Freshwater fungi from karst landscapes in China and Thailand

Freshwater fungi comprises a highly diverse group of organisms occurring in freshwater habitats throughout the world.Dur-ing a survey of freshwater fungi on submerged wood in streams and lakes,a wide range of sexual and asexual species were collected mainly from karst regions in China and Thailand.Phylogenetic inferences using partial gene regions of LSU,ITS,SSU,TEF1α,and RPB2 sequences revealed that most of these fungi belonged to Dothideomycetes and Sordariomycetes and a few were related to Eurotiomycetes.Based on the morphology and multi-gene phylogeny,we introduce four new genera,viz.Aquabispora,Neocirrenalia,Ocellisimilis and Uvarisporella,and 47 new species,viz.Acrodictys chishuiensis,A.effusa,A.pyriformis,Actinocladium aquaticum,Annulatascus tratensis,Aquabispora setosa,Aqualignicola setosa,Aquimassario-sphaeria vermiformis,Ceratosphaeria flava,Chaetosphaeria polygonalis,Conlarium muriforme,Digitodesmium chishuiense,Ellisembia aquirostrata,Fuscosporella atrobrunnea,Halobyssothecium aquifusiforme,H.caohaiense,Hongkongmyces aquisetosus,Kirschsteiniothelia dushanensis,Monilochaetes alsophilae,Mycoenterolobium macrosporum,Myrmecridium splendidum,Neohelicascus griseoflavus,Neohelicomyces denticulatus,Neohelicosporium fluviatile,Neokalmusia aquib-runnea,Neomassariosphaeria aquimucosa,Neomyrmecridium naviculare,Neospadicoides biseptata,Ocellisimilis clavata,Ophioceras thailandense,Paragaeumannomyces aquaticus,Phialoturbella aquilunata,Pleurohelicosporium hyalinum,Pseudodactylaria denticulata,P.longidenticulata,P.uniseptata,Pseudohalonectria aurantiaca,Rhamphoriopsis aquimi-crospora,Setoseptoria bambusae,Shrungabeeja fluviatilis,Sporidesmium tratense,S.versicolor,Sporoschisma atroviride,Stanjehughesia aquatica,Thysanorea amniculi,Uvarisporella aquatica and Xylolentia aseptata,with an illustrated account,discussion of their taxonomic placement and comparison with morphological similar taxa.Seven new combinations are introduced,viz.Aquabispora grandispora(≡Boerlagiomyces grandisporus),A.websteri(≡Boerlagiomyces websteri),Cer-atosphaeria suthepensis(≡Pseudohalonectria suthepensis),Gamsomyces aquaticus(≡Pseudobactrodesmium aquaticum),G.malabaricus(≡Gangliostilbe malabarica),Neocirrenalia nigrospora(≡Cirrenalia nigrospora),and Rhamphoriopsis glauca(≡Chloridium glaucum).Ten new geographical records are reported in China and Thailand and nine species are first reported from freshwater habitats.Reference specimens are provided for Diplocladiella scalaroides and Neocirrenalia nigrospora(≡Cirrenalia nigrospora).Systematic placement of the previously introduced genera Actinocladium,Aqualigni-cola,and Diplocladiella is first elucidated based on the reference specimens and new collections.Species recollected from China and Thailand are also described and illustrated.The overall trees of freshwater Dothideomycetes and Sordariomycetes collected in this study are provided respectively and genera or family/order trees are constructed for selected taxa.

Refined families of Dothideomycetes:orders and families incertae sedis in Dothideomycetes

