Newly recognised lineages of perithecial ascomycetes: The new orders Conioscyphales and Pleurotheciales

Abstract

Phylogenetic analyses of DNA sequences from nuclear ribosomal and protein-coding loci support the placement of several perithecial ascomycetes and dematiaceous hyphomycetes from freshwater and terrestrial environments in two monophyletic clades closely related to the Savoryellales. One clade formed by five species of Conioscypha, and a second clade containing several genera of uncertain taxonomic status centred on Pleurothecium, represent two distinct taxonomic groups at the ordinal systematic rank. They are proposed as new orders, the Conioscyphales and Pleurotheciales. Several taxonomic novelties are introduced in the Pleurotheciales, i.e. two new genera (Adelosphaeria and Melanotrigonum), three novel species (A. catenata, M. ovale, Phaeoisaria fasciculata) and a new combination (Pleurotheciella uniseptata). A new combination is proposed for Savoryella limnetica in Ascotaiwania s. str. based on molecular data and culture characters. A strongly supported lineage containing a new genus Plagiascoma, species of Bactrodesmiastrum and Ascotaiwania persoonii, was identified as a sister to the Conioscyphales/Pleurotheciales/ Savoryellales clade in our multilocus phylogeny. Together, they are nested in a monophyly in the Hypocreomycetidae, significantly supported by Bayesian inference and Maximum Likelihood analyses. Members of this clade share a few morphological characters, such as the absence of stromatic tissue or clypeus, similar anatomies of the two-layered ascomatal walls, thin-walled unitunicate asci with a distinct, non-amyloid apical annulus, symmetrical, transversely septate ascospores and holoblastic conidiogenesis. They represent the only fungi in the Hypocreomycetidae with apically free, filiform to cylindrical, persistent or partially disintegrating paraphyses. The systematic placement of two other dematiaceous hyphomycetes was resolved based on DNA sequences; Phragmocephala stemphylioides is a member of the Pleurotheciales and Triadelphia uniseptata is within the Savoryellales.

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Persoonia37,2016:57– 81
www.ingentaconnect.com/content/nhn/pimj http://dx.doi.org/10.3767/003158516X689819
RESEARCH ARTICLE
INTRODUCTION
The subclass Hypocreomycetidae(Sordariomycetes)includes
non-lichenised ascomycetes with perithecial and cleistothecial
ascomata.Manyspeciesareparasiticonplants,insectsand
otherfungi.Someareendophytesinplantsor saprobes on
decaying wood and herbs, and some are involved in obligate
mutualismwithwood-boringbeetles.BasedonDNAsequences
from nuclear ribosomal and protein-coding loci, the Hypocreo-
mycetidae was recognised as a strongly supported monophy-
leticcladeencompassing five orders(Spataforaetal. 2007,
Zhangetal.2007),i.e.theCoronophorales, Halosphaeriales,
Hypocreales, Melanosporales, Microascales, and one family
not then placed in an order, the Glomerellaceae.Theabsence
ofparaphyseswasusedtodelimitthissubclass(Zhangetal.
2007).Inthemorerecentclassification,theHypocreomycetidae
compriseseightorders,i.e.theCoronophorales, Falcocladiales,
Glomerellales including the Plectosphaerellaceae(Zareetal.
2007,Réblová et al. 2011),Hypocreales, Melanosporales,
a revised Microascales(DeBeeret al. 2013), Savoryellales
(Boonyuenetal.2011)and Torpedosporales(Schochetal.
2007,Jonesetal.2014,2015).Hamathecialelementsinthe
Hypocreomycetidaecompriseseveraltypes,i.e.apical,centri-
petal and lateral paraphyses, catenophyses, a reticulate net-
workoffiliformfilamentsattachedatthetopandbottomofthe
ascomatalcavity;sometimesinterthecialfilamentsarelacking.
Theonlygroupcharacterisedbyparaphyses,i.e.sterilefiliform,
apicallyfreefilamentsemergingfromthehymeniumamong
asci and growing upwards, is the Savoryellales, placed in this
subclass based on a combined analysis of six nuclear loci
(Boonyuenetal.2011).
The Savoryellales comprises three genera, Ascotaiwania, Canali-
sporium5 and Savoryella from freshwater, brackish, marine and
terrestrialhabitats.They share a setofcharactersincluding
non-stromatic,immersed,semi-immersedtosuperficial,dark,
coriaceous ascomata, often lying horizontally to the host, unitu-
nicate asci with a non-amyloid apical annulus, partly disintegra-
ting paraphyses and fusiform to ellipsoidal, transversely septate
ascosporeswithhyalinepolarcellsandbrownmiddlecells.
Asexual morphs were experimentally proven for two species
of Ascotaiwania (asMonotosporella,Ranghoo&Hyde1998,
Sivichaiet al.1998)andonespeciesofCanalisporium(with
Ascothailandiasexualmorph;Sri-indrasutdhietal.2010).The
distant placement of Helicoön farinosum, the asexual morph
of Ascotaiwania hughesii(Fallahetal.1999),frommembersof
the SavoryellaleswasrevealedbyrDNAdata (Boonyuenetal.
2011,Réblováetal.2012).Theasexualmorphslinkedtothe
Savoryellales are dematiaceous hyphomycetes characterised
by semi-macronematous conidiophores and monoblastic coni-
Newly recognised lineages of perithecial ascomycetes:
the new orders Conioscyphales and Pleurotheciales
M.Réblová1,K.A.Seifert2,J.Fournier3,V.Štěpánek4
1 DepartmentofTaxonomy,InstituteofBotanyoftheAcademyofSciences,
Průhonice,CzechRepublic;
 correspondingauthore-mail:martina.reblova@ibot.cas.cz.
2 Biodiversity(Mycology and Botany),Agricultureand Agri-FoodCanada,
Ottawa,Ontario,Canada.
3 LasMuros,Rimont09420,France.
4 LaboratoryofEnzymeTechnology,InstituteofMicrobiologyoftheAcademy
ofSciences,Prague,CzechRepublic.
Key words
freshwater fungi
holoblastic conidiogenesis
Hypocreomycetidae
multigene analysis
Phaeoisaria
systematics
AbstractPhylogeneticanalysesofDNAsequencesfromnuclearribosomalandprotein-codinglocisupportthe
placement of several perithecial ascomycetes and dematiaceous hyphomycetes from freshwater and terrestrial
environments in two monophyletic clades closely related to the Savoryellales.Onecladeformedbyfivespeciesof
Conioscypha and a second clade containing several genera of uncertain taxonomic status centred on Pleurothe-
cium,representtwodistincttaxonomicgroupsattheordinalsystematicrank.Theyareproposedasnew orders,
the Conioscyphales and Pleurotheciales.Severaltaxonomicnovelties areintroducedinthePleurotheciales, i.e.
twonew genera(Adelosphaeria and Melanotrigonum),three novelspecies (A. catenata, M. ovale, Phaeoisaria
fasciculata)and anewcombination (Pleurotheciella uniseptata).Anew combinationisproposed forSavoryella
limnetica in Ascotaiwania s.str.based on molecular data andculturecharacters.Astronglysupported lineage
containing a new genus Plagiascoma, species of Bactrodesmiastrum and Ascotaiwania persoonii, wasidentified
as a sister to the Conioscyphales/Pleurotheciales /Savoryellalescladeinourmultilocusphylogeny.Together,they
are nested in a monophyly in the Hypocreomycetidae,significantlysupportedbyBayesianinferenceandