Families of Sordariomycetes

Sordariomycetes is one of the largest classes of Ascomycota that comprises a highly diverse range of fungi characterized mainly by perithecial ascomata and inoperculate unitunicate asci.The class includes many important plant pathogens,as well as endophytes,saprobes,epiphytes,coprophilous and fungicolous,lichenized or lichenicolous taxa.They occur in terrestrial,freshwater and marine habitats worldwide.This paper reviews the 107 families of the class Sordariomycetes and provides a modified backbone tree based on phylogenetic analysis of four combined loci,with a maximum five representative taxa from each family,where available.This paper brings together for the first time,since Barrs’1990 Prodromus,descriptions,notes on the history,and plates or illustrations of type or representative taxa of each family,a list of accepted genera,including asexual genera and a key to these taxa of Sordariomycetes.Delineation of taxa is supported where possible by molecular data.The outline is based on literature to the end of 2015 and the Sordariomycetes now comprises six subclasses,32 orders,105 families and 1331 genera.The family Obryzaceae and Pleurotremataceae are excluded from the class.

An updated phylogeny of Sordariomycetes based on phylogenetic and molecular clock evidence

The previous phylogenies of Sordariomycetes by M.E.Barr,O.E.Eriksson and D.L.Hawksworth,and T.Lumbsch and S.Huhndorf,were mainly based on morphology and thus were somewhat subjective.Later outlines by T.Lumbsch and S.Huhndorf,and Maharachchikumbura and co-authors,took into account phylogenetic evidence.However,even these phylogenetic driven arrangements for Sordariomycetes,were somewhat subjective,as the arrangements in trees depended on many variables,such as number of taxa,different gene regions and methods used in the analyses.What is needed is extra evidence to help standardize ranking in the fungi.Estimation of divergence times using molecular clock methods has been proposed for providing additional rational for higher ranking of taxa.Thus,in Sordariomycetes,a divergence period(i.e.200–300 MYA)can be used as criteria to judge when a group of related taxa evolved and what rank they should be given.In this paper,we provide an updated classification of accepted subclasses,orders of Sordariomycetes and use divergence times to provide additional evidence to stabilize ranking of taxa in the class.We point out and discuss discrepancies where the phylogenetic tree conflicts with the molecular clock.

Towards a natural classification of Annulatascaceae-like taxa:introducing Atractosporales ord.nov.and six new families

Species with relatively small,membraneous,black ascomata,with or without long necks,unitunicate,cylindrical asci with apical rings and fusiform,hyaline ascospores with or without mucilaginous sheaths are common in freshwater habitats in tropical and temperate regions.Many of these taxa have originally been recorded as Annulatascaceae-like taxa.Twenty genera have been included in the family Annulatascaceae,mostly based on morphological characters,while molecular work and phylogenetic analyses are lacking for many genera.In this study,nine new Annulatascaceae-like taxa collected from Thailand were morphologically examined.Pure cultures obtained from single ascospores were used in molecular studies.The nine new strains and several other strains of Annulatascaceae-like Sordariomycetes species were used to establish phylogenetic and evolution relationships among the taxa,based on combined LSU,SSU,ITS and RPB2 sequence data.Phylogenetic analyses provide evidence to introduce one new order and six new families,to accommodate taxa excluded from Annulatascaceae sensu stricto.A new order Atractosporales is established based on the molecular study,including three new introduced families Conlariaceae,Pseudoproboscisporaceae and Atractosporaceae.Conlariaceae is introduced for the genus Conlarium which comprises two species,Conlarium duplumascosporun and a new Hyphomycetous asexual morph taxon Conlarium aquaticum which has subglobose or irregular,brown,clathrate,muriform conidia.Pseudoproboscisporaceae includes Pseudoproboscispora and Diluviicola,while Atractosporaceae includes the genera Rubellisphaeria and Atractospora.Barbatosphaeria,Xylomelasma and Ceratostomella form a distinct stable lineage which is introduced as a new family Barbatosphaeriaceae in Diaporthomycetidae families incertae sedis.A new family Lentomitellaceae is introduced in Diaporthomycetidae families incertae sedis,to accommodate the genus Lentomitella.Woswasiaceae is introduced to accommodate Woswasia,Xylochrysis and Cyanoannulus in Diaporthomycetidae families incertae sedis.Three new species of Fluminicola viz.F.saprophytica,F.thailandensis and F.aquatica are introduced.A new sexual morph,Dictyosporella thailandensis,is reported and Dictyosporella is excluded from Annulatascaceae and placed in Diaporthomycetidae genera incertae sedis.The first sexual morph of Sporidesmium,S.thailandense is also described.The new species Atractospora thailandensis,Diluviicola aquatica and Pseudoproboscispora thailandensis are also introduced.Platytrachelon is added to Papulosaceae based on phylogenetic analysis and morphological characters.Aquaticola,Fusoidispora and Pseudoannulatascus are excluded from Annulatascaceae and placed in Diaporthomycetidae genera incertae sedis.Mirannulata is accommodated in Sordariomycetes,genera incertae sedis.

The numbers of fungi: is the descriptive curve flattening?

