Introduction {#sec1}
============

People spend up to 90 % of their time indoors ([@bib10]).  Fungi present in these indoor environments can produce toxins or carry allergens which cause health hazards.  Therefore it is important to know which fungal species are present indoors.  Several reports are made on the presence of *Microascaceae* in the indoor environment.  The species *Scopulariopsis brevicaulis*, *S. candida*, *S. fusca* (= *S. asperula*), *S. brumptii* (= *Microascus paisii*) and *S. sphaerospora* (= *M. paisii*) are often mentioned as indoor fungi ([@bib28]).  However, in most of the indoor reports of scopulariopsis-like isolates, morphological examination has not been confirmed with molecular studies.  Also the absence of thorough phylogenetic studies in these genera made it difficult to accurately identify the indoor *Microascaceae*.  The first phylogenetic study of scopulariopsis-like species was based on the large subunit 28S nrDNA (LSU, [@bib12]).  Here the potential relationship between asexual and sexually reproducing species was assessed, with a focus on clinically occurring species.  The main microascoid clade, which contained all *Microascus* and *Scopulariopsis* species studied, was divided into seven clades.  Further taxonomic study is suggested to redefine or split the genus *Microascus*.  A study of clinical isolates in Poland confirmed that the LSU sequence alone is insufficient for species delimitation in *Scopulariopsis* ([@bib13]).  A taxonomic study of cheese fungi used the beta-tubulin (*tub2*) and translation elongation factor 1-alpha (*tef1*) gene regions next to LSU to identify their *Scopulariopsis* species ([@bib26]).  The internal transcribed spacer 1 and 2 and intervening 5.8S nrDNA (ITS) gave problems with amplification, and displayed a high variability, which made it not useful for phylogenetic study of their isolates.  Translation elongation factor 1-alpha showed to be the most phylogenetically informative genomic region and was proposed for identifying *Scopulariopsis* species.  A subsequent phylogenetic study on clinical *Microascus* and *Scopulariopsis* species made a combined phylogeny of the LSU and *tef1* gene region ([@bib30]).  They concluded that this combined analysis is useful for the identification of the most common clinically relevant *Scopulariopsis* species.  However, further phylogenetic studies testing more genetic markers and reference strains are suggested, since nine phylogenetic clades in their combined phylogenies could not be properly named.  This follow-up study with the aim to clarify the taxonomy and phylogeny of *Microascus*, *Scopulariopsis* and allied genera was published recently ([@bib29]).  On a large set of clinical and environmental isolates, including ex-type strains of multiple species, a phylogenetic study was conducted based on the ITS, LSU, *tef1* and *tub2* gene regions, in combination with morphological and physiological analyses.  In this polyphasic approach study the genera *Microascus* and *Scopulariopsis* are separated, the genus *Pithoascus* reinstated, and the new genus *Pseudoscopulariopsis* proposed.  Seven new *Microascus* species and one new *Scopulariopsis* species are described, nine new name combinations are introduced, and several species are neotypified ([@bib29]).  A second taxonomic study on a set of clinical and environmental scopulariopsis-like fungi followed soon ([@bib14]).  Here another three new *Microascus* species, one new *Scopulariopsis* species and one new *Pithoascus* species are described, *S. albo-flavescens* is reinstated, *M. trigonosporus* var. * terreus* recombined in *M. terreus*, and the new genus *Fuscoannellis* proposed. 

Although these two recent phylogenetic studies ([@bib14], [@bib29]) make molecular identification of scopulariopsis-like isolates upon species level possible, the involved name changes can cause a lot of confusion.  Commonly mentioned species from the indoor environment, like *S. brumptii* (now *M. paisii*), *S. fusca* (now *S. asperula*) and *S. sphaerospora* (now *M. paisii*), are renamed.  The aim of this project is to molecularly identify the scopulariopsis-like taxa, which occur in the indoor environment.  Simultaneously, a phylogenetic overview of these genera is constructed, and the species habitats are studied.  All available *Microascus* and *Scopulariopsis* isolates from the Westerdijk Fungal Biodiversity Institute culture collection (CBS collection) and working collection of the Applied and Industrial Mycology department (DTO collection) are included in the study.  Species phylogenetic inferences were conducted on sequence data of parts of the ITS, *tub2* and *tef1* gene regions, and a genus phylogenetic inference on the LSU, ITS and *tef1* gene regions.  Phylogenetic clades which contain indoor isolates are highlighted as indoor species.  New species are described, and an overview of the current species and their habitats in the genera *Microascus*, *Scopulariopsis* and *Yunnania* is provided.  Furthermore, a table showing recent name changes and a key to common species of *Scopulariopsis* and scopulariopsis-like genera found in the indoor environment is provided. 

Materials and methods {#sec2}
=====================

Isolates {#sec2.1}
--------

In total 248 isolates were included in this study, comprising of 152 *Microascus* isolates, 88 *Scopulariopsis* isolates, four *Yunnania* isolates, and four out-group isolates.  The isolates were obtained from the culture collection of the Westerdijk Fungal Biodiversity Institute (former CBS-KNAW Fungal Biodiversity Centre), Utrecht, the Netherlands and the working collection of the Applied and Industrial Mycology department (DTO) housed at the Westerdijk Institute ([Table 1](#tbl1){ref-type="table"}).  Isolates from the culture collection of the Westerdijk Institute (CBS collection) have a world-wide distribution and are isolated from a diverse range of substrates.  Isolates from the working collection of DTO are mostly isolated from indoor environments or food, and include swab and air samples mainly from Europe, and house dust samples collected world-wide ([@bib1]).  Freeze-dried strains from the CBS culture collection were revived in 2 mL malt/peptone (50 % / 50 %) and subsequently transferred to oatmeal agar (OA) ([@bib28]).  Strains stored in the liquid nitrogen (CBS collection) or the DTO collection were transferred to OA directly from the −185 °C or −80 °C storage, respectively.  They were cultured for 14 d at 25 °C in the dark.  From eight isolates only their DNA sequences from GenBank were obtained ([Table 1](#tbl1){ref-type="table"}, isolates without a DTO number).Table 1Isolates used in this study and their GenBank accession numbers.  Bold accession numbers were generated in other studies. Table 1NameOld name[1](#tbl1fn1){ref-type="table-fn"}Strain numbers[2](#tbl1fn2){ref-type="table-fn"}Host/SubstrateCountryGenBank accession numberITS*tub2tef1*LSU*Cephalotrichum asperulum***CBS 582.71**^IT^**, DTO 104-B7, ATCC 26885**SoilArgentinaKX923818KX924043KX924027*C. stemonitis***CBS 103.19**^NT^**, DTO 170-B3, MUCL 6960**SeedNetherlandsKX923819**LN850953LN850952***Microascus alveolarisM. trigonosporus*CBS 268.49, DTO 342-D8*Avena sativa*, grainUSAKX923823KX924257KX924047*M. trigonosporus*CBS 269.49, DTO 342-C2*Glycine soja*, seedUSAKX923824KX924258KX924048*M. trigonosporus*CBS 270.49, DTO 345-F7*Hordeum vulgare*, seedUSAKX923825KX924259KX924049*M. trigonosporus*CBS 271.49, DTO 342-D9*Hordeum vulgare*, seedUSAKX923826KX924260KX924050*M. trigonosporus*CBS 272.49, DTO 342-C3*Avena sativa*, leafUSAKX923827KX924261KX924051*M. trigonosporus*CBS 150.64, DTO 342-E2*Allium cepa*, seedUSAKX923828KX924262KX924052*M. trigonosporus*CBS 494.70, DTO 342-C8Marine sedimentNorway**LN850757LN850855LN850903CBS 139501**^T^**, DTO 351-E2, FMR 12252, UTHSC 07-3491**Human, BAL fluidUSAKX923829KX924263KX924053KX924029DTO 223-A7IndoorUruguayKX923830KX924264KX924054*M. appendiculatus* sp.  nov.*M. senegalensis***CBS 594.78**^T^**, DTO 354-C3**Human, skinAlgeria**LN850781LN850878**KX924055**LN850830***M. atrogriseus* nom.  nov.*Masonia grisea***CBS 295.52**^T^**, DTO 103-H6, IFO 6795, IMI 049908, MUCL 9003**Culture contaminantUK**LM652433**KX924265KX924056KX924030CBS 897.68, DTO 356-C3, ATCC 16279, IFO 31245, MUCL 8993Wheat field soilGermany**LM652436LM652649LM652571***S. chartarum*CBS 410.76, DTO 345-B1Burnt soilNetherlandsKX923831KX924266KX924057DTO 139-D7IndoorGermanyKX923832KX924267KX924058DTO 191-C2Indoor horse arenaNetherlandsKX923833KX924268KX924059*M. brunneosporus***CBS 138276**^T^**, DTO 351-D8, UTHSC 06-4312, FMR 12343**Human, BAL fluidUSAKX923834KX924269[**HG380420**](ncbi-n:HG380420){#intref0110}[**HG380497**](ncbi-n:HG380497){#intref0115}*M. chartarusMasonia chartarum***CBS 294.52**^T^**, IMI 049909, MUCL 9001**Wall paperUK**LM652393LM652607**[**HG380386**](ncbi-n:HG380386){#intref0120}[**HG380463**](ncbi-n:HG380463){#intref0125}*M. chinensis***CBS 139628**^T^**, BMU 01837**Human, nailChina**LN850760LN850858LN850906LN850809***M. cinereusM. griseus*^T^CBS 365.65, DTO 104-A3, DTO 104-A4, ATCC 16204, HACC 1252, IMI 113680SoilIndia**LM652399**KX924270KX924060CBS 664.71, DTO 104-B8Human, lungUSAKX923835**LN850860**KX924061CBS 324.72, DTO 342-G2ClayNamibiaKX923836KX924271KX924062**CBS 138709**^NT^**, DTO 351-E1, UTHSC 10-2805, FMR 12217**Human, BAL fluidUSAKX923837KX924272KX924063KX924031*M. cirrosus***CBS 217.31**^T^**, DTO 103-G1***Prunus* sp.,  leafItalyKX923838KX924273KX924064KX924032CBS 277.34, DTO 345-A7, MUCL 9050, MUCL 9055*Vitis vinifera*, rootItalyKX923839KX924274**LM652556***M. longirostris*CBS 267.49, DTO 170-B9, IFO 7029*Sciurus vulgaris*, skinNetherlandsKX923840KX924275KX924065CBS 240.58, DTO 342-E1, IMI 086913Compost soilGermanyKX923841KX924276KX924066CBS 301.61, DTO 345-A8, IMI 086914, NRRL 1689, MUCL 9054UnknownUKKX923842KX924277KX924067CBS 302.61, DTO 345-D1UnknownCanadaKX923843KX924278KX924068CBS 424.62, DTO 345-A9PolyvinylchlorideNetherlandsKX923844KX924279KX924069CBS 541.74, DTO 342-I8Rodent dungUSAKX923845KX924280KX924070CBS 157.92, DTO 342-G6, FRR 4174*Arachis hypogaea*, nutIndonesiaKX923846KX924281KX924071CBS 115860, DTO 345-D4, FMR 8575AirSpainKX923847KX924282KX924072CBS 116405, DTO 345-D6Antique tapestriesPoland**LN850763LN850861LN850909**DTO 342-D4, RGR 84.0007*Helianthus annuus*, oilseedUSAKX923848KX924283KX924073DTO 342-D5, RGR 84.0033UnknownUnknownKX923849KX924284KX924074DTO 342-D7, RGR 84.0051UnknownUnknownKX923850KX924285KX924075*M. cleistocarpus* sp.  nov.**CBS 134638**^T^**, DTO 342-D2, CGMCC 3.15222**Discarded clothChinaKX923851KX924286KX924076KX924033*M. crociS. croci***CBS 158.44**^T^**, DTO 103-H3, IMI 078261, MUCL 9002***Crocus* sp. NetherlandsKX923852KX924287KX924077**LM652508***Masoniella tertia*^T^CBS 296.61, DTO 103-I6, IMI 109550, MUCL 9005AirBrazilKX923853KX924288KX924078*S. chartarum*CBS 522.69, DTO 342-C7Forest soilCanadaKX923854KX924289KX924079DTO 220-I5IndoorIndonesiaKX923855KX924290KX924080DTO 252-D8IndoorGermanyKX923856KX924291KX924081DTO 305-B3IndoorMexicoKX923857KX924292KX924082DTO 305-B5IndoorMexicoKX923858KX924293KX924083*M. expansus***CBS 138127**^T^**, DTO 351-D6, UTHSC 06-4472, FMR 12266**Human, sputumUSAKX923859KX924294KX924084[**HG380492**](ncbi-n:HG380492){#intref0130}*M. fusisporus* sp.  nov.*M. paisii***CBS 896.68**^T^**, DTO 356-C2, ATCC 16278, IFO 31244, MUCL 8989**Wheat-field soilGermany**LM652432LM652645**[**HG380372**](ncbi-n:HG380372){#intref0135}**LN850825***M. gracilisM. cinereus*CBS 126.14, DTO 347-C4, IMI 086916UnknownUnknownKX923860KX924295KX924085*M. cinereus*CBS 195.61, DTO 345-C8, IMI 075542, MUCL 9048SoilUK**LM652416LM652629**[**HG380391**](ncbi-n:HG380391){#intref0140}CBS 300.61, DTO 345-C9, MUCL 9049*Zea mays*, stored seedUSA**LM652417**KX924296**LM652563CBS 369.70**^T^**, DTO 104-B3, IFO 7561**Wheat flourJapanKX923861KX924297KX924086[**HG380467**](ncbi-n:HG380467){#intref0145}CBS 794.91, DTO 345-D2Rice flourAustraliaKX923862KX924298KX924087CBS 156.92, DTO 345-D3*Arachis hypogaea*, nutThailandKX923863KX924299KX924088*M. cinereus*CBS 116059, DTO 345-D5Polyethylene with starchPolandKX923864**LN850863**KX924089CBS 120886, DTO 342-D1*Prunus persica*South AfricaKX923865KX924300KX924090DTO 220-I4IndoorIndonesiaKX923866KX924301KX924091*M. cinereus*DTO 342-D6, RGR 84.0038UnknownUnknownKX923867KX924302KX924092*M. cinereus*DTO 342-G8, RGR 84.0040UnknownUnknownKX923868KX924303KX924093*M. hollandicus* sp.  nov.**CBS 141582**^T^**, DTO 191-C3**Indoor horse arenaNetherlandsKX923869KX924304KX924094KX924034*M. hyalinus***CBS 766.70**^T^**, DTO 170-F2**Cow dungUSAKX923870KX924305**LM652564LM652513**CBS 134639, DTO 342-I9Goat dungChinaKX923871KX924306KX924095*M. intricatus***CBS 138128**^T^**, DTO 351-D7, UTHSC 07-156, FMR 12264**Human BAL fluidUSAKX923872KX924307[**HG380419**](ncbi-n:HG380419){#intref0150}[**HG380496**](ncbi-n:HG380496){#intref0155}DTO 223-A6IndoorMicronesiaKX923873KX924308KX924096*M. longicollis*CBS 752.97, DTO 338-G8*Anacardium occidentale*, nutBrazilKX923874KX924309KX924097KX924035*M. longirostris***CBS 196.61**^NT^**, IMI 086908, MUCL 9058, NRRL 1717**Wasp\'s nestUSA**LM652421LM652634LM652566LM652515**CBS 415.64, IFO 7554SoilJapan**LM652422LM652635LM652567***M. macrosporus*CBS 662.71, DTO 170-F6, NRRL A-8018SoilUSA**LM652423LM652636LM652568LM652517***M. cirrosus*CBS 540.74, DTO 347-D1SoilUSAKX923875KX924310KX924098*M. melanosporus* comb.  nov.*S. melanospora***CBS 272.60**^T^**, DTO 103-I1, IFO 6441, IMI 078257, LCP 59.1590, MUCL 9040, NHL 6045***Oryza sativa*, milledUSAKX923876KX924311**LM652572**KX924036CBS 854.68, DTO 220-H7Compost soilGermanyKX923877KX924312KX924099*S. fusca*CBS 102829, DTO 342-E7Cheese warehouseNetherlandsKX923878KX924313KX924100CBS 116060, DTO 136-G8Antique tapestriesPoland**LN850775LN850872**KX924101DTO 043-A1UnknownUnknownKX923879KX924314KX924102DTO 043-A2UnknownUnknownKX923880KX924315KX924103DTO 049-E4ArchiveNetherlandsKX923881KX924316KX924104DTO 049-E5ArchiveNetherlandsKX923882KX924317KX924105DTO 049-F2OfficeNetherlandsKX923883KX924318KX924106DTO 053-H2Between concrete Floor and carpetNetherlandsKX923884KX924319KX924107DTO 067-G7BakeryNetherlandsKX923885KX924320KX924108DTO 138-B6IndoorGermanyKX923886KX924321KX924109DTO 220-H9IndoorSouth AfricaKX923887KX924322KX924110DTO 220-I1IndoorSouth AfricaKX923888KX924323KX924111DTO 220-I2IndoorSouth AfricaKX923889KX924324KX924112DTO 220-I3IndoorSouth AfricaKX923890KX924325KX924113DTO 223-A9IndoorGermanyKX923891KX924326KX924114DTO 240-A9ArchiveNetherlandsKX923892KX924327KX924115DTO 240-B1ArchiveNetherlandsKX923893KX924328KX924116DTO 240-B3ArchiveNetherlandsKX923894KX924329KX924117DTO 240-B4ArchiveNetherlandsKX923895KX924330KX924118DTO 252-D7IndoorGermanyKX923896KX924331KX924119DTO 255-A5AirsamplingGermanyKX923897KX924332KX924120DTO 255-A6AirsamplingGermanyKX923898KX924333KX924121DTO 255-A7AirsamplingGermanyKX923899KX924334KX924122DTO 255-B1PlasterGermanyKX923900KX924335KX924123DTO 255-B3PolystyreneGermanyKX923901KX924336KX924124DTO 255-B5Oriented strand boardGermanyKX923902KX924337KX924125DTO 255-B6WoodGermanyKX923903KX924338KX924126DTO 255-C3UnknownGermanyKX923904KX924339KX924127*M. micronesiensis* sp.  nov.**CBS 141523**^T^**, DTO 220-I9**IndoorMicronesiaKX923905KX924340KX924128KX924037DTO 223-A5IndoorMicronesiaKX923906KX924341KX924129*M. murinus*CBS 621.70, DTO 347-C8Composted municipal wasteGermanyKX923907**LN850868**KX924130**CBS 830.70**^T^**, DTO 104-B5, DTO 170-F5, IMI 161540**Composted municipal wasteGermanyKX923908KX924342KX924131[**HG380481**](ncbi-n:HG380481){#intref0160}CBS 864.71, DTO 347-C9Municipal wasteGermanyKX923909**LN850867**KX924132*M. onychoides***CBS 139629**^T^**, BMU 03911**Human, nailChina**LN850774LN850871LN850920LN850823***M. paisiiTorula paisii***CBS 213.27**^T^**, DTO 103-F9, IMI 036480, MUCL 7915, VKM F-424**HumanItaly**LM652434**KX924343KX924133**LM652518***S. sphaerospora*^T^CBS 402.34, DTO 103-G8, MUCL 9045UnknownAustria**LM652437**KX924344**LM652651***S. brumptii*^IT\ (?) ^CBS 333.35, DTO 220-H5Small-pox vaccineFranceKX923910KX924345KX924134CBS 345.58, DTO 220-H6Human, skin and hairGermany**LN850777LN850874LN850923**DTO 073-F1Moist wall of archiveNetherlandsKX923911KX924346KX924135DTO 109-G6IndoorDenmarkKX923912KX924347KX924136DTO 220-I6IndoorNew ZealandKX923913KX924348KX924137DTO 220-I7IndoorNew ZealandKX923914KX924349KX924138DTO 252-D9IndoorGermanyKX923915KX924350KX924139DTO 255-A8AirsamplingGermanyKX923916KX924351KX924140DTO 255-A9AirsamplingGermanyKX923917KX924352KX924141DTO 255-B2PlasterGermanyKX923918KX924353KX924142DTO 255-B4Oriented strand boardGermanyKX923919KX924354KX924143DTO 255-B7PlasterGermanyKX923920KX924355KX924144DTO 255-B8PolystyreneGermanyKX923921KX924356KX924145DTO 255-B9AirsamplingGermanyKX923922KX924357KX924146*M. pseudolongirostrisM. cirrosus***CBS 462.97**^T^**, DTO 351-D5**Human, nailNetherlands**LN850782LN850879**KX924147**LN850831***M. pseudopaisii* sp.  nov.**CBS 141581**^T^**, DTO 116-A3**Air, basementNetherlandsKX923923KX924358KX924148KX924038DTO 116-A4Air, basementNetherlandsKX923924KX924359KX924149*M. pyramidus***CBS 212.65**^T^**, DTO 104-A1, DTO 104-A2, ATCC 36763, IMI 109887**Desert soilUSAKX923925KX924360KX924150[**HG380435**](ncbi-n:HG380435){#intref0165}CBS 668.71, DTO 342-E4Pocket mouse, hairUSAKX923926**LN850876LN850925***M. trigonosporus*CBS 663.71, DTO 342-G1SoilUSAKX923927KX924361KX924151*M. restrictus***CBS 138277**^T^**, DTO 347-B4, UTHSC 09-2704, FMR 12227**Human, left halluxUSAKX923928KX924362KX924152[**HG380494**](ncbi-n:HG380494){#intref0170}*M. senegalensis***CBS 277.74**^T^**, DTO 351-E4**Mangrove soilSenegalKX923929KX924363KX924153**LM652523**CBS 760.84, DTO 347-D2*Helianthus annuus*, seedUSAKX923930KX924364KX924154CBS 761.84, DTO 342-G5*Helianthus annuus*, seedUSAKX923931KX924365KX924155CBS 775.84, DTO 342-C9*Helianthus annuus*, seedGermanyKX923932KX924366KX924156DTO 342-F1, RGR 84.0112UnknownUnknownKX923933KX924367KX924157DTO 342-G9, RGR 84.0113UnknownUnknownKX923934KX924368KX924158DTO 342-H1, RGR 84.0158UnknownUnknownKX923935KX924369KX924159DTO 342-H2, RGR 84.0159UnknownUnknownKX923936KX924370KX924160DTO 342-H4, RGR 85.0058unknownUnknownKX923937KX924371KX924161*M. terreus***CBS 601.67**^T^**, DTO 104-A8, ATCC 22360, NRRL A-18283, VKM F-1144**SoilUkraine**LN850783LN850880LN850928LN850832***M. trigonosporus*CBS 665.71, DTO 342-E3SoilUSAKX923938KX924372KX924162*M. trigonosporus*CBS 807.73, DTO 342-E5Saline desert soilKuwaitKX923939KX924373KX924163CBS 138275, UTHSC 07-1823, FMR 12342Human, sputumUSA**LM652384LM652600**[**HG380412**](ncbi-n:HG380412){#intref0175}*M. trigonosporus*DTO 342-D3, RGR 84.0004, ATCC 62716*Helianthus annuus*, confectionary seedUSAKX923940KX924374KX924164*M. trigonosporus*DTO 343-A1, RGR 84.0003*Helianthus annuus*, seedUSAKX923941KX924375KX924165*M. trautmannii* sp.  nov.**CBS 141583**^T^**, DTO 255-C1**oriented strand boardGermanyKX923942KX924376KX924166KX924039*M. trigonosporus***CBS 218.31**^T^**, DTO 103-G2, HACC 178, IMI 113702**UnknownPuerto RicoKX923943KX924377[**HG380359**](ncbi-n:HG380359){#intref0180}[**HG380436**](ncbi-n:HG380436){#intref0185}CBS 198.61, DTO 342-C5, IMI 086911, NRRL 1570UnknownUnknownKX923944KX924378KX924167CBS 199.61, DTO 342-C6, IFO 7027, IMI 086912, MUCL 9061, NHL 2265*Oryza sativa*, milledBurma**LM652444**KX924379[**HG380361**](ncbi-n:HG380361){#intref0190}CBS 366.65, DTO 342-F9, ATCC 16203, HACC 178UnknownIndiaKX923945KX924380KX924168CBS 158.92, DTO 342-E6, FRR 4046*Arachis hypogaea*, nutThailandKX923946KX924381KX924169DTO 220-I8IndoorMicronesiaKX923947KX924382KX924170DTO 223-A1IndoorMicronesiaKX923948KX924383KX924171DTO 223-A2IndoorMicronesiaKX923949KX924384KX924172*M. verrucosus***CBS 138278**^T^**, DTO 351-D9, UTHSC 10-2601, FMR 12219**Human, BAL fluidUSAKX923950**LM652658**[**HG380416**](ncbi-n:HG380416){#intref0195}[**HG380493**](ncbi-n:HG380493){#intref0200}*S. sphaerospora*CBS 210.61, DTO 347-B3, IMI 086939UnknownUnknownKX923951KX924385KX924173*Pithoascus stoveri***CBS 176.71**^T^**, DTO 104-B6, ATCC 11173***Beta vulgaris*, root seedlingUSAKX923952KX924386KX924174**LM652532***Pseudoscopulariopsis schumacheri***CBS 435.86**^NT^**, DTO 170-H5**SoilSpainKX923953KX924387KX924175**LM652534***Scopulariopsis africana* sp.  nov.*M. manginii***CBS 118736**^T^**, DTO 336-D1**Mud, salt panSouth AfricaKX923954KX924388KX924176KX924040*S. albida* sp.  nov.*S. flava***CBS 119.43**^T^**, DTO 334-G7**SoilNetherlands**LN850800LN850897LM652592LN850849***S. acremonium*CBS 415.51, DTO 334-H2UnknownGermanyKX923955KX924389KX924177*S. alboflavescensS. koningii*CBS 152.22, DTO 347-C5, IMI 086928, MUCL 9044UnknownFrance**LN850785LN850882**KX924178**CBS 399.34**^T^**, DTO 103-G6, UAMH 934**Human, skinAustriaKX923956[**JQ434537**](ncbi-n:JQ434537){#intref0205}KX924179**LM652539***S. koningii*CBS 208.61, DTO 170-C6, IMI 086926, FMR 3654ElephantUnknown**LN850786LN850883LN850931**DTO 104-C7UnknownUnknownKX923957KX924390KX924180*S. asperulaS. arnoldii*^T^CBS 204.27, DTO 334-F5, MUCL 9009, UAMH 923UnknownFranceKX923958KX924391KX924181*S. fusca*^T^CBS 401.34, DTO 103-G7, IFO 8181, IMI 086934, MUCL 9032, UAMH 930Rabbit carcassAustria**LM652463**KX924392KX924182CBS 105.35, DTO 334-F8, IMI 086925UnknownUnknownKX923959KX924393KX924183*Torula bestae*^T^CBS 289.38, DTO 103-H1, IMI 086927, MUCL 9012, UAMH 924HumanItalyKX923960KX924394KX924184**LM652538***S. fusca*CBS 351.49, DTO 342-C4, MUCL 9033*Sciurus vulgaris*, deadUnknownKX923961KX924395KX924185CBS 390.52, DTO 334-H4, IMI 086924, LCP 224, MUCL 9043*Calliphora vomitoria*, proboscisFranceKX923962KX924396KX924186*S. fusca*CBS 334.53, DTO 334-H5Human, nailNetherlands**LN850788LN850885**KX924187CBS 298.67, DTO 334-I4*Triticum aestivum*Turkey**LN850789LN850886LN850934**CBS 853.68, DTO 334-I5Compost soilGermanyKX923963[**JQ434558**](ncbi-n:JQ434558){#intref0210}KX924188CBS 872.68, DTO 334-I6, ATCC 16281Wheat field soilGermanyKX923964KX924397KX924189CBS 668.74, DTO 335-A5SoilEgyptKX923965KX924398KX924190CBS 373.76, DTO 170-G6UnknownNetherlandsKX923966KX924399KX924191CBS 114063, DTO 038-B3, DTO 335-B3Wood sampleGermanyKX923967KX924400KX924192CBS 117767, DTO 038-B6Wood sampleGermanyKX923968**LN850884**KX924193*S. fusca*CBS 138116, DTO 335-D1Alkaline soilRussiaKX923969KX924401KX924194*S. brevicaulis*CBS 120.20, DTO 334-F4, MUCL 9021UnknownUnknownKX923970KX924402KX924195CBS 273.30, DTO 334-F6, VKM F-175UnknownUnknownKX923971KX924403KX924196*S. flava*CBS 334.35, DTO 334-G1*Arge berberidis*, pupaCzech Republic**LN850790LN850887LN850935***S. flava*CBS 335.35, DTO 334-G2, IMI 086922, MUCL 9035*Pteronus pini*, pupaNetherlands**LM652477**KX924404KX924197CBS 340.39, DTO 336-C4BoneSouth AfricaKX923972KX924405KX924198CBS 341.39, DTO 334-G4UnknownUnknownKX923973KX924406KX924199CBS 147.41, DTO 334-G5Human, nailNetherlandsKX923974KX924407KX924200CBS 467.48, DTO 103-H4, ATCC 7903, IMI 040026, IMI 061534, NRRL 1096UnknownUnknownKX923975KX924408KX924201CBS 398.54, DTO 170-C3, IMI 086919Human, toe nailUKKX923976KX924409KX924202CBS 112377, DTO 011-H5, DTO 038-A8IndoorGermanyKX923977KX924410KX924203CBS 115540, DTO 335-B4Air biofilterMexicoKX923978KX924411KX924204CBS 116112, DTO 335-B5Tattoo-paintCzech RepublicKX923979KX924412KX924205CBS 117277, DTO 001-F7, DTO 012-E7Hat-rack in museumNetherlandsKX923980KX924413KX924206CBS 118469, DTO 338-H1Tattoo-paintUKKX923981KX924414KX924207CBS 118470, DTO 336-C8Tattoo-paintUKKX923982KX924415KX924208CBS 118471, DTO 335-B6Tattoo-paintUKKX923983KX924416KX924209CBS 118472, DTO 335-B7Tattoo-paintUKKX923984KX924417KX924210CBS 118473, DTO 338-H2Tattoo-paintUKKX923985KX924418KX924211CBS 118474, DTO 336-C9Tattoo-paintUKKX923986KX924419KX924212CBS 118993, DTO 335-B8UnknownFranceKX923987KX924420KX924213CBS 119549, DTO 335-B9Human, skin biopsyUSAKX923988KX924421KX924214CBS 119550, DTO 335-C1Human, blood cultureUSAKX923989KX924422KX924215*M. brevicaulis*^T^CBS 127812, DTO 138-E6, DTO 138-E7, UAMH 7770, MUCL 40726Indoor airCanada**LM652465**KX924423[**HG380363**](ncbi-n:HG380363){#intref0215}[**HG380440**](ncbi-n:HG380440){#intref0220}CBS 127825, DTO 138-E8, DTO 138-E9, UAMH 7880Indoor airCanadaKX923990KX924424KX924216CBS 137631, DTO 335-C8Alkaline soilRussiaKX923991KX924425KX924217CBS 137632, DTO 335-C9Alkaline soilRussiaKX923992KX924426KX924218DTO 012-C7Indoor airGermanyKX923993KX924427KX924219DTO 012-D9Wall paperUnknownKX923994KX924428KX924220DTO 012-F6PlasterGermanyKX923995KX924429KX924221DTO 106-B6Indoor, giraffes stayNetherlandsKX923996KX924430KX924222DTO 109-H4IndoorDenmarkKX923997KX924431KX924223DTO 145-C7IndoorGermanyKX923998KX924432KX924224DTO 168-A4Indoor air, poultry housePolandKX923999KX924433KX924225DTO 168-A5Indoor air, poultry housePolandKX924000KX924434KX924226DTO 168-A6Indoor air, poultry housePolandKX924001KX924435KX924227DTO 195-A1Indoor, swab sample bakeryNetherlandsKX924002KX924436KX924228DTO 197-F3Indoor air sampleNetherlandsKX924003KX924437KX924229DTO 240-A8ArchiveNetherlandsKX924004KX924438KX924230DTO 305-A2IndoorSouth AfricaKX924005KX924439KX924231DTO 305-A3IndoorUSAKX924006KX924440KX924232DTO 305-A4IndoorUSAKX924007KX924441KX924233DTO 305-A5IndoorUSAKX924008KX924442KX924234DTO 305-A8IndoorUSAKX924009KX924443KX924235*S. candidaNephrospora manginii*^T^CBS 170.27, DTO 347-C6, IMI 086931, UAMH 9135UnknownFrance**LM652488LM652694LM652594***M. manginii*CBS 205.27, DTO 347-C7, MUCL 9026UnknownFrance**LM652483**KX924444KX924236*S. acremonium*CBS 305.31, DTO 336-C3UnknownUSAKX924010KX924445KX924237*M. manginii*CBS 353.36, DTO 334-G3Indoor air, hospitalunknownKX924011KX924446KX924238*S. acremonium*CBS 389.52, DTO 334-H3UnknownItalyKX924012KX924447KX924239*M. manginii*CBS 254.69, DTO 335-A1Greenhouse soilNetherlandsKX924013KX924448KX924240*M. manginii*CBS 132.78, DTO 342-G3Human, dentineFranceKX924014**LN850898**KX924241DTO 032-A6Indoor air, houseNetherlandsKX924015KX924449KX924242DTO 138-B7Indoor airGermanyKX924016KX924450KX924243DTO 139-D8IndoorGermanyKX924017KX924451KX924244DTO 139-E7IndoorGermanyKX924018KX924452KX924245DTO 139-E8IndoorGermanyKX924019KX924453KX924246**UAMH 9004**^ET^**, MUCL 40743**Indoor airCanada**LM652484LM652690**[**HG380381**](ncbi-n:HG380381){#intref0225}[**HG380458**](ncbi-n:HG380458){#intref0230}*S. caseicola* sp.  nov.*S. acremonium***CBS 480.62**^T^**, DTO 342-F8**Cheese-coatingNetherlandsKX924020KX924454KX924247KX924041*S. cordiaeM. manginii*CBS 816.73, DTO 335-A4SoilAustraliaKX924021KX924455KX924248**CBS 138129**^T^**, DTO 335-D2, UTHSC 09-866, FMR 12338**Human, fingerUSAKX924022KX924456KX924249[**HG380499**](ncbi-n:HG380499){#intref0235}*S. flava***CBS 207.61**^NT^**, DTO 170-C5, IMI 086921, MUCL 9031**CheeseUKKX924023KX924457[**HG380387**](ncbi-n:HG380387){#intref0240}[**HG380464**](ncbi-n:HG380464){#intref0245}*S. brevicaulis*CBS 108960, DTO 335-B2CheeseDenmark**LN850804LN850901LN850949***S. macuraeM. manginii***CBS 506.66**^T^**, DTO 334-I3, ATCC 16685**Chicken litterCanada**LN850805LN850902**KX924250**LN850854***S. sexualis* sp.  nov.*M. manginii***CBS 250.64**^T^**, DTO 338-G3, IFO 7555, NHL 2278, UAMH 1923***Oryza sativa*, milledBurmaKX924024KX924458KX924251KX924042*M. manginii*CBS 667.71, DTO 335-A3, NRRL A-8022Bat dungUSAKX924025KX924459KX924252*M. manginii*CBS 332.78, DTO 342-G4*Brassica oleracea*, seedIndiaKX924026KX924460KX924253*S. soppii***UAMH 9169**^T^*Populus tremuloides*, woodCanada**LM652495LM652698LM652595LM652552***Yunnania carbonariaS. carbonaria***CBS 205.61**^T^**, DTO 103-I3, IFO 8116, IMI 086941, MUCL 9027, NRRL 1860**SoilPanamaKX923820KX924254KX924044[**HG380462**](ncbi-n:HG380462){#intref0250}*S. brumptii*CBS 121662, DTO 220-H8Dead hardwood branchUSAKX923821KX924255KX924045*Y. penicillata***CBS 130296**^T^**, DTO 139-F4**Molded pork sampleChina[**JN831361**](ncbi-n:JN831361){#intref0255}KY659807KY659808KY659809*Y. smithii* sp.  nov.*S. carbonaria***CBS 855.68**^T^**, DTO 354-C2**Garden soilGermanyKX923822KX924256KX924046KX924028[^1][^2]

