidus,Muyocopron lithocarpi,Neoroussoella solani,Periconia cortaderiae,Phragmocamarosporium hederae,Sphaerellopsis paraphysata and Sphaeropsis eucalypticola.

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.

Fungal diversity notes 603–708: taxonomic and phylogenetic notes on genera and species

This is the sixth in a series of papers where we bring collaborating mycologists together to produce a set of notes of several taxa of fungi.In this study we introduce a new family Fuscostagonosporaceae in Dothideomycetes.We also introduce the new ascomycete genera Acericola,Castellaniomyces,Dictyosporina and Longitudinalis and new species Acericola italica,Alternariaster trigonosporus,Amarenomyces dactylidis,Angustimassarina coryli,Astrocystis bambusicola,Castellaniomyces rosae,Chaetothyrina artocarpi,Chlamydotubeufia krabiensis,Colletotrichum lauri,Collodiscula chiangraiensis,Curvularia palmicola,Cytospora mali-sylvestris,Dictyocheirospora cheirospora,Dictyosporina ferruginea,Dothiora coronillae,Dothiora spartii,Dyfrolomyces phetchaburiensis,Epicoccum cedri,Epicoccum pruni,Fasciatispora calami,Fuscostagonospora cytisi,Grandibotrys hyalinus,Hermatomyces nabanheensis,Hongkongmyces thailandica,Hysterium rhizophorae,Jahnula guttulaspora,Kirschsteiniothelia rostrata,Koorchalomella salmonispora,Longitudinalis nabanheensis,Lophium zalerioides,Magnibotryascoma mali,Meliola clerodendri-infortunati,Microthyrium chinense,Neodidymelliopsis moricola,Neophaeocryptopus spartii,Nigrograna thymi,Ophiocordyceps cossidarum,Ophiocordyceps issidarum,Ophiosimulans plantaginis,Otidea pruinosa,Otidea stipitata,Paucispora kunmingense,Phaeoisaria microspora,Pleurothecium floriforme,Poaceascoma halophila,Periconia aquatica,Periconia submersa,Phaeosphaeria acaciae,Phaeopoacea muriformis,Pseudopithomyces kunmingnensis,Ramgea ozimecii,Sardiniella celtidis,Seimatosporium italicum,Setoseptoria scirpi,Torula gaodangensis and Vamsapriya breviconidiophora.We also provide an amended account of Rhytidhysteron to include apothecial ascomata and a J?hymenium.The type species of Ascotrichella hawksworthii(Xylariales genera incertae sedis),Biciliopsis leptogiicola(Sordariomycetes genera incertae sedis),Brooksia tropicalis(Micropeltidaceae),Bryochiton monascus(Teratosphaeriaceae),Bryomyces scapaniae(Pseudoperisporiaceae),Buelliella minimula(Dothideomycetes genera incertae sedis),Carinispora nypae(Pseudoastrosphaeriellaceae),Cocciscia hammeri(Verrucariaceae),Endoxylina astroidea(Diatrypaceae),Exserohilum turcicum(Pleosporaceae),Immotthia hypoxylon(Roussoellaceae),Licopolia franciscana(Vizellaceae),Murispora rubicunda(Amniculicolaceae)and Doratospora guianensis(synonymized under Rizalia guianensis,Trichosphaeriaceae)were reexamined and descriptions,illustrations and discussion on their familial placement are given based on phylogeny and morphological data.New host records or new country reports are provided for Chlamydotubeufia huaikangplaensis,Colletotrichum fioriniae,Diaporthe subclavata,Diatrypella vulgaris,Immersidiscosia eucalypti,Leptoxyphium glochidion,Stemphylium vesicarium,Tetraploa yakushimensis and Xepicula leucotricha.Diaporthe baccae is synonymized under Diaporthe rhusicola.A reference specimen is provided for Periconia minutissima.Updated phylogenetic trees are provided for most families and genera.We introduce the new basidiomycete species Agaricus purpurlesquameus,Agaricus rufusfibrillosus,Lactifluus holophyllus,Lactifluus luteolamellatus,Lactifluus pseudohygrophoroides,Russula benwooii,Russula hypofragilis,Russula obscurozelleri,Russula parapallens,Russula phoenicea,Russula pseudopelargonia,Russula pseudotsugarum,Russula rhodocephala,Russula salishensis,Steccherinum amapaense,Tephrocybella constrictospora,Tyromyces amazonicus and Tyromyces angulatus and provide updated trees to the genera.We also introduce Mortierella formicae in Mortierellales,Mucoromycota and provide an updated phylogenetic tree.