Numerous new taxa and classifications of Dothideomycetes have been published following the last monograph of families of Dothideomycetes in 2013.A recent publication by Honsanan et al.in 2020 expanded information of families in Dothideo-mycetidae and Pleosporomycetidae with modern classifications.In this paper,we provide a refined updated document on orders and families incertae sedis of Dothideomycetes.Each family is provided with an updated description,notes,including figures to represent the morphology,a list of accepted genera,and economic and ecological significances.We also provide phylogenetic trees for each order.In this study,31 orders which consist 50 families are assigned as orders incertae sedis in Dothideomycetes,and 41 families are treated as families incertae sedis due to lack of molecular or morphological evidence.The new order,Catinellales,and four new families,Catinellaceae,Morenoinaceae Neobuelliellaceae and Thyrinulaceae are introduced.Seven genera(Neobuelliella,Pseudomicrothyrium,Flagellostrigula,Swinscowia,Macroconstrictolumina,Pseudobogoriella,and Schummia)are introduced.Seven new species(Acrospermum urticae,Bogoriella complexoluminata,Dothiorella ostryae,Dyfrolomyces distoseptatus,Macroconstrictolumina megalateralis,Patellaria microspora,and Pseu-domicrothyrium thailandicum)are introduced base on morphology and phylogeny,together with two new records/reports and five new collections from different families.Ninety new combinations are also provided in this paper.

Freshwater fungal number

A comprehensive account of fungal classification from freshwater habitats is outlined and discussed in the present review based on literature of biodiversity studies and recent morpho-phylogenetic analyses.A total of 3,870 freshwater fungal species are listed with additional details on the isolation source,habitat,geographical distribution,and molecular data.The Ascomycota(2,968 species,1,018 genera)dominated the freshwater fungal taxa wherein Sordariomycetes(823 species,298 genera)had the largest number,followed by Dothideomycetes(677 species,229 genera),Eurotiomycetes(276 species,49 genera),and Leotiomycetes(260 species,83 genera).Other phyla included in the updated classification of freshwater fungi are:Chytridiomycota(333 species,97 genera),Rozellomycota(221 species,105 genera),Basidiomycota(218 species,100 genera),Blastocladiomycota(47 species,10 genera),Monoblepharomycota(29 species,6 genera),Mucoromycota(19 spe-cies,10 genera),Aphelidiomycota(15 species,3 genera),Entomophthoromycota(6 species,4 genera),Mortierellomycota(5 species,3 genera),Olpidiomycota(4 species,1 genus),Zoopagomycota(3 species,2 genera),and Sanchytriomycota(2 species,2 genera).The freshwater fungi belong to 1,361 genera,386 families and 145 orders.The Pleosporales and Laboulbeniaceae are the largest freshwater fungal order and family comprised of 391 and 185 species,respectively.The most speciose genera are Chitonomyces(87,Laboulbeniomycetes),Verrucaria(50,Eurotiomycetes),Rhizophydium(52,Rhizophydiomycetes),Penicillium(47,Eurotiomycetes),and Candida(42,Saccharomycetes).

The numbers of fungi:contributions from traditional taxonomic studies and challenges of metabarcoding

The global diversity of fungi has been estimated using several different approaches.There is somewhere between 2–11 million estimated species,but the number of formally described taxa is around 150,000,a tiny fraction of the total.In this paper,we examine 12 ascomycete genera as case studies to establish trends in fungal species descriptions,and introduce new species in each genus.To highlight the importance of traditional morpho-molecular methods in publishing new species,we introduce novel taxa in 12 genera that are considered to have low species discovery.We discuss whether the species are likely to be rare or due to a lack of extensive sampling and classification.The genera are Apiospora,Bambusicola,Beltrania,Capronia,Distoseptispora,Endocalyx,Neocatenulostroma,Neodeightonia,Paraconiothyrium,Peroneutypa,Phaeoacremonium and Vanakripa.We discuss host-specificity in selected genera and compare the number of species epithets in each genus with the number of ITS(barcode)sequences deposited in GenBank and UNITE.We furthermore discuss the relationship between the divergence times of these genera with those of their hosts.We hypothesize whether there might be more species in these genera and discuss hosts and habitats that should be investigated for novel species discovery.