DNA isolation, PCR and sequencing {#sec2.2}
---------------------------------

DNA extraction was performed using the Ultraclean^®^ Microbial DNA Isolation Kit (MoBio laboratories, Carlsbad, CA, USA), according to the manufacturer\'s instructions.  The LSU, ITS, *tub2* and *tef1* gene regions were amplified and sequenced with respectively the primers LR0R ([@bib24])/LR5 ([@bib34]), V9G ([@bib6])/LS266 ([@bib17]), Bt2a/Bt2b ([@bib8]) and EF1-983F/EF1-2218R ([@bib23]).  The PCRs were performed in an Applied Biosystems^®^ 2720 Thermal Cycler (Thermo Fisher Scientific, Bleiswijk, the Netherlands) in a total volume of 12.5 μl.  The PCR mixture consisted of 1 μl genomic DNA, 1 × NH~4~ reaction buffer (Bioline, Luckenwalde, Germany), 0.2 μM of each primer, 5 % dimethyl sulfoxide (DMSO), 20 μM (*tub2*) or 40 μM (LSU/ITS/*tef1*) of each dNTP, 1 mM (ITS) or 1.6 mM (*tef1*) or 2 mM (LSU/*tub2*) MgCl~2~, and 0.25 U *Taq* DNA polymerase (Bioline).  The PCR conditions for LSU, ITS and *tub2* consisted of an initial denaturation step of 5 min at 94 °C followed by 35 cycles of 30 s at 94 °C, 30 s at 47 °C (LSU) or 55 °C (ITS) or 59 °C (*tub2*) and 1 min at 72 °C, and a final elongation step of 7 min at 72 °C.  For *tef1* a touchdown PCR protocol of 9 cycles of 30 s at 94 °C, 30 s at 66 °C (−1 °C every cycle) and 90 s at 72 °C, followed by 30 cycles of 30 s at 94 °C, 30 s at 56 °C and 90 s at 72 °C and a final elongation step of 7 min at 72 °C was used.  The PCR products were sequenced in both directions using the BigDye Terminator v. 3.1 Cycle Sequencing Kit (Thermo Fisher Scientific) and analysed with an ABI Prism 3730xl DNA Analyser (Thermo Fisher Scientific) according to the manufacturer\'s instructions.  Consensus sequences were computed from forward and reverse sequences using the Bionumerics v. 4.61 software package (Applied Maths, St-Marthens-Latem, Belgium).  Several sequences obtained in this study had one or multiple nucleotide differences and length differences with already published sequenced of the same isolates.  All new sequences and sequences which were longer in length or had nucleotide differences with already published sequenced were submitted to GenBank ([Table 1](#tbl1){ref-type="table"}). 

Phylogenetic analyses {#sec2.3}
---------------------

Multiple sequence alignments of the separate LSU, ITS, *tub2* and *tef1* sequences were generated with MAFFT v. 7.271 (<http://mafft.cbrc.jp/alignment/server/index.html>) using the L-INS-i method.  With Findmodel (<http://www.hiv.lanl.gov/content/sequence/findmodel/findmodel.html>) the best nucleotide substitution models were determined.  On both the single gene-sequence alignments and the combined gene-sequence alignment Bayesian analyses were performed with MrBayes v. 3.1.2 ([@bib11], [@bib25]).  The Markov Chain Monte Carlo (MCMC) analysis used four chains and started from a random tree topology.  The sample frequency was set at 1 000 and the temperature value of the heated chain was set at 0.1.  The run stopped when the average standard deviation of split frequencies reached below 0.01.  Burn-in was set to 25 % after which the likelihood values were stationary.  Tracer v. 1.5.0 ([@bib21]) was used to confirm the convergence of chains.  Maximum-likelihood analyses including 500 bootstrap replicates using RAxML v. 7.2.6 ([@bib32]) were additionally run on both the single gene-sequence alignments and the combined gene-sequence alignment.  The resulting trees were printed with TreeView v. 1.6.6 ([@bib20]) and, together with the alignments, deposited into TreeBASE (<http://www.treebase.org>). 

In order to get optimal sequence alignments, the dataset was divided in three different phylogenies.  Based on the ITS, *tub2* and *tef1* sequences a separate species phylogeny for *Microascus* (157 isolates) and for *Scopulariopsis* (89 isolates) was constructed.  Following these species phylogenies a genus phylogeny including one isolate per recognised species (when present the ex-type isolate) was constructed based on the LSU, ITS and *tef1* sequences (52 isolates).  The *tub2* sequences were omitted in the genus phylogeny because of alignment difficulties.  Based on former phylogenetic studies ([@bib14], [@bib29]) *Pithoascus stoveri* (CBS 176.71) was used as out-group in the *Microascus* phylogeny, *Pseudoscopulariopsis schumacheri* (CBS 435.86) in the *Scopulariopsis* phylogeny, and *Cephalotrichum stemonitis* (CBS 103.19) in the genus phylogeny. 

Habitat study {#sec2.4}
-------------

All studied *Microascus*, *Scopulariopsis* and *Yunnania* isolates are assigned to one of nine different habitats (animal, dung, food, human, indoor, insect, plant, soil, others) based on their origin of isolation.  Subsequently, these habitats are plotted behind the species name, to which the isolate belongs, in the genus phylogeny tree.  The habitats from isolates studied by others ([@bib26], [@bib14], [@bib29]) are also included.  Finally, a table showing which *Microascus*, *Scopulariopsis* and *Yunnania* species are present in which habitat is constructed. 

Morphology {#sec2.5}
----------

Cultures were incubated on oatmeal agar (OA), malt extract agar (MEA) and dichloran 18 % glycerol agar (DG18) plates (recipes [@bib28]) at 25 °C in the dark.  After 14 d the colony diameters were measured and the colony characters noted.  Colony colours were rated according to [@bib22].  Measurements and descriptions of microscopic structures were made from cultures grown on synthetic nutrient agar (SNA, [@bib28]) at 25 °C in the dark for 14 d or longer to ensure ascomata development.  Slide preparations of the asexual morph structures were made with the sellotape technique ([@bib31]) or mounted in 85 % lactic acid, like the sexual morph structures.  Photographs of characteristic structures were made with a Zeiss Axio Imager A2 microscope equipped with a Nikon DS-Ri2 high-definition colour camera head using differential interference contrast (DIC) optics and the Nikon software NIS-elements D v. 4.50.  Furthermore, growth at 36 °C and 40 °C in the dark on OA was tested. 