The recent realistic estimate of fungal numbers which used various algorithms was between 2.2 and 3.8 million.There are nearly 100,000 accepted species of Fungi and fungus-like taxa,which is between 2.6 and 4.5%of the estimated species.Several forums such as Botanica Marina series,Fungal Diversity notes,Fungal Biodiversity Profiles,Fungal Systematics and Evolution-New and Interesting Fungi,Mycosphere notes and Fungal Planet have enhanced the introduction of new taxa and nearly 2000 species have been introduced in these publications in the last decade.The need to define a fungal species more accurately has been recognized,but there is much research needed before this can be better clarified.We address the evidence that is needed to estimate the numbers of fungi and address the various advances that have been made towards its understanding.Some genera are barely known,whereas some plant pathogens comprise numerous species complexes and numbers are steadily increasing.In this paper,we examine ten genera as case studies to establish trends in fungal description and introduce new species in each genus.The genera are the ascomycetes Colletotrichum and Pestalotiopsis(with many species or complexes),Atrocalyx,Dothiora,Lignosphaeria,Okeanomyces,Rhamphoriopsis,Thozetella,Thyrostroma(rela-tively poorly studied genera)and the basidiomycete genus Lepiota.We provide examples where knowledge is incomplete or lacking and suggest areas needing further research.These include(1)the need to establish what is a species,(2)the need to establish how host-specific fungi are,not in highly disturbed urban areas,but in pristine or relatively undisturbed forests,and(3)the need to establish if species in different continents,islands,countries or regions are different,or if the same fungi occur worldwide?Finally,we conclude whether we are anywhere near to flattening the curve in new species description.

Uncovering the hidden taxonomic diversity of fungi in Oman

Hot desert regions are undoubtedly challenging to fungal survival on the Arabian Peninsula.Fungi are,however,recognized as the most stress-resistant organisms among all eukaryotes,which could be a result of the rapid evolution of distinct species.Our current understanding of these microorganisms is derived from studies examining only a fraction of the overall fungal diversity.Therefore,further studies are needed to understand the diversity of fungi in desert regions.This paper highlights the taxonomy of several unusual fungal genera collected in a range of aquatic and terrestrial environments in Oman.These taxa were identified based on phylogenetic analyses of nuclear ribosomal DNA(rDNA)(LSU,SSU and ITS)and protein-coding genes(TEF,RPB2 and TUB),plus morphological comparisons.Phylogenetic analyses,including the presently recognized genera in Patellariales resulted in splitting the order into four clades in Dothideomycetes.The new family Holmiellaceae and new order Holmiellales are introduced to include Holmiella.Species of Homortomycetaceae form a well-supported and distinct clade and raise it to Homortomycetales ordo novus.Omania(Halojulellaceae),Desertiserpentica(Lophiostomata-ceae)and Montanitestudina(Testudinaceae)are described as novel genera in Dothideomycetes.A synnematous hyphomycete with basidiomycetous affinity(Corticiales)was also identified and described as Basidiodesertica.Additionally,an asexual morph was observed for Holmiella junipericola;Patellaria quercus is synonymized under P.atrata,and seven new species are described.

The ranking of fungi:a tribute to David L.Hawksworth on his 70th birthday

The history of assigning ranks to fungi,as well as the relative importance of using divergence time estimates is reviewed.The paper pays tribute to the major mycological players,and especially to David Hawksworth on his 70th birthday and his contribution to fungal ranking in Systema Ascomycetum from 1982 to 1998.Following the conclusion of the latter series,the ranking continued with the Outlines of Ascomycota in 2007 and 2010 and more recently with specific classes in‘Towards an outline of Sordariomycetes’and‘Families of Dothideomycetes’.Earlier classifications based on phenotype were certainly more subjective;however,remarkably many of these old arrangements have stood the test of time.More recently,phylogenetic analyses have provided evidence towards a natural classification,resulting in significant changes in many lineages.The classification arrangements however,are still subjective and dependent on the taxa analysed,resulting in different taxonomic interpretations and schemes,particularly when it comes to ranking.Thus,what have been considered as genera by some,have been introduced as families by others.More recently,estimation of divergence times using molecular clock methods have been used as objective evidence for higher ranking of taxa.A divergence period(i.e.200–300 MYA)can be used as a criterion to infer when a group of related taxa evolved and what rank they should be given.We compiled data on divergence times for various higher ranking taxa in the Kingdom Fungi.The kingdom evolved 1000–1600 MYA(Stenian–Calymmian),while the presently accepted phyla evolved between 358 and 541 MYA(Devonian–Cambrian).Divergence times for subphyla are generally between 358 and 485 MYA(Devonian–Ordovician),those of classes 145–358 MYA(Jurassic–Carboniferous),subclasses 66–358 MYA(Cretaceous–Carboniferous),orders 23–252 MYA(Paleogene–Triassic),families 2.8–145 MYA(Neogene–Cretaceous),and genera 2.8–66 MYA(Neogene–Paleogene).Thus,there are wide discrepancies in the times different taxa diverged.We provide an overview over Ascomycota,showing how application of temporal banding could affect the recognition of higher taxa at certain rank levels.We then use Sordariomycetes as an example where we use divergence times to provide additional evidence to stabilize ranking of taxa below class level.We propose a series of evolutionary periods that could be used as a guide to determine the various higher ranks of fungi:phyla[550 MYA,subphyla 400–550 MYA;classes 300–400 MYA;subclasses 250–300 MYA,orders 150–250 MYA,and families 50–150 MYA.It is proposed that classification schemes and ranking of taxa should,where possible,incorporate a polyphasic approach including phylogeny,phenotype,and estimate of divergence times.