Evolution of freshwater Diaporthomycetidae(Sordariomycetes)provides evidence for five new orders and six new families

Cancellidium is a remarkable fungal genus which has been collected from wood submerged in freshwater and has unique conidia that are important in dispersal in running streams.With such a remarkable morphology,one would have expected it to be a distinct family or order.However,due to the dearth of molecular evidence in related taxa,this genus has previ-ously been placed in the order Hypocreales,subclass Hypocreomycetidae of Sordariomycetes.In this study,we made three new collections of this remarkable aquatic genus from streams in China and Thailand,isolated them into culture,extracted DNA and carried out multigene phylogenetic analysis and divergence time estimation which placed the genus in Diaporthomycetidae.This is one of the seven subclasses of Sordariomycetes and contains 30 lineages that are only known from freshwater.The subclass is therefore of interest when considering the evolution of freshwater fungi.Several lineages of Diaporthomycetidae are morphologically unique and taxa cluster with strong support,but have weak support at the base of the trees.The phylogenetic and MCC trees generated in this study indicate that Aquapteridospora,Barbatosphaeriaceae,Bullimyces,Cancellidium,Ceratolenta,Conlarium,Phialemoniopsis,Pseudostanjehughesia and Rhamphoriaceae are distinct genera/families that evolved in the family/order time frame.The new orders Barbatosphaeriales(177 MYA),Cancellidiales(137 MYA),Ceratolentales(147 MYA),Conlariales(138 MYA)and Rhamphoriales(133 MYA)and six new families,Aquapteridosporaceae(110 MYA),Cancellidiaceae(137 MYA),Ceratolentaceae(81 MYA),Bullimycetaceae(81 MYA),Phialemoniopsaceae(59 MYA),and Pseudostanjehughesiaceae(111 MYA)are introduced with evidence from phylogenies,divergence estimates and distinct morphologies.Of these families,Aquapteridosporaceae,Cancellidiaceae,Bullimycetaceae,and Pseudostanjehughesiaceae are only known from freshwater.

Fungal diversity notes 1387-1511:taxonomic and phylogenetic contributions on genera and species of fungal taxa

This article is the 13th contribution in the Fungal Diversity Notes series,wherein 125 taxa from four phyla,ten classes,31 orders,69 families,92 genera and three genera incertae sedis are treated,demonstrating worldwide and geographic distri-bution.Fungal taxa described and illustrated in the present study include three new genera,69 new species,one new com-bination,one reference specimen and 51 new records on new hosts and new geographical distributions.Three new genera,Cylindrotorula(Torulaceae),Scolecoleotia(Leotiales genus incertae sedis)and Xenovaginatispora(Lindomycetaceae)are introduced based on distinct phylogenetic lineages and unique morphologies.Newly described species are Aspergillus lan-naensis,Cercophora dulciaquae,Cladophialophora aquatica,Coprinellus punjabensis,Cortinarius alutarius,C.mammil-latus,C.quercoflocculosus,Coryneum fagi,Cruentomycena uttarakhandina,Cryptocoryneum rosae,Cyathus uniperidiolus,Cylindrotorula indica,Diaporthe chamaeropicola,Didymella azollae,Diplodia alanphillipsii,Dothiora coronicola,Efibula rodriguezarmasiae,Erysiphe salicicola,Fusarium queenslandicum,Geastrum gorgonicum,G.hansagiense,Helicosporium sexualis,Helminthosporium chiangraiensis,Hongkongmyces kokensis,Hydrophilomyces hydraenae,Hygrocybe boertmannii,Hyphoderma australosetigerum,Hyphodontia yunnanensis,Khaleijomyces umikazeana,Laboulbenia divisa,Laboulbenia triarthronis,Laccaria populina,Lactarius pallidozonarius,Lepidosphaeria strobelii,Longipedicellata megafusiformis,Lophiotrema lincangensis,Marasmius benghalensis,M.jinfoshanensis,M.subtropicus,Mariannaea camelliae,Mel-anographium smilaxii,Microbotryum polycnemoides,Mimeomyces digitatus,Minutisphaera thailandensis,Mortierella solitaria,Mucor harpali,Nigrograna jinghongensis,Odontia huanrenensis,O.parvispina,Paraconiothyrium ajrekarii,Par-afuscosporella niloticus,Phaeocytostroma yomensis,Phaeoisaria synnematicus,Phanerochaete hainanensis,Pleopunctum thailandicum,Pleurotheciella dimorphospora,Pseudochaetosphaeronema chiangraiense,Pseudodactylaria albicolonia,Rhexoacrodictys nigrospora,Russula paravioleipes,Scolecoleotia eriocamporesi,Seriascoma honghense,Synandromyces makranczyi,Thyridaria aureobrunnea,Torula lancangjiangensis,Tubeufia longihelicospora,Wicklowia fusiformispora,Xenovaginatispora phichaiensis and Xylaria apiospora.One new combination,Pseudobactrodesmium stilboideus is pro-posed.A reference specimen of Comoclathris permunda is designated.New host or distribution records are provided for Acrocalymma fici,Aliquandostipite khaoyaiensis,Camarosporidiella laburni,Canalisporium caribense,Chaetoscutula juniperi,Chlorophyllum demangei,C.globosum,C.hortense,Cladophialophora abundans,Dendryphion hydei,Diaporthe foeniculina,D.pseudophoenicicola,D.pyracanthae,Dictyosporium pandanicola,Dyfrolomyces distoseptatus,Ernakula-mia tanakae,Eutypa flavovirens,E.lata,Favolus septatus,Fusarium atrovinosum,F.clavum,Helicosporium luteosporum,Hermatomyces nabanheensis,Hermatomyces sphaericoides,Longipedicellata aquatica,Lophiostoma caudata,L.clematidis-vitalbae,Lophiotrema hydei,L.neoarundinaria,Marasmiellus palmivorus,Megacapitula villosa,Micropsalliota globocys-tis,M.gracilis,Montagnula thailandica,Neohelicosporium irregulare,N.parisporum,Paradictyoarthrinium diffractum,Phaeoisaria aquatica,Poaceascoma taiwanense,Saproamanita manicata,Spegazzinia camelliae,Submersispora variabi-lis,Thyronectria caudata,T.mackenziei,Tubeufia chiangmaiensis,T.roseohelicospora,Vaginatispora nypae,Wicklowia submersa,Xanthagaricus necopinatus and Xylaria haemorrhoidalis.The data presented herein are based on morphological examination of fresh specimens,coupled with analysis of phylogenetic sequence data to better integrate taxa into appropriate taxonomic ranks and infer their evolutionary relationships.