Results {#sec3}
=======

*Microascus* phylogeny {#sec3.1}
----------------------

For the phylogeny 157 isolates were selected to represent the genus *Microascus* ([Table 1](#tbl1){ref-type="table"}) including the outgroup-isolate *Pithoascus stoveri* (CBS 176.71).  The aligned sequences of the ITS (474 characters), *tub2* (529 characters), and *tef1* (898 characters) gene regions had a total length of 1 901 characters, with respectively 147, 253, and 221 unique site patterns.  The GTR model with a gamma-distributed rate variation was suggested as model for the ITS and *tef1* alignments and the HKY model with a gamma-distributed rate variation for the *tub2* alignment.  After discarding the burn-in phase trees, the multi-gene Bayesian analysis resulted in 4 172 trees from two runs from which the majority rule consensus tree and posterior probabilities were calculated.  The multi-gene phylogeny divided the isolates in 33 *Microascus* species (clades, [Fig.  1](#fig1){ref-type="fig"}), and three *Yunnania* species (clades, [Fig.  1](#fig1){ref-type="fig"}).  As a result of this study, seven new *Microascus* species (*M. appendiculatus*, *M. cleistocarpus*, *M. fusisporus*, *M. hollandicus*, *M. micronesiensis*, *M. pseudopaisii*, and *M. trautmannii*), one new *Yunnania* species (*Y. smithii*), one new name (*M. atrogriseus*) and two new name combinations (*M. melanosporus*, and *Y. carbonaria*) are proposed.  The recently established genus *Fuscoannellis* is synonymised under *Yunnania*.  All descriptions are provided below in the taxonomy section.  For the genus *Microascus*, only the *tef1* phylogeny can distinguish all identified species.  This is in congruence with [@bib26] and [@bib14].  With *tub2*, the species *M. restrictus* and *M. verrucosus* cannot be separated, they can molecularly only clearly be distinguished based on their *tef1* sequence (ITS 2 nt difference, *tub2* 1 nt difference, *tef1* 20 nt difference).  The ITS single gene phylogeny is least distinctive.  Besides *M. restrictus* and *M. verrucosus*, *M. alveolaris* and *M. terreus*, *M. intricatus* and *M. onychoides*, *M. paisii* and *M. melanosporus* and the three *Yunnania* species cannot be separated based on their ITS sequences.  Furthermore, the four isolates of *M. cinereus* are split into two clades based on their ITS sequence alone (data not shown, all single gene phylogenies are submitted to TreeBase).Fig.  1Maximum likelihood tree based on the ITS, *tub2* and *tef1* sequences of 157 isolates representing the genera *Microascus* and *Yunnania*.  The RAxML bootstrap support values ≥75 % (BS) and Bayesian posterior probabilities ≥0.95 (PP) are given at the nodes.  Thickened lines indicate a BS of 100 % and a PP of 1.0.  Ex-type strain numbers are in **bold** face and indicated with ^T^ (or ^NT^ when ex-neotype).  Species names between parentheses represent synonymised species names.  The tree was rooted to *Pithoascus stoveri* (CBS 176.71).Fig.  1

*Scopulariopsis* phylogeny {#sec3.2}
--------------------------

For the phylogeny 89 isolates were selected to represent the genus *Scopulariopsis* ([Table 1](#tbl1){ref-type="table"}) including the outgroup-isolate *Pseudoscopulariopsis schumacheri* (CBS 435.86).  The aligned sequences of the ITS (441 characters), *tub2* (502 characters), and *tef1* (887 characters) gene regions had a total length of 1 830 characters, with respectively 58, 143, and 109 unique site patterns.  The TrN model with a gamma-distributed rate variation was suggested as model for the ITS alignment, the GTR model with a gamma-distributed rate variation as model for the *tef1* alignment and the HKY model with a gamma-distributed rate variation for the *tub2* alignment.  After discarding the burn-in phase trees, the multi-gene Bayesian analysis resulted in 2 214 trees from both runs from which the majority rule consensus tree and posterior probabilities were calculated.  The multi-gene phylogeny divided the isolates in 12 species (clades, [Fig.  2](#fig2){ref-type="fig"}) of which four are proposed as new; *S. africana*, *S. albida*, *S. caseicola* and *S. sexualis*.  Their descriptions are provided below in the taxonomy section.  Both with only the *tub2* or *tef1* sequence all 12 species can be identified, although the *S. candida* isolates do not form a monophyletic clade.  Based on ITS alone, only five species can be identified, *S. alboflavescens*, *S. brevicaulis*, *S. flava*, *S. macurae* and *S. soppii* (data not shown, all single gene phylogenies are submitted to TreeBase).Fig.  2Maximum likelihood tree based on the ITS, *tub2* and *tef1* sequences of 89 isolates representing the genus *Scopulariopsis*.  The RAxML bootstrap support values ≥75 % (BS) and Bayesian posterior probabilities ≥0.95 (PP) are given at the nodes.  Thickened lines indicate a BS of 100 % and a PP of 1.0.  Ex-type strain numbers are in **bold** face and indicated with ^T^ (or ^NT^ or ^ET^ when ex-neotype or ex-epitype respectively).  Species names between parentheses represent synonymised species names.  The tree was rooted to *Pseudoscopulariopsis schumacheri* (CBS 435.86).Fig.  2

Genus phylogeny with habitat study {#sec3.3}
----------------------------------

For the genus phylogeny ([Fig.  3](#fig3){ref-type="fig"}) 52 isolates were selected to represent all above recognised species in the genera *Microascus*, *Scopulariopsis* and *Yunnania* together with *Pithoascus stoveri* (CBS 176.71), *Pseudoscopulariopsis schumacheri* (CBS 435.86), *Cephalotrichum asperulum* (CBS 582.71) and the out-group isolate *Cephalotrichum stemonitis* (CBS 103.19).  The phylogeny was constructed based on the LSU, ITS and *tef1* sequences.  The aligned sequences of the LSU (533 characters), ITS (474 characters), and *tef1* (814 characters) gene regions had a total length of 1 821 characters, with respectively 93, 175, and 209 unique site patterns.  The GTR model with a gamma-distributed rate variation was suggested as model for the ITS and *tef1* alignments and the TrN model with a gamma-distributed rate variation for the LSU alignment.  After discarding the burn-in phase trees, the multi-gene Bayesian analysis resulted in 1 180 trees from two runs from which the majority rule consensus tree and posterior probabilities were calculated.  The habitats are plotted behind the species names in the genus tree ([Fig.  3](#fig3){ref-type="fig"}) and placed in an overview table depicting the species per habitat ([Table 2](#tbl2){ref-type="table"}).  No specific clustering of habitat preference related to phylogenetic relationships can be found, the different habitats are scattered over the phylogenetic tree ([Fig.  3](#fig3){ref-type="fig"}).Fig.  3Maximum likelihood tree based on the LSU, ITS and *tef1* sequences of 52 isolates.  The RAxML bootstrap support values ≥75 % (BS) and Bayesian posterior probabilities ≥0.95 (PP) are given at the nodes.  Thickened lines indicate a BS of 100 % and a PP of 1.0.  When no collection number is mentioned behind the species name, the type-isolate is used.  The tree was rooted to *Cephalotrichum stemonitis* (CBS 103.19).  The habitats where the species are found are plotted behind the species names: ![]( fx1.gif) = indoor; ![]( fx2.gif) = animal; ![]( fx3.gif) = dung; ![]( fx4.gif) = food; ![]( fx5.gif) = human; ![]( fx6.gif) = insect; ![]( fx7.gif) = plant; ![]( fx8.gif) = soil; ![]( fx9.gif) = others. Fig.  3Table 2Current species within *Microascus*, *Scopulariopsis* and *Yunnania* per habitat. Table 2HabitatSpeciesIndoor*M. alveolaris*, *M. atrogriseus*, *M. chartarus*, *M. cirrosus*, *M. croci*, *M. gracilis*, *M. hollandicus*, *M. intricatus*, *M. melanosporus*, *M. micronesiensis*, *M. paisii*, *M. pseudopaisii*, *M. trautmannii*, *M. trigonosporus*, *S. asperula*, *S. brevicaulis*, *S. candida*Animal*M. cirrosus*, *M. pyramidus*, *S. alboflavescens*, *S. asperula*Dung*M. cirrosus*, *M. hyalinus*, *S. asperula*, *S. sexualis*, *S. macurae*Food*M. gracilis*, *M. longicollis*, *M. trigonosporus*, *S. asperula*, *S. brevicaulis*, *S. candida*, *S*. * caseicola*, *S. flava*, *Y. penicillata*Human*M. alveolaris*, *M. appendiculatus*, *M. brunneosporus*, *M. chinensis*, *M. cinereus*, *M. cirrosus*, *M. croci*, *M. expansus*, *M. gracilis*, *M. intricatus*, *M. longicollis*, *M. melanosporus*, *M. onychoides*, *M. paisii*, *M. restrictus*, *M. terreus*, *M. verrucosus*, *M. pseudolongirostris*, *S. alboflavescens*, *S. asperula*, *S. brevicaulis*, *S. candida*, *S. cordiae*Insect*M. longirostris*, *S. asperula*, *S. brevicaulis*Plant*M. alveolaris*, *M. cirrosus*, *M. croci*, *M. gracilis*, *M. melanosporus*, *M. paisii*, *M. senegalensis*, *M. terreus*, *M. trigonosporus*, *S. asperula*, *S. sexualis*, *S. soppii*, *Y. carbonaria*Soil*M. alveolaris*, *M. atrogriseus*, *M. cinereus*, *M. cirrosus*, *M. croci*, *M. fusisporus*, *M. gracilis*, *M. intricatus*, *M. longirostris*, *M. macrosporus*, *M. melanosporus*, *M. murinus*, *M. pyramidus*, *M. senegalensis*, *M. terreus*, *S. africana*, *S. albida*, *S. asperula*, *S. brevicaulis*, *S. candida*, *S. cordiae*, *Y. carbonaria*, *Y. smithii*Others*M. paisii*, *M. cleistocarpus*, *S. brevicaulis*

Seventeen species are found in the indoor environment, 14 *Microascus* species and three *Scopulariopsis* species ([Table 2](#tbl2){ref-type="table"}).  Most of them are only occasionally found in the indoor environment. * Scopulariopsis brevicaulis* (22 indoor isolates) and *M. melanosporus* (19 indoor isolates) are the most common indoor species, in number of isolates, followed by *M. paisii* (8 indoor isolates), *S. candida* (7 indoor isolates), and *M. croci* (5 indoor isolates).  Ten species which are found indoor are also found in relation with humans ([Fig.  3](#fig3){ref-type="fig"}), but mostly only from skin or nail infections, and more rarely in other tissues like pulmonary tissue (e.g. *M. cirrosus*) or blood culture (e.g. *S. brevicaulis*).  This needs to be taken into account when trying to indicate the risk for human health. 

*Scopulariopsis asperula* can be found in all included habitats, followed by *S. brevicaulis* and *M. cirrosus* which both are found in six different habitats and *M. gracilis* found in five different habitats ([Fig.  3](#fig3){ref-type="fig"}).  These species are all also found indoor, which is not surprisingly considering their non-selective habitats.  Five species, *M. chartarus*, *M. hollandicus*, *M. micronesiensis*, *M. pseudopaisii* and *M. trautmannii*, are only found in the indoor environment.  Of these five species, three are single isolate species and the other two only include two isolates (*M. micronesiensis* and *M. pseudopaisii*).  The two isolates of *M. pseudopaisii* are isolated from the same place, and could be seen as duplicates. * Microascus micronesiensis* has been found in two different houses in Micronesia in different cities on separate occasions, and has therefore the most potential in being a true indoor species. 

Taxonomy {#sec4}
========

Based on the multi-gene species phylogenies ([Fig.  1](#fig1){ref-type="fig"}, [Fig.  2](#fig2){ref-type="fig"}) 33 *Microascus* species, 12 *Scopulariopsis* species, and three *Yunnania* species are recognised.  In total 12 new species (four *Scopulariopsis*, seven *Microascus* and one *Yunnania* species), and a new name and two new name combination are proposed, which descriptions are provided below.  Additionally, all recent name changes are summarised in an overview table ([Table 3](#tbl3){ref-type="table"}).Table 3Overview of recent name changes in *Scopulariopsis* and scopulariopsis-like species. Table 3Old nameCurrent nameOld nameCurrent name*Kernia hyalinusMicroascus hyalinusScopulariopsis brevicaulis* var. * glabraScopulariopsis candidaMasonia chartarumMicroascus chartarusS. brumptiiMicroascus paisiiM. griseaMicroascus atrogriseusS. carbonariaYunnania carbonariaMasoniella chartarumMicroascus chartarusS. caseiScopulariopsis flavaM. crociMicroascus crociS. chartarumMicroascus chartarusM. griseaMicroascus atrogriseusS. crociMicroascus crociM. tertiaMicroascus crociS. fuscaScopulariopsis asperulaMicroascus brevicaulisScopulariopsis brevicaulisS. gracilisMicroascus gracilisM. exsertusPithoascus exsertusS. grylliScopulariopsis flavaM. griseusMicroascus cinereusS. hibernicaPseudoscopulariopsis hibernicaM. intermediusPithoascus intermediusS. hominisScopulariopsis brevicaulisM. manginiiScopulariopsis candidaS. insectivora*Scopulariopsis *brevicaulisM. nidicolaPithoascus nidicolaS. ivorensisScopulariopsis asperulaM. nigerScopulariopsis asperulaS. koningiiScopulariopsis brevicaulisM. schumacheriPseudoscopulariopsis schumacheriS. murinaMicroascus murinusM. soppiiScopulariopsis soppiiS. paisiiMicroascus paisiiM. stoveriPithoascus stoveriS. penicilloidesScopulariopsis brevicaulisM. stysanophorusPseudoscopulariopsis schumacheriS. roseolaScopulariopsis asperulaM. trigonosporus* var. * terreusMicroascus terreusS. rufulusScopulariopsis brevicaulisNephrospora manginiiScopulariopsis candidaS. sphaerosporaMicroascus paisiiPithoascus schumacheriPseudoscopulariopsis schumacheriS. stercorariaScopulariopsis brevicaulisP. stysanophorusPesudoscopulariopsis schumacheriS. trigonosporaMicroascus trigonosporusScopulariopsis arnoldiiScopulariopsis asperulaS. versicolorMicroascus paisiiS. atraPithoascus aterTorula asperulaScopulariopsis asperulaS. aureaScopulariopsis flavaT. bestaeScopulariopsis asperulaS. bestaeScopulariopsis asperulaT. paisiiMicroascus paisiiS. brevicaulis* var. * albaScopulariopsis flava*

***Microascus appendiculatus*** Woudenb. &  Samson **sp.  nov.**  MycoBank [MB818278](mycobank:818278){#intref0025}. [ Fig.  4](#fig4){ref-type="fig"}.Fig.  4*Microascus appendiculatus* sp.  nov.  CBS 594.78. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D.** Ascoma. ** E.** Ascomatal wall. ** F.** Asci and ascospores. ** G.** Ascospores. ** H, J--K.** Conidiophores, annellides and conidia. ** I.** Conidia.  Scale bars = 10 μm. Fig.  4

*Etymology*: name refers to its conidia with a basal appendage. 

*Ascomata* abundant, immersed, ostiolate, globose to subglobose with a short (up to 45 μm long) cylindrical ostiolar neck, (134--)158--208(--218) μm diam.,  black, glabrous; peridium with a *textura angularis*. * Asci* irregularly ellipsoidal, (19.5--)21--24.5(--25) × (10--)12.5--17.5(--20) μm. * Ascospores* fusiform, (5.5--)6.5--7.5(--8) × (3.5--)4--4.5(--5) μm, honey, pale luteous in mass, smooth, with a single inconspicuous germ pore. * Conidiophores* arising from substrate mycelium, indistinctive or simple, rarely branched, bearing terminally a single annellide. * Annellides* lageniform to ampulliform, (6--)7.5--11(--13.5) μm long, (2--)2.5--3(--3.5) μm broad at the widest part, tapering abruptly to a cylindrical annellate zone 0.5--1(--1.5) μm wide, hyaline to subhyaline, smooth-walled. * Conidia* subglobose with small basal appendage, (5--)5.5--7 × (3.5--)4--5(--5.5) μm, subhyaline, older conidia covered with hazel mucilaginous coating, smooth, thick-walled, arranged in short chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 27--28 mm after 14 d at 25 °C, flat, white to cream-coloured with smoke grey zones and olivaceous grey ascomata, margin undulated.  On MEA attaining a diameter of 17--19 mm, convex, white to cream-coloured, radially striated with dentate margin.  On DG18 attaining a diameter of 19 mm, low convex, white to cream-coloured with partly a grey olivaceous ring close to the edge, margin undulate.  On OA able to grow at 36 and 40 °C. 

*Specimen examined*: **Algeria**, from human skin, collection date and collector unknown, (**holotype** CBS H-22744, culture **ex-type** CBS 594.78). 

*Notes*: The ex-type strain of *M. appendiculatus* (CBS 594.78) was recently published as *M. senegalensis* ([@bib14]).  However, the sequences of their two included *M. senegalensis* isolates deposited on GenBank (which are confirmed by resequencing the isolate) only have 94 % identity based on ITS, 99 % on LSU, 90 % on *tub2* and 96 % on *tef1*.  Although they seem to cluster together in their phylogenetic tree, our phylogenies (all single gene phylogenies, and the combined phylogeny [Fig.  2](#fig2){ref-type="fig"}) places CBS 594.78 as a separate species, which is described here as *M. appendiculatus*.  Also morphologically it is distinct from *M. senegalensis* with the subglobose conidia with small basal appendage and the hazel mucilaginous coating around the older conidia. 

***Microascus atrogriseus*** Woudenb. &  Samson **nom.  nov.**  MycoBank [MB818284](mycobank:818284){#intref0030}. [ Fig.  5](#fig5){ref-type="fig"}.Fig.  5*Microascus atrogriseus* nom.  nov. ** A--C.** Fourteen day old colonies of DTO 139-D7 on OA (A), MEA (B) and DG18 (C). ** D--F.** Conidiophores, annellides and conidia CBS 295.52. ** G.** Conidiophores, annellides and conidia DTO 139-D7. ** H.** Conidia DTO 191-C2.  Scale bars = 10 μm. Fig.  5

*Basionym*: *Masonia grisea* G. Sm.,  Trans.  Brit.  Mycol.  Soc.  35: 149.  1952, non *Microascus griseus* P.N. Mathur & Thirum,.  1963. 

≡ *Masoniella grisea* (G. Sm.)  G. Sm.,  Trans.  Brit.  Mycol.  Soc.  35: 237.  1952. 

*Etymology*: name refers to the original description of the basionym where "atro-griseis coloniis" are described. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, simple or indistinctive, bearing one or multiple annellides. * Annellides* ampulliform, (4.5--)5.5--8.5(--10) μm long, 2--3 μm broad at the widest part, tapering abruptly to a cylindrical annellate zone 1--1.5(--2) μm wide, hyaline, smooth-walled. * Conidia* broadly ellipsoidal to short clavate with truncate base, (3--)3.5--4(--4.5) × (2.5--)3(--3.5) μm, hyaline when young turning hazel when ageing, smooth, thick-walled, arranged in chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 17--20 mm after 14 d at 25 °C, flat, white to cream-coloured with (pale) olivaceous grey to iron grey centre, margin crenated.  On MEA attaining a diameter of 13--14 mm, convex, white to very pale olivaceous grey, radially striated edge with crenated margin.  On DG18 attaining a diameter of 10--20 mm, low convex, white to pale olivaceous grey, margin entire.  On OA no growth at 36 and 40 °C. 

*Specimens examined*: **England**, London, isolated as culture contaminant, 1946, G. smith, (culture **ex-type** CBS 295.52). ** Germany**, indoor environment, before Aug. 2010, collector unknown, DTO 139-D7. ** Netherlands**, from a swab sample of an indoor horse arena, Mar. 2012, Houba, DTO 191-C2. 

*Notes*: *Microascus atrogriseus* is morphologically indistinguishable from *M. paisii*.  Sequence data is necessary to distinguish it from *M. paisii*.  All three genes used in this manuscript can separate the two species (ITS 5 nt difference, *tub2* 20 nt difference, and *tef1* 9 nt difference between the type isolates of both species). 

***Microascus cleistocarpus*** Woudenb.,  X. Wei Wang & Samson **sp.  nov.**  MycoBank [MB803264](mycobank:803264){#intref0035}. [ Fig.  6](#fig6){ref-type="fig"}.Fig.  6*Microascus cleistocarpus* sp.  nov.  CBS 134638. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D, G.** Ascomata. ** E.** Ascomatal wall. ** F.** Asci and ascospores. ** H--J.** Conidiophores, annellides and conidia. ** K.** Conidia.  Scale bars = 10 μm. Fig.  6

*Etymology*: named after the non-ostiolate ascomata. 

*Ascomata* abundant, immersed, non-ostiolate, globose to subglobose, (50--)52--71(--83) μm diam.,  dark brown, glabrous; peridium with a *textura angularis*. * Asci* ovoid to subglobose, (11--)11.5--14(--15) × (7--)7.5--9.5(--10.5) μm. * Ascospores* broad fusiform to ellipsoidal, (6.5--)7--8 × 4--5 μm, buff to honey, smooth, with a single inconspicuous germ pore. * Conidiophores* arising from substrate mycelium, indistinctive, simple or occasionally branched, bearing terminally a single annellide. * Annellides* lageniform to ampulliform, (8--)9--14.5(--17.5) μm long, (2--)2.5--3 μm broad at the widest part, tapering gradually to a cylindrical annellate zone 1--1.5(--2) μm wide, hyaline to subhyaline, smooth-walled. * Conidia* obovoid with truncate base, (4.5--)5--6(--6.5) × 3.5--4.5 μm, hyaline, turning to hazel when ageing, smooth or finely roughened, thick-walled, arranged in chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 21--24 mm after 14 d at 25 °C, flat, white to cream-coloured with greenish olivaceous to olivaceous buff and pale olivaceous grey zones, radially striated with dentate margin.  On MEA attaining a diameter of 18--19 mm, crateriform, pale olivaceous grey to olivaceous grey and greyish sepia, radially striated and folded in the centre, margin crenated.  On DG18 attaining a diameter of 13--14 mm, crateriform, buff to rosy buff, radially striated and folded in the centre, margin undulate.  On OA still growth at 36 °C, no growth at 40 °C. 

*Specimen examined*: **China**, Inner Mongolia, Ulanqab city, Huade county, from discarded cloth, 24 Jul. 2011, Y-Y Huo, (**holotype** HMAS 2444424, culture **ex-type** CBS 134638 = CGMCC 3.15222). 

*Notes*: The newly described *M. cleistocarpus* is closely related to *M. hyalinus* ([Fig.  1](#fig1){ref-type="fig"}). * Microascus cleistocarpus* and *M. hyalinus* are the only two species in *Microascus* producing cleistothecial ascocarps.  However, *M. hyalinus* produces hyaline conidia and *M. cleistocarpus* has hazel conidia.  Based on sequence data *M. cleistocarpus* can be distinguished from *M. hyalinus* on all three genes (ITS 3 nt difference, *tub2* 5 nt difference, *tef1* 10 nt difference). 

***Microascus fusisporus*** Woudenb. &  Samson **sp.  nov.**  MycoBank [MB818280](mycobank:818280){#intref0040}. [ Fig.  7](#fig7){ref-type="fig"}.Fig.  7*Microascus fusisporus* sp.  nov.  CBS 896.68. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D--G.** Conidiophores, annellides and conidia. ** H.** Conidia.  Scale bars = 10 μm. Fig.  7

*Etymology*: name refers to the fusiform conidia. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, simple or branched, occasionally indistinctive, bearing one to multiple annellides. * Annellides* ampulliform, (7--)9--12(--14) μm long, 2--3(--3.5) μm broad at the widest part, tapering gradually to a cylindrical annellate zone, sometimes thickened, 1--1.5(--2) μm wide, hyaline, smooth-walled. * Conidia* obovoid to broad clavate or fusiform, with truncate base, (5--)5.5--6.5 (--7) × 2.5--3.5 μm, hyaline or subhyaline, smooth, thick-walled, arranged in chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 15 mm after 14 d at 25 °C, flat, white at the margin with grey olivaceous to olivaceous centre, margin crenated.  On MEA attaining a diameter of 10--12 mm, convex, olivaceous grey with white to cream-coloured sectors and margin, margin crenated.  On DG18 attaining a diameter of 15--18 mm, crateriform, cinnamon with buff tufts of mycelium at the outer ring, olivaceous grey with white velvet mycelium at the centre, margin dentate.  On OA no growth at 36 and 40 °C. 

*Specimen examined*: **Germany**, Schleswig-Holstein, Kiel-Kitzeberg, from wheat-field soil, collection date unknown, K.H. Domsch & W. Gams, (**holotype** CBS H-22743, culture **ex-type** CBS 896.68 = ATCC 16278 = IFO 31244 = MUCL 8989). 

*Notes*: Morphologically *M. fusisporus* resembles *M. trautmannii*, but can be distinguish based on its shorter annellides (9--12 μm long in *M. fusisporus* against 16--22 μm long in *M. trautmannii*) and the ability to grow on OA at 36 °C of *M. trautmannii*.  Both species can easily be distinguished from the other *M. paisii*-like species based on their obovoid to broad clavate or fusiform conidia with truncate base. 

***Microascus hollandicus*** Woudenb. &  Samson **sp.  nov.**  MycoBank [MB818279](mycobank:818279){#intref0045}. [ Fig.  8](#fig8){ref-type="fig"}.Fig.  8*Microascus hollandicus* sp.  nov.  CBS 141582. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D--F.** Conidiophores, annellides and conidia. ** H.** Conidia.  Scale bars = 10 μm. Fig.  8

*Etymology*: name refers to the country of isolation, the Netherlands. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, simple or indistinctive, bearing one or multiple annellides. * Annellides* ampulliform, (3.5--)4--6(--8) μm long, (2.0--)2.5--3(--3.5) μm broad at the widest part, tapering abruptly to a cylindrical annellate zone 1--1.5 μm wide, hyaline, smooth-walled. * Conidia* broadly ellipsoidal to short clavate with truncate base, (3.5--)4--4.5(--5) × (2.5--)3--3.5(--4) μm, hyaline when young turning honey when ageing, smooth, thick-walled, arranged in chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 17--18 mm after 14 d at 25 °C, flat to slightly raised, white to cream-coloured with olivaceous grey to olivaeous buff centre, margin dentate.  On MEA attaining a diameter of 12--13 mm, raised, white to very pale olivaceous grey, radially striated with crenated margin.  On DG18 attaining a diameter of 10--11 mm, raised, olivaceous grey, woolly with long white mycelium hairs growing out, margin entire.  On OA no growth at 36 and 40 °C. 

*Specimen examined*: **Netherlands**, from a swab sample of an indoor horse arena, Mar. 2012, Houba, (**holotype** CBS H-22716, culture **ex-type** CBS 141582). 

*Notes*: *Microascus hollandicus* morphologically resembles *M. pseudopaisii*.  Sequence data is necessary to distinguish both species.  All three genes used in this manuscript can separate the two species (ITS 8 nt difference, *tub2* 21 nt difference, and *tef1* 10 nt difference between the type isolates of both species). * Microascus hollandicus* and *M. pseudopaisii* can be differentiated from the other *M. paisii*-like species by their shorter annellides (4--6 μm long). 

***Microascus melanosporus*** (Udagawa) Woudenb. &  Samson **comb.  nov.**  MycoBank [MB817657](mycobank:817657){#intref0050}. [ Fig.  9](#fig9){ref-type="fig"}.Fig.  9*Microascus melanosporus* comb.  nov.  DTO 255-B1. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D--F.** Conidiophores, annellides and conidia. ** G.** Conidia.  Scale bars = 10 μm. Fig.  9

*Basionym*: *Scopulariopsis melanospora* Udagawa, J. agric.  Sci. ( Tokyo) 5: 18.  1959. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, simple or indistinctive, bearing one to multiple annellides. * Annellides* ampulliform, (5.5--)7.5--11(--13) μm long, (2--)2.5--3.5(--4) μm broad at the widest part, tapering abruptly to a cylindrical annellate zone 1--1.5(--2) μm wide, hyaline, smooth-walled. * Conidia* broadly ellipsoidal to short clavate with truncate base, 4--4.5(--5) × (2.5--)3--3.5 μm, hyaline when young turning hazel when ageing, smooth, thick-walled, arranged in chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 21--25 mm after 14 d at 25 °C, low convex, white to cream-coloured with olivaceous grey to iron grey zones, margin crenated.  On MEA attaining a diameter of 19--23 mm, convex or crateriform, pale olivaceous grey to olivaceous grey, margin undulate.  On DG18 attaining a diameter of 20--24 mm, crateriform, white to cream-coloured with grey olivaceous to greyish sepia and vinaceous grey zones, margin undulate.  On OA some isolates grow at 36 °C, no growth at 40 °C. 