Fungal diversity notes 1036-1150:taxonomic and phylogenetic contributions on genera and species of fungal taxa

This article is the tenth series of the Fungal Diversity Notes,where 114 taxa distributed in three phyla,ten classes,30 orders and 53 families are described and illustrated.Taxa described in the present study include one new family(viz.Pseudoberkleasmiaceae in Dothideomycetes),five new genera(Caatingomyces,Cryptoschizotrema,Neoacladium,Paramassaria and Trochilispora)and 71 new species,(viz.Acrogenospora thailandica,Amniculicola aquatica,A.guttulata,Angustimassarina sylvatica,Blackwellomyces lateris,Boubovia gelatinosa,Buellia viridula,Caatingomyces brasiliensis,Calophoma humuli,Camarosporidiella mori,Canalisporium dehongense,Cantharellus brunneopallidus,C.griseotinctus,Castanediella meliponae,Coprinopsis psammophila,Cordyceps succavus,Cortinarius minusculus,C.subscotoides,Diaporthe italiana,D.rumicicola,Diatrypella delonicis,Dictyocheirospora aquadulcis,D.taiwanense,Digitodesmium chiangmaiense,Distoseptispora dehongensis,D.palmarum,Dothiorella styphnolobii,Ellisembia aurea,Falciformispora aquatic,Fomitiporia carpinea,F.lagerstroemiae,Grammothele aurantiaca,G.micropora,Hermatomyces bauhiniae,Jahnula queenslandica,Kamalomyces mangrovei,Lecidella yunnanensis,Micarea squamulosa,Muriphaeosphaeria angustifoliae,Neoacladium indicum,Neodidymelliopsis sambuci,Neosetophoma miscanthi,N.salicis,Nodulosphaeria aquilegiae,N.thalictri,Paramassaria samaneae,Penicillium circulare,P.geumsanense,P.mali-pumilae,P.psychrotrophicum,P.wandoense,Phaeoisaria siamensis,Phaeopoacea asparagicola,Phaeosphaeria penniseti,Plectocarpon galapagoense,Porina sorediata,Pseudoberkleasmium chiangmaiense,Pyrenochaetopsis sinensis,Rhizophydium koreanum,Russula prasina,Sporoschisma chiangraiense,Stigmatomyces chamaemyiae,S.cocksii,S.papei,S.tschirnhausii,S.vikhrevii,Thysanorea uniseptata,Torula breviconidiophora,T.polyseptata,Trochilispora schefflerae and Vaginatispora palmae).Further,twelve new combinations(viz.Cryptoschizotrema cryptotrema,Prolixandromyces australi,P.elongatus,P.falcatus,P.longispinae,P.microveliae,P.neoalardi,P.polhemorum,P.protuberans,P.pseudoveliae,P.tenuistipitis and P.umbonatus),an epitype is chosen for Cantharellus goossensiae,a reference specimen for Acrogenospora sphaerocephala and new synonym Prolixandromyces are designated.Twenty-four new records on new hosts and new geographical distributions are also reported(i.e.Acrostalagmus annulatus,Cantharellus goossensiae,Coprinopsis villosa,Dothiorella plurivora,Dothiorella rhamni,Dothiorella symphoricarposicola,Dictyocheirospora rotunda,Fasciatispora arengae,Grammothele brasiliensis,Lasiodiplodia iraniensis,Lembosia xyliae,Morenoina palmicola,Murispora cicognanii,Neodidymelliopsis farokhinejadii,Neolinocarpon rachidis,Nothophoma quercina,Peroneutypa scoparia,Pestalotiopsis aggestorum,Pilidium concavum,Plagiostoma salicellum,Protofenestella ulmi,Sarocladium kiliense,Tetraploa nagasakiensis and Vaginatispora armatispora).

Fungal diversity notes 1277-1386:taxonomic and phylogenetic contributions to fungal taxa

This is the twelfth contribution to the Fungal Diversity Notes series on fungal taxonomy,based on materials collected from many countries which were examined and described using the methods of morphology,anatomy,and strain culture,combined with DNA sequence analyses.110 taxa are described and illustrated,including five new genera,92 new species,eight new combinations and other taxonomic contributions(one new sequenced species,one new host and three new records)which are accommodated in 40 families and 1 incertae sedis in Dothideomycetes.The new genera are Amyloceraceomyces,Catenuliconidia,Hansenopezia,Ionopezia and Magnopulchromyces.The new species are Amyloceraceomyces angustisporus,Amylocorticium ellipsosporum,Arthrinium sorghi,Catenuliconidia uniseptata,Clavulina sphaeropedunculata,Colletotrichum parthenocissicola,Coniothyrium triseptatum,Cortinarius indorusseus,C.paurigarhwalensis,C.sinensis,C.subsanguineus,C.xiaojinensis,Diaporthe pimpinel-lae,Dictyosporella guizhouensis,Diplodia torilicola,Fuscoporia marquesiana,F.semiarida,Hansenopezia decora,Helicoarcta-tus thailandicus,Hirsutella hongheensis,Humidicutis brunneovinacea,Lentaria gossypina,L.variabilis,Lycoperdon lahorense,L.pseudocurtisii,Magnopulchromyces scorpiophorus,Moelleriella gracilispora,Neodevriesia manglicola,Neodidymelliopsis salvia,N.urticae,Neoroussoella magnoliae,Neottiella gigaspora,Ophiosphaerella chiangraiensis,Phaeotremella yunnanensis,Podosphaera yulii,Rigidoporus juniperinus,Rhodofomitopsis pseudofeei,Russula benghalensis,Scleroramularia vermispora,Scytinopogon minisporus,Sporormurispora paulsenii,Thaxteriellopsis obliqus,Tomentella asiae-orientalis,T.atrobadia,T.atrocastanea,T.aureomarginata,T.brevis,T.brunneoflava,T.brunneogrisea,T.capitatocystidiata,T.changbaiensis,T.citri-nocystidiata,T.coffeae,T.conclusa,T.cystidiata,T.dimidiata,T.duplexa,T.efibulata,T.efibulis,T.farinosa,T.flavidobadia,T.fuscocrustosa,T.fuscofarinosa,T.fuscogranulosa,T.fuscopelliculosa,T.globospora,T.gloeocystidiata,T.griseocastanea,T.griseofusca,T.griseomarginata,T.inconspicua,T.incrustata,T.interrupta,T.liaoningensis,T.longiaculeifera,T.longiechinuli,T.megaspora,T.olivacea,T.olivaceobrunnea,T.pallidobrunnea,T.pallidomarginata,T.parvispora,T.pertenuis,T.qingyuanensis,T.segregata,T.separata,T.stipitata,T.storea,Trichoderma ceratophylletum,Tyromyces minutulus,Umbelopsis heterosporus and Xylolentia reniformis.The new combinations are Antrodiella descendena,Chloridium macrocladum,Hansenopezia retrocurvata,Rhodofomitopsis monomitica,Rh.oleracea,Fuscoporia licnoides,F.scruposa and Ionopezia gerardii.A new sequenced species(Graphis supracola),one new host(Aplosporella prunicola)and three new geographical records(Golovinomyces monardae,Paradictyoarthrinium diffractum and Prosthemium betulinum),are reported.