*Specimens examined*: **Germany**, from indoor air sample, 2013, C. Trautmann, DTO 255-A5; from indoor air sample, 2013, C. Trautmann, DTO 255-A7; from plaster, 2013, C. Trautmann, DTO 255-B1. ** South Africa**, Somerset West, from house dust, 12 Feb. 2009, Karin Jacobs, DTO 220-H9. ** USA**, from milled *Oryza sativa*, 1955, S. Udagawa, (culture **ex-type** CBS 272.60 = MUCL 9040 = IMI 078257). 

*Notes*: *Microascus melanosporus* is morphologically and phylogenetically closely related to *M. paisii*.  Morphologically it can be differentiated from the other *M. paisii*-like species by its faster growth on OA and MEA at 25 °C (21--25 mm versus 16--20 mm and 19--23 versus 15--18 mm in diam.  respectively).  Based on sequence data both the *tub2* and *tef1* can separate *M. melanosporus* from the other *M. paisii*-like species.  The ITS sequence is identical to *M. paisii*. 

***Microascus micronesiensis*** Woudenb.,  Seifert & Samson **sp.  nov.**  MycoBank [MB818281](mycobank:818281){#intref0055}. [ Fig.  10](#fig10){ref-type="fig"}.Fig.  10*Microascus micronesiensis* sp.  nov.  CBS 141523. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D--G.** Conidiophores, annellides and conidia. ** H.** Conidia.  Scale bars = 10 μm. Fig.  10

*Etymology*: name refers to the country of isolation, Micronesia. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, indistinctive, simple or branched, bearing terminally one or occasionally two annellides. * Annellides* ampulliform, (6--)7--11(--14) μm long, 2--2.5(--3) μm broad at the widest part, tapering gradually to a cylindrical annellate zone (0.5--)1--1.5 μm wide, hyaline, smooth-walled. * Conidia* broadly obovoid with truncate base, 3--4(--4.5) × (2--)2.5--3(--3.5) μm, hyaline or subhyaline, smooth, thick-walled, arranged in chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 15--17 mm after 14 d at 25 °C, flat, white to cream-coloured with pale grey olivaceous to grey olivaceous rings, margin undulated.  On MEA attaining a diameter of 16--17 mm, low convex, white to cream-coloured, margin undulate.  On DG18 attaining a diameter of 10--11 mm, low convex, white to cream-coloured, margin undulate.  On OA reduced growth at 36, no growth at 40 °C. 

*Specimens examined*: **Micronesia**, Kosrae, Kosrae Island, Malem, from house dust, 15 Mar. 2009, Wayne Law, (**holotype** CBS H-22739 culture **ex-type** CBS 141523); Kosrae, Kosrae Island, Tofol, from house dust, 2009, Wayne Law, DTO 223-A5. 

*Notes*: Phylogenetically *M. micronesiensis* is closely related to the two recently described sexual species *M. brunneosporus* ([@bib29]) and *M. chinensis* ([@bib14]).  Morphologically *M. micronesiensis* can be distinguished from *M. brunneosporus* and *M. chinensis* by the lack of producing sexual structures in culture and its much slower growth on OA at 25 °C (15--17 mm for *M. micronesiensis* versus 21--25 and 25--28 mm for *M. brunneosporus* and *M. chinensis* respectively after 14 d). * Microascus micronesiensis* has been found in house-dust samples from two different houses in Micronesia in different cities on separate occasions. 

***Microascus paisii*** (Pollacci) Sandoval-Denis, Gené & Guarro, Persoonia 36:21.  2016. [ Fig.  11](#fig11){ref-type="fig"}.Fig.  11*Microascus paisii* DTO 255-B2. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D--G.** Conidiophores, annellides and conidia. ** H.** Conidia.  Scale bars = 10 μm. Fig.  11

*Basionym*: *Torula paisii* Pollacci (as 'pais'), Atti Ist.  Bot.  Univ.  Pavia, ser.  2, 18:130.  1921. 

≡ *Phaeoscopulariopsis paisii* (Pollacci) M. Ota, Jap.  J. Dermatol.  Urol.  28:5.  1928.  nom.  inval. 

≡ *Scopulariopsis paisii* (Pollacci) Nann.,  Repert.  Sist.  dei Miceti dell\'Uomo e degli Anim.:  259.  1934. 

= *Scopulariopsis sphaerospora* Zach, Oesterr.  Bot.  Z. 83: 180.  1934. 

= *Scopulariopsis brumptii* Salv.-Duval, Thèse Fac.  Pharm.  Paris.  23: 58.  1935. 

= *Scopulariopsis versicolor* Salv.-Duval, Thèse Fac.  Pharm.  Paris 23: 63.1935. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, simple or indistinctive, bearing one or multiple annellides. * Annellides* ampulliform, (5.5--)6.5--9.5(--12) μm long, (2--)2.5--3(--3.5) μm broad at the widest part, tapering abruptly to a cylindrical annellate zone 1--1.5(--2) μm wide, hyaline, smooth-walled. * Conidia* broadly ellipsoidal to short clavate with truncate base, 3.5--4(--4.5) × 3--3.5(--4) μm, hyaline when young turning hazel when ageing, smooth, thick-walled, arranged in chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 17--20 mm after 14 d at 25 °C, low convex, white to cream-coloured with olivaceous grey to iron grey centre, margin crenated.  On MEA attaining a diameter of 15--18 mm, crateriform, olivaceous grey to iron grey, radially striated, margin crenated.  On DG18 attaining a diameter of 19--22 mm, crateriform, vinaceous buff with purplish grey zones at the margin and greyish sepia centre, margin entire.  On OA some isolates grow at 36 °C, no growth at 40 °C. 

*Specimens examined*: **Austria**, from unknown substrate, 1934, F. Zach, (*S. sphaerospora* culture **ex-type** CBS 402.34 = MUCL 9045). ** France**, from small-pox vaccine, 1935, M. Langeron, (probably *S. brumptii* culture **ex-type** CBS 333.35). ** Germany**, from plaster, 2013, C. Trautmann, DTO 255-B2; from oriented strand board, 2013, C. Trautmann, DTO 255-B8. ** Italy**, from human, 1927, G. Pollacci (*T. paisii* culture **ex-type** CBS 213.27 = MUCL 7915). 

*Notes*: The new name combination *Microascus paisii* for *Torula paisii* was recently proposed, together with the synonymy of several well-known species underneath it ([@bib29]).  In this manuscript two synonymies are reinstated, namely *Masonia grisea* as *Microascus atrogriseus* and *Scopulariopsis melanospora* as *Microascus melanosporus*, and four new species are described, *M. fusisporus*, *M. hollandicus*, *M. pseudopaisii* and *M. trautmannii*.  Morphologically *M. fusisporus* and *M. trautmannii* can be distinguished from *M. paisii* by their shape of conidia (see notes of the respective species), and *M. pseudopaisii* and *M. hollandicus* by their shorter annellides (see notes of the respective species). * Microascus melanosporus* can be distinguished by its faster growth rate on OA and MEA at 25 °C (see notes of *M. melanosporus*). * Microascus atrogriseus* is morphological identical to *M. paisii*, molecular data is necessary to distinguish the species (see notes of *M. atrogriseus*).  CBS 333.35 isolated from small-pox vaccine in France is recognised here as the probable ex-type isolate of *S. brumpti*.  It was deposited to the CBS in 1935 by Prof. Dr. Langeron who worked in the Université de Paris, Faculté de Médecine.  The original description of *S. brumptii* by Salvanet-Duval was published in Thése Faculté de Pharmacie at the Université de Paris, on research on the small-pox vaccine ([@bib27]).  All three studied ex-type isolates (CBS 213.27, CBS 402.34 and CBS 333.35) showed reduced growth and are therefore excluded from the culture descriptions. 

***Microascus pseudopaisii*** Woudenb. &  Samson **sp.  nov.**  MycoBank [MB818282](mycobank:818282){#intref0060}. [ Fig.  12](#fig12){ref-type="fig"}.Fig.  12*Microascus pseudopaisii* sp.  nov.  CBS 141581. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D--H.** Conidiophores, annellides and conidia.  Scale bars = 10 μm. Fig.  12

*Etymology*: name refers to the morphological and phylogenetic close relationship to *M. paisii*. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, simple or branched, occasionally indistinctive, bearing one to multiple annellides. * Annellides* lageniform to ampulliform, (3.5--)4.5--6(--6.5) μm long, 2--3(--3.5) μm broad at the widest part, tapering abruptly to a cylindrical annellate zone 1--1.5 μm wide, hyaline, smooth-walled. * Conidia* broadly ellipsoidal to short clavate with truncate base, (3--)3.5--4.5(--5) × 2.5--3(--3.5) μm, hyaline when young turning honey when ageing, smooth, thick-walled, arranged in chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 17--20 mm after 14 d at 25 °C, slightly raised, white to cream-coloured with olivaceous grey centre, margin crenated.  On MEA attaining a diameter of 14--15 mm, crateriform, olivaceous grey with white to cream-coloured edge, radially striated with crenated margin.  On DG18 attaining a diameter of 10--12 mm, crateriform, pale olivaceous grey, woolly with long white mycelium hairs growing out, margin entire.  On OA no growth at 36 and 40 °C. 

*Specimens examined*: **Netherlands**, Nederwetten, from an air sample of the basement of a house, 16 Dec. 2009, J. Houbraken, (**holotype** CBS H-22715, culture **ex-type** CBS 141581); additional strain from the same source, DTO 116-A4. 

*Notes*: *Microascus pseudopaisii* morphologically resembles *M. hollandicus*.  Sequence data is necessary to distinguish both species (see notes *M. hollandicus*). * Microascus hollandicus* and *M. pseudopaisii* can be differentiated from the other *M. paisii*-like species by their shorter annellides (4--6 μm long). 

***Microascus trautmannii*** Woudenb. &  Samson **sp.  nov.**  MycoBank [MB818283](mycobank:818283){#intref0065}. [ Fig.  13](#fig13){ref-type="fig"}.Fig.  13*Microascus trautmannii* sp.  nov.  CBS 141583. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D--G.** Conidiophores, annellides and conidia. ** H.** Conidia.  Scale bars = 10 μm. Fig.  13

*Etymology*: named after Dr. Christoph Trautmann, who collected numerous *Microascus* isolates from the indoor environment. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, simple or indistinctive, bearing terminally one or multiple annellides. * Annellides* slender ampulliform, (13.5--)16--22(--25) μm long, (1.5--)2--2.5(--3) μm broad at the widest part, with a sometimes thickened cylindrical annellate zone (1--)1.5--2(--2.5) μm wide, hyaline, smooth-walled. * Conidia* obovoid to broad clavate or fusiform, with truncate base, (5--)5.5--6.5(--7) × (2--)2.5--3 μm, hyaline or subhyaline, smooth, thick-walled, arranged in chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 17 mm after 14 d at 25 °C, flat to slightly raised, white to cream-coloured with olivaceous grey centre, margin dentate.  On MEA attaining a diameter of 13--16 mm, raised, buff to (pale) olivaceous grey with zones of white woolly mycelium, edge radially striated with crenated margin.  On DG18 attaining a diameter of 10--11 mm, crateriform, pale olivaceous grey to pale greenish grey, margin entire.  On OA reduced growth at 36 °C, no growth at 40 °C. 

*Specimen examined*: **Germany**, from oriented strand board, C. Trautmann, 2013 (**holotype** CBS H-22717, culture **ex-type** CBS 141583). 

*Notes*: Morphologically *M. trautmannii* resembles *M. fusisporus*, but they can be distinguished based on the size of their annellides and the growth at 36 °C on OA (see notes *M. fusisporus*).  Both species can easily be distinguished from the other *M. paisii*-like species based on their obovoid to broad clavate or fusiform conidia with truncate base. 

***Scopulariopsis africana*** Woudenb. &  Samson **sp.  nov.**  MycoBank [MB818274](mycobank:818274){#intref0070}. [ Fig.  14](#fig14){ref-type="fig"}.Fig.  14*Scopulariopsis africana* sp.  nov.  CBS 118736. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D--F.** Conidiophores, annellides and conidia. ** G--H.** Conidia.  Scale bars = 10 μm. Fig.  14

*Etymology*: name refers to the country of isolation, South Africa. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, simple to indistinctive, occasionally branched. * Annellides* cylindrical to slight ampulliform, (5--)8--15(--21.5) μm long, (2--)3--4(--4.5) μm broad at the widest part, tapering gradually to a cylindrical annellate zone 2-- 3(--3.5) μm wide, hyaline, smooth-walled. * Conidia* subglobose to broadly ovoid with truncate base, (5.5--)6--7(--8) × (4--)4.5--5.5(--6) μm, hyaline, smooth, thick-walled, arranged in chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 35 mm after 14 d at 25 °C, flat, white to cream-coloured with olivaceous zones, margin crenated.  On MEA attaining a diameter of 12--14 mm, crateriform, white to cream-coloured, folded, margin crenated.  On DG18 attaining a diameter of 25--28 mm, low convex, white to cream-coloured, margin undulate to erose to fimbriate.  On OA no growth at 36 and 40 °C. 

*Specimen examined*: **South Africa**, Free State, Lemoenskloof, from mud sample from salt pan, before Sep. 2004, M.E. Setati, (**holotype** CBS H-22741, culture **ex-type** CBS 118736). 

*Notes*: Morphologically *S. africana* resembles *S. albida*, *S. candida* and *S. alboflavescens*, although *S*. * africana* shows olivaceous zones on OA (*S. albida* and *S. candida* are characterised by white colonies, *S. alboflavescens* by white-cream to pale yellowish colonies).  Molecularly, *S. africana* can be distinguished from other *Scopulariopsis* species based on its *tub2* and *tef1* sequence. 