Freshwater Sordariomycetes

Sordariomycetes is one of the largest classes of Ascomycota that comprises a highly diverse range of fungi mainly characterized by perithecial ascomata and inoperculate unitunicate asci.Freshwater Sordariomycetes play an important role in ecosystems and some of them have the potential to produce bioactive compounds.This study documents and reviews the freshwater Sordariomycetes,which is one of the largest and important groups of fungi in aquatic habitats.Based on evidence from DNA sequence data and morphology,we introduce a new order Distoseptisporales,two new families,viz.Ceratosphaeriaceae and Triadelphiaceae,three new genera,viz.Aquafiliformis,Dematiosporium and Neospadicoides,47 new species,viz.Acrodictys fluminicola,Aquafiliformis lignicola,Aquapteridospora fusiformis,Arthrinium aquaticum,Ascosacculus fusiformis,Atractospora aquatica,Barbatosphaeria lignicola,Ceratosphaeria aquatica,C.lignicola,Chaetosphaeria aquatica,Ch.catenulata,Ch.guttulata,Ch.submersa,Codinaea yunnanensis,Conioscypha aquatica,C.submersa,Cordana aquatica,C.lignicola,Cosmospora aquatica,Cylindrotrichum submersum,Dematiosporium aquaticum,Dictyochaeta cangshanensis,D.ellipsoidea,D.lignicola,D.submersa,Distoseptispora appendiculata,D.lignicola,D.neorostrata,D.obclavata,Hypoxylon lignicola,Lepteutypa aquatica,Myrmecridium aquaticum,Neospadicoides aquatica,N.lignicola,N.yunnanensis,Ophioceras submersum,Peroneutypa lignicola,Phaeoisaria filiformis,Pseudostanjehughesia lignicola,Rhodoveronaea aquatica,Seiridium aquaticum,Sporidesmiella aquatica,Sporidesmium lageniforme,S.lignicola,Tainosphaeria lunata,T.obclavata,Wongia aquatica,two new combinations,viz.Acrodictys aquatica,Cylindrotrichum aquaticum,and 9 new records,viz.Chaetomium globosum,Chaetosphaeria cubensis,Ch.myriocarpa,Cordana abramovii,Co.terrestris,Cuspidatispora xiphiago,Sporidesmiella hyalosperma,Stachybotrys chartarum,S.chlorohalonata.A comprehensive classification of the freshwater Sordariomycetes is presented based on updated literature.Phylogenetic inferences based on DNA sequence analyses of a combined LSU,SSU,RPB2 and TEF1a dataset comprising species of freshwater Sordariomycetes are provided.Detailed information including their habitats distribution,diversity,holotype,specimens collected and classification are provided.