***Scopulariopsis albida*** Woudenb. &  Samson **sp.  nov.**  MycoBank [MB818275](mycobank:818275){#intref0075}. [ Fig.  15](#fig15){ref-type="fig"}.Fig.  15*Scopulariopsis albida* sp.  nov.  CBS 119.43. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D--G.** Conidiophores, annellides and conidia. ** H.** Conidia.  Scale bars = 10 μm. Fig.  15

*Etymology*: name refers to the white colonies. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, simple to indistinctive. * Annellides* cylindrical to slight ampulliform, (6)8.5--19.5(--29.5) μm long, (2.5--)3--5(5.5) μm broad at the widest part, annellate zone (2--)2.5--3.5(--4) μm wide, single, hyaline, smooth-walled. * Conidia* globose to subglobose with truncate base, (5--)6.5--8(--8.5) × (6--)6.5--7.5(--8) μm, hyaline, smooth, thick-walled, arranged in chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 55--60 mm after 14 d at 25 °C, flat, white to cream-coloured, margin entire.  On MEA attaining a diameter of 20--23 mm, crateriform, white to cream-coloured, folded, margin undulate.  On DG18 attaining a diameter of 21--25 mm, low convex, white to cream-coloured, folded, margin undulate to erose to fimbriate.  On OA no growth at 36 and 40 °C. 

*Specimens examined*: **Netherlands**, from soil, collection date and collector unknown (**holotype** CBS H-22740, culture **ex-type** CBS 119.43). ** Germany**, substrate and collection date unknown, P. Höhle, CBS 415.51. 

*Notes*: Morphologically *S. albida* resembles *S. candida*.  Although *S. candida* does not form a monophyletic clade, *S. albida* can molecularly be distinguished from *S. candida* based on its *tub2* and *tef1* sequence.  Also *S. africana* and *S. alboflavescens* morphologically resemble *S. albida*.  Here the olivaceous zones on OA of *S. africana* isolates and the cream-white to pale yellowish colonies and subhyaline conidia of *S. alboflavescens* isolates can be used to distinguish the species. 

***Scopulariopsis caseicola*** Woudenb. &  Samson **sp.  nov.**  MycoBank [MB818276](mycobank:818276){#intref0080}. [ Fig.  16](#fig16){ref-type="fig"}.Fig.  16*Scopulariopsis caseicola* sp.  nov.  CBS 480.62. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D--G.** Conidiophores, annellides and conidia. ** H.** Conidia.  Scale bars = 10 μm. Fig.  16

*Etymology*: name refers to the substrate of isolation, cheese. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, simple or branched, bearing terminally a single annellide (at each branch). * Annellides* cylindrical, (10.5--)22.5--47.5(--67.5) × (2.5--)3--4(--5) μm, tapering gradually to a cylindrical annellate zone 2--3.5(--4) μm wide, subhyaline becoming darker with age, smooth-walled. * Conidia* broad ovoid with truncate base, (4.5--)6--7(--8) × (4--)5--6(--7) μm, buff to honey, smooth, thick-walled, arranged in long chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 30 mm after 14 d at 25 °C, flat, white to opaque, margin entire.  On MEA attaining a diameter of 22--23 mm, low convex, white to cream-coloured, margin undulate.  On DG18 attaining a diameter of 7 mm, flat, pale olivaceous grey to smoke grey, margin fimbriate.  On OA no growth at 36 and 40 °C. 

*Specimen examined*: **Netherlands**, from cheese-coating, collection date unknown, M.B. Schol-Schwarz, (**holotype** CBS H-22738, culture **ex-type** CBS 480.62). 

*Note*: Sporulation was only observed on OA after 2 months cultivation, re-isolation of a fresh culture might influence the morphological description. [ @bib19] discussed the synonyms of *S. flava*, a species frequently found on cheese.  Among the synonyms they also discussed *S. casei* Loubière of which no type material is known to exist. 

***Scopulariopsis sexualis*** Woudenb. &  Samson **sp.  nov.**  MycoBank [MB818277](mycobank:818277){#intref0085}. [ Fig.  17](#fig17){ref-type="fig"}.Fig.  17*Scopulariopsis sexualis* sp.  nov.  CBS 250.64. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D.** Ascomata. ** E.** Ascomatal wall. ** F.** Asci. ** G.** Ascospores.  Scale bars = 10 μm. Fig.  17

*Etymology*: name refers to the presence of only sexual structures and lack of asexual structures. 

*Ascomata* abundant, superficial or immersed, ostiolate, globose with a short cylindrical ostiolar neck (up to 20 μm) or ovoid, 108--145(171) μm diam.,  dark brown to black, glabrous; peridium with a *textura angularis*. * Asci* irregularly ellipsoidal, (9.5--)10.5--12(--13) × 9--11.5(--13) μm. * Ascospores* reniform to broadly lunate, (4.5--)5--5.5(--6.5) × 3.5--4.5(--5) μm, buff to honey, luteous to orange in mass, smooth, with a single inconspicuous germ pore.  Asexual morph not observed. 

*Culture characteristics*: Colonies on OA attaining a diameter of 60--67 mm after 14 d at 25 °C, flat, dull green with white edge, margin entire.  On MEA attaining a diameter of 47--57 mm, crateriform, white to cream-coloured with olivaceous grey to iron grey centre, radially striated with entire margin.  On DG18 attaining a diameter of 38--40 mm, flat, white to cream-coloured, margin undulate to crenated.  On OA still growth at 36 °C, no growth at 40 °C. 

*Specimens examined*: **Burma**, from milled rice, 1954, S. Udagawa, (**holotype** CBS H-14445, culture **ex-type** CBS 250.64 = IFO 7555 = UAMH 1923 = NHL 2278). ** India**, Delhi, from seed of *Brassica oleracea* (*Brassicaceae*), collection date unknown, K.G. Mukerji, CBS 332.78. ** USA**, Arizona, Tucson, from bat dung, collection date unknown, G.F. Orr, CBS 667.71 = NRRL A-8022. 

*Note*: Morphologically *S. sexualis* resembles the sexual morph of *S. cordiae*. * Scopulariopsis sexualis* can be differentiated from *S. cordiae* by its much faster growth on OA (60--70 mm for *S. sexualis*, 35--36 mm for *S. cordiae* at 25 °C after 14 d) and shorter cylindrical ostiolar neck (*S. sexualis* up to 20 μm, *S. cordiae* up to 390 μm). 

***Yunnania*** H.Z. Kong, Mycotaxon 69: 320.  1998. 

= *Fuscoannellis* Sandoval-Denis, Jagielski, Jin Yu & Gené, Fungal Biol.  120: 593.  2016. 

***Yunnania carbonaria*** (F.J. Morton & G. Sm.)  Woudenb.,  Houbraken & Samson **comb.  nov.**  MycoBank [MB820189](mycobank:820189){#intref0090}. 

*Basionym*: *Scopulariopsis carbonaria* F.J. Morton & G. Sm.,  Mycol.  Pap.  86: 59.  1963. 

≡ *Fuscoannellis carbonaria* (F.J. Morton & G. Sm.)  Sandoval-Denis, Jagielski, Jin Yu & Gené, Fungal Biol.  120: 593.  2016. 

*Specimens examined*: **Panama**, from soil, collection date unknown, R. Cogill, (culture **ex-type** CBS 205.61 = NRRL 1860 = IFO 8116 = MUCL 9027 = IMI 086941); **USA**, Hawaii, on dead hardwood branch, 3 Nov. 2002, D.T. Wicklow, CBS 121662. 

***Yunnania smithii*** Woudenb. &  Samson **sp.  nov.**  MycoBank [MB818273](mycobank:818273){#intref0095}. [ Fig.  18](#fig18){ref-type="fig"}.Fig.  18*Yunnania smithii* sp.  nov.  CBS 855.68. ** A--C.** Fourteen day old colonies on OA (A), MEA (B) and DG18 (C). ** D--F.** Conidiophores, annellides and conidia. ** G--H.** Conidia.  Scale bars = 10 μm. Fig.  18

*Etymology*: named after late George Smith (1895--1967), a British mycologist who extensively studied the closely related fungal genera *Microascus* and *Scopulariopsis*, and collected the type isolate. 

Sexual morph not observed. * Conidiophores* arising from substrate mycelium, frequently branched, bearing terminally a group of annellides (at each branch). * Annellides* ampulliform, (4--)5--7 μm long, 2--2.5(--3) μm broad at the widest part, with a short annellate zone, (1--)1.5--2 μm wide, hyaline to subhyaline, smooth-walled. * Conidia* ovoid to ellipsoidal with truncate base, 4--5(--5.5) × 2--2.5(--3) μm, vinaceous buff to hazel, smooth, thick-walled, arranged in (long) chains. 

*Culture characteristics*: Colonies on OA attaining a diameter of 33--35 mm after 14 d at 25 °C, flat, olivaceous to olivaceous grey, margin crenated.  On MEA attaining a diameter of 26--30 mm, crateriform, greyish blue to (pale) olivaceous grey and slate blue, radially striated with dentate margin.  On DG18 attaining a diameter of 29--30 mm, crateriform, pale olivaceous grey to olivaceous grey, radially striated, margin entire.  On OA no growth at 36 and 40 °C. 

*Specimen examined*: **Germany**, Kiel-Kitzeberg, from garden soil, 1963, G. Smith, (**holotype** CBS H-22742, culture **ex-type** CBS 855.68). 

*Notes*: *Yunnania smithii* morphologically resembles the other two species in the genus *Yunnania*, *Y. carbonaria* and *Y. penicillata*.  It can be distinguished from *Y. penicillata* by its faster growth on OA in 7 d (*Y. smithii* 33--35 mm, *Y. penicillata* 20 mm).  The colour of the conidiophores and annellides can be used to distinguish it from *Y. carbonaria* (hyaline in *Y. smithii*, pale brown to brown in *Y. carbonaria*).  Molecularly *Y. smithii* can best be distinguished based on its *tef1* sequence (19 nt difference *Y*. * carbonaria*, 19 nt difference *Y. penicillata*), followed by its *tub2* sequence (12 nt difference *Y*. * carbonaria*, 14 nt difference *Y. penicillata*).  The LSU and ITS sequences are not suited for identification, the LSU sequences are all 100 % identical and the ITS sequence of *Y. smithii* is identical to *Y. penicillata* and has only 1 nt difference with the type isolate of *Y. carbonaria*. 

Discussion {#sec5}
==========

This manuscript presents a molecular phylogenetic study of species in the genera *Microascus* and *Scopulariopsis* known from culture, with the intention to identify the common indoor species.  Since fungi present in indoor environments can produce toxins or carry allergens which cause health hazards, it is important to know which fungal species are present indoors. * Scopulariopsis* and scopulariopsis-like species are mainly found in soil, but also frequently isolated from food and building materials like drywall paper and wood ([@bib28]).  Little is known about the health effects of these fungi, although several species seem to be able to cause human onychomycosis and superficial tissue infections (*e.g*. [ @bib33], [@bib35]).  Rare cases of more severe diseases are reported, but only in immunocompromised patients (*e.g*. [ @bib2], [@bib18]).  The ability of *Scopulariopsis* species to deteriorate building materials ([@bib9], [@bib16]), and to accumulate various elements and turning these into toxic volatiles ([@bib4], [@bib3]) also makes them an important group to study in the indoor environment. 

As stated in the results section, 17 species are mentioned in this manuscript to occur in the indoor environment ([Table 2](#tbl2){ref-type="table"}), but most of them are only occasionally found indoors.  Besides the number of isolates found indoors, the substrate of isolation should also be taken into consideration when labelling species as indoor species.  The isolates assigned to the indoor habitat include swab samples and house dust or air samples.  Since swab sample are mostly taken from sites suspicious of fungal growth, they can (often) be related to actual indoor growth.  A dust or air sample only implies the presence of the fungus.  Since the concentration of fungal spores in the indoor air is to certain extent dependent on the outside spore concentration, it is recommended to also sample the outside air for comparison ([@bib28]).  This information is not known for our isolates, and also information on the abundance of the species in the air or dust sample is unknown.  Ten of the species which are mentioned here to occur in the indoor environment are also found in relation with humans ([Fig.  3](#fig3){ref-type="fig"}).  However, isolates assigned in this study to the human habitat are mainly isolated from human-derived specimens, which is merely an indication of a possible pathogenic role.  For the majority of the species known from human specimens there is no proven relation with disease ([@bib30]).  Especially for isolates obtained from superficial sites and the upper respiratory tracts it should be taken into account that these can be environmental contaminants.  Another point of attention is the indoor environment in which the species are found.  How much time people spend in the different indoor environments (archives, homes, offices, stables, animal pens, etc.)  varies, although all of them are treated here as indoor habitat.  As example we will take *M. alveolaris*, only 1 out of 9 studied isolates is isolated from the indoor environment.  This indoor isolate came from a house-dust sample, but no additional information is known on the abundance of the fungus in the dust sample.  Additional information is needed to label *M. alveolaris* as true indoor species, although this study shows it can be found in homes.  One other *M. alveolaris* isolate was human-derived, although an earlier study linked multiple human-derived isolates to this species ([@bib29]).  Most of them are bronchoalveolar lavage isolates, as the name of the fungus already applies.  Although for most of these isolates their relation with disease is not known, the finding of multiple isolates from the respiratory tract of human patients, and the ability of the species to grow at 40 °C are good indications of the potential pathogenicity of the species. 

The most commonly found indoor species, both in swab and air/dust samples are *M. melanosporus*, *M. paisii*, *S. brevicaulis* and *S. candida*.  All four are also placed in relation with the human habitat. * Scopulariopsis brevicaulis* and *S. candida* are known to be involved in onychomycosis, and *S. brevicaulis* is also recognised as important human opportunistic pathogens, as well as *S. brumptii* (now *M. paisii*) ([@bib7]).  For *M. melanosporus*, which was previously regarded a synonym of *S. brumptii* ([@bib19], as *S. melanospora*), the pathogenic abilities are unknown.  However, one can expect that it can also act as opportunistic pathogen, as it close relatives, especially with the ability of some isolates to grow at 36 °C. 

Based on the multi-gene phylogeny ([Fig.  2](#fig2){ref-type="fig"}) and congruent single gene trees ([Fig.  19](#fig19){ref-type="fig"}) the newly combined *M. paisii* ([@bib29]) is split in this study into seven species of which four (*M. fusisporus*, *M. hollandicus*, *M. pseudopaisii*, and *M. trautmannii*) are newly described, one was given a new name (*M. atrogriseus*) and one a new name combination (*M. melanosporus*). * Scopulariopsis brumptii* is still regarded as synonym of *M. paisii* ([Table 3](#tbl3){ref-type="table"}).  Based on their *tub2* and *tef1* sequences all seven *M. paisii*-like species can be molecularly identified ([Fig.  19](#fig19){ref-type="fig"}). * Microascus melanosporus* seems to be the most prevalent species found indoor, and *M. paisii* is recognised as second most common indoor *Microascus* species.  For the studied isolates *M. melanosporus* can morphologically be distinguished from *M. paisii* based on the growth rate and colony colour after 2 wk incubation on OA at 25 °C ([Fig.  20](#fig20){ref-type="fig"}). * Microascus melanosporus* grows slightly faster than *M. paisii* (21--25 mm versus 16--20 mm in diam.  respectively), and *M. melanosporus* has slightly lighter grey colonies than *M. paisii* ([Fig.  20](#fig20){ref-type="fig"}A, B). * Microascus fusisporus* and *M. trautmannii* can be distinguished from the other *M. paisii*-like species based on their obovoid to broad clavate or fusiform conidia versus the broadly ellipsoidal to short clavate conidia from the other *M. paisii-*like isolates. * Microascus hollandicus* and *M. pseudopaisii* can be distinguished by their shorter annellides (4--6 μm long versus 6--11 μm long on average for the other *M. paisii-*like isolates).  To distinguish *M. atrogriseus* from *M. paisii* and *M. hollandicus* from *M. pseudopaisii* molecular data is needed.  The two most common indoor *Scopulariopsis* species, *S. brevicaulis* and *S. candida*, are both molecularly as morphologically easy to distinguish.  Morphologically the growth rate and colony colour after 2 wk incubation on OA at 25 °C, and the conidia morphology can be used to distinguish the species ([Fig.  20](#fig20){ref-type="fig"}). * Scopulariopsis brevicaulis* grows faster than *S. candida* (75 mm and 38--48 mm in diam.  respectively), and *S. brevicaulis* has buff to rosy buff colonies versus white colonies in *S. candida* ([Fig.  20](#fig20){ref-type="fig"}C, D).  Another distinction are the roughened conidia of *S. brevicaulis*, versus the smooth conidia of *S. candida* ([Fig.  20](#fig20){ref-type="fig"}G, H).  A growth test at 36 °C can also be used to distinguish the species since *S. brevicaulis* is able to grow at 36 °C and *S. candida* is not. Fig.  19Maximum likelihood trees based on respectively the ITS, *tub2* or *tef1* sequences of 56 isolates, representing the *Microascus paisii* clade and the out-group isolate *M. hyalinus* (CBS 766.70).  The RAxML bootstrap support values ≥75 % (BS) and Bayesian posterior probabilities ≥0.95 (PP) are given at the nodes.  Thickened lines indicate a BS of 100 % and a PP of 1.0.  Species names between parentheses indicate the ex-type isolates of those species names.  The red and orange printed isolates represent *M. melanosporus*, the blue printed isolates represent *M. paisii*, the green printed isolates represent *M. atrogriseus* and the pink printed isolates represent *M. pseudopaisii*.Fig.  19Fig.  20Most common indoor *Microascus* and *Scopulariopsis* species. ** A, E.***M. melanosporus* DTO 255-B1. ** B, F.***M. paisii* DTO 255-B2. ** C, G.***S. brevicaulis* CBS 118474. ** D, H.***S. candida* DTO 138-B7. ** A--D.** Fourteen day old colonies on OA. ** E--H.** Conidiophores, annellides and conidia.  Scale bars = 10 μm. Fig.  20

Below we will discuss some phylogenetic unclarities which we encountered during this molecular phylogenetic study of *Scopulariopsis* and scopulariopsis-like species. 

The phylogenetic position of *M. longirostris* and *M. pseudolongirostris* is doubtful.  In our *Microascus* phylogeny, based on the ITS, *tub2* and *tef1* sequences, *M. longirostris* and *M. pseudolongirostris* cluster closest to the genus *Yunnania* rather than *Microascus* but without phylogenetic support ([Fig.  2](#fig2){ref-type="fig"}).  In the genus tree, based on the LSU, ITS and *tef1* sequences, *M. longirostris* and *M. pseudolongirostris* cluster closest to *Microascus*, although again without phylogenetic support ([Fig.  3](#fig3){ref-type="fig"}).  We choose to keep them in the genus *Microascus* following earlier publications ([@bib14], [@bib29]) although they will need further study.  The genus *Yunnania* is supported as separate genus in the genera tree, which is congruent with the study of [@bib14].  They already stated that the *S. carbonaria* isolates did not belong to *Scopulariopsis*, and proposed the new genus *Fuscoannellis*.  However, since the genus *Yunnania* was described earlier ([@bib15]), this genus name has priority according to the rules of the ICN and *Fuscoannellis* will become a synonym of *Yunnania*.  The CBS collection contained four isolates named *Scopulariopsis carbonaria*, including the type isolate CBS 205.61.  However, the type isolate only clustered together with CBS 121662 ([Fig.  2](#fig2){ref-type="fig"}), which was originally stored as *S. brumptii* in the CBS collection (as was already noticed by [@bib14]).  Two other '*S. carbonaria*' isolates, CBS 687.68 and 253.69, cluster together in the genus *Kernia* (data not shown).  CBS 855.68 does cluster with the type isolate of *S. carbonaria* based on its ITS sequence (only 1 nt difference), but based on its *tub2* (12 nt difference) and *tef1* (19 nt difference) sequence, in combination with morphological study, we describe it here as a new species *Y. smithii*.  These "*S. carbonaria*" isolates form a good example of the problems with morphological identification in these genera.  The isolate DTO 223-A6 clusters close to the type isolates of the recent described species *M. intricatus* and *M. onychoides*.  These two species have identical ITS sequences, but differ 9 nt in their *tub2* sequences, and 10 nt in their *tef1* sequences.  Isolate DTO 223-A6 clusters closest to *M. onychoides*, although it is not 100 % molecularly identical.  Also based on morphology we could not clearly place DTO 223-A6 in one of the two species, since the measurements of the spores did not exactly match one of them.  We choose however to name DTO 223-A6 *M. onychoides* for now, but collection of more isolates will be necessary to establish the species boundaries of *M. intricatus* and *M. onychoides*.  The phylogenetic species *M. terreus* contains two supported clades ([Fig.  2](#fig2){ref-type="fig"}).  Because the isolates in these two clades are morphologically identical, and are isolated from the same substrates, we choose to keep them as one species.  Also in the species *M. croci* there is some sequence variation.  Isolates CBS 158.44 and DTO 220-I5 deviate from the other *M. croci* isolates in their ITS sequence only (3 nt in a short stretch).  DTO 305-B5 deviates on all three loci from the other isolates (ITS 3 nt, *tub2* 6 nt and *tef* 8 nt difference).  We choose to keep it as a *M. croci* for now, since there is no support in the tree for the split.  With the collection of more isolates, *M. croci* might split into two or maybe three *Microascus* species.  The new species *M. longicollis* was published while preparing this manuscript ([@bib5], Fungal Planet description sheet 444).  The morphological description matches the morphology of isolate CBS 752.97, stored as *S. gracilis* in the CBS collection.  Sequence comparison of CBS 752.97 with the ex-type isolate of *M. longicollis* gave 99 % identity matches between all three sequenced loci.  We therefore named CBS 752.97 *M. longicollis*.  Multiple isolates stored as *M. trigonosporus* are now identified as *M. alveolaris* ([Table 1](#tbl1){ref-type="table"}).  Also many stored as *M. manginii* now cluster in different species clades than the type isolate of *M. manginii* which falls within the *S. candida* clade.  Four of them actually cluster within *Scopulariopsis*, corresponding to two new species (*S. sexualis* and *S. africana*).  These are more examples of the problems with identification based on morphology in these genera. * Microascus campaniformis* CBS 138126 has been omitted from the study awaiting a new culture deposit.  The isolate deposited to the CBS collection turned out to be a *M. melanosporus* isolate.  The published sequences from the ex-type isolate were also not included in this study, since the *tef1* sequence (GenBank [HG380418](ncbi-n:HG380418){#intref0100}) seems to belong to the *M. melanosporus* clade.  The deposited ITS (GenBank LM652391), and *tub2* (GenBank LM652606) sequences are unique sequences suggesting it is indeed a new species in *Microascus*.  The ex-type isolate need to be recovered and re-sequenced to place it into the phylogenetic tree. * Scopulariopsis candida* has a lot of molecular variation and is non-monophyletic in the species phylogeny ([Fig.  1](#fig1){ref-type="fig"}).  This is congruent with previous publications ([@bib14], [@bib29]).  No solution for this problem could be provided in this study, since the single gene trees were not congruent.  Further study including more isolates will be necessary to solve this (for now) non-monophyletic species. 

Conclusions {#sec6}
===========

In the genus *Microascus* 33 phylogenetic species can be distinguished based on (parts) of the ITS, *tub2* and *tef1* gene regions.  From these 33 species, seven are described here as new species, and one new name and new combination are proposed.  Thirteen *Microascus* species are found in the indoor environment of which *M. melanosporus* is most commonly found, followed by *M. paisii*.  In the genus *Scopulariopsis* 12 phylogenetic species can be distinguished based on (parts) of the ITS, *tub2* and *tef1* gene regions.  From these 12 species, four are described here as new species.  Three *Scopulariopsis* species are found in the indoor environment of which *S. brevicaulis* and *S. candida* are most common.  No correlation was found between phylogenetic relationships and habitat preference in the genera *Microascus* and *Scopulariopsis*.  The genus *Fuscoannellis* is placed in synonymy with *Yunnania*.  The genus *Yunnania*, which is not known for indoor environments, now includes three species of which one is described here as new and one has a new name combination proposed. 

Key to the most common *Microascus*, *Scopulariopsis* and *Cephalotrichum* species from the indoor environment {#sec6.1}
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1a. Synnemata absent ..................................................................... 21b. Synnemata present .......................................... ***Cephalotrichum***, 82a. Annellides lageniform or ampulliform .......................... ***Microascus***, 32b. Annellides cylindrical .......................................\... ***Scopulariopsis***, 73a. Colony diam.  on OA after 14 d at 25 °C \> 20 mm .............................................. ***M. melanosporus***3b. Colony diam.  on OA after 14 d at 25 °C \< 20 mm ..................................................................... 44a. Conidia obovoid to broad clavate or fusiform .................................. 54b. Conidia broadly ellipsoidal to short clavate .................................... 65a. Annellides 9--12 μm long .......................................... ***M. fusisporus***5b. Annellides 16--22 μm long ....................................... ***M. trautmannii***6a. Annellides 4--6 μm long ............\... ***M. hollandicus*** or ***M. pseudopaisii***6b. Annellides 6--11 μm long ........................ ***M. atrogriseus*** or ***M. paisii***7a. Roughened conidia, able to grow at 36 °C on OA ...................................................... ***S. brevicaulis***7b. Smooth conidia, no growth at 36 °C on OA ........................................................... ***S. candida***8a. Coiled setae present on the upper part of the synnemata ......................................................\... ***C. gorgonifer***8b. Setae absent ............................................................................ 99a. Conidia distinctively rough .................................\... ***C. verrucisporum***9b. Conidia smooth .....................................................................\... 1010a. Conidia 3.5--5 × 2--3 μm ....................................\... ***C. microsporum***10b. Conidia larger ...\... ***C. pseudopurpureofuscum*** or ***C. purpureofuscum***

This research was supported by a grant from the Alfred P. Sloan Foundation Program on the Microbiology of the Built Environment (Grant No.  G-2014-14529).  The authors would like to thank Dr. Christoph Trautmann (Umweltmykologie, Germany), Prof. Dr. Keith Seifert (Agriculture and Agri-Food, Canada), Dr. Kinga Plewa (Poland) and the Technical University of Denmark (DTU) for sending/depositing their indoor isolates into the working collection of the Westerdijk Fungal Biodiversity Institute Applied and Industrial Mycology Group.  Dr. Konstanze Bensch (Westerdijk Institute, the Netherlands) is thanked for her help with naming the new species and Dr. Marcelo Sandoval-Denis (Westerdijk Institute) for helpful discussions on the manuscript. 

Peer review under responsibility of Westerdijk Fungal Biodiversity Institute. 

[^1]: The ^T^ indicates the ex-type isolate of the synonymised species. 

[^2]: ATCC: American Type Culture Collection, Manassas, VA, USA; BMU: Beijing Medical University (Peking University), Beijing China; CBS: Culture Collection of the Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CGMCC: China General Microbiological Culture Collection Center, Institute of Microbiology, Chinese Academy of Sciences, Beijing, China; DTO: Working Collection of the Applied and Industrial Mycology Group of the Westerdijk Fungal Biodiversity Institute, Utrecht, Netherlands; FMR: Facultat de Medicina i Ciències de la Salut, Reus, Spain; FRR: Division of Food Research, Food Research Laboratory, CSIRO, North Ryde, Australia; HACC: Research Laboratory, Hindustan Antibiotics Ltd., Pimpri, Pune, India; IFO: Institute for Fermentation Culture Collection, Osaka, Japan; IMI: Culture Collection of CABI Europe-UK, Egham, UK; LCP: Laboratory of Cryptogamy, National Museum of Natural History, Paris, France; MUCL: (Agro)Industrial Fungi and Yeast Collection of the Belgian Co-ordinated Collections of Micro-organisms (BCCM), Louvain-la Neuve, Belgium; NHL: National Institute of Hygienic Sciences, Tokyo, Japan; NRRL: ARS Culture Collection, U.S. Department of Agriculture, Peoria, IL, USA; RGR: Personal Collection of Rodney G. Roberts; UAMH: University of Toronto, UAMH Centre for Global Microfungal Biodiversity, Toronto, Canada; UTHSC: Fungus Testing Laboratory, Department of Pathology, University of Texas Health Science Center, San Antonio, TX, USA; VKM: All-Russian Collection of Microorganisms, Moscow, Russia.  Ex-epitype, -isotype, -type, and -neotype isolates are indicated with ^ET^, ^IT^, ^T^ and ^NT^, respectively, and printed in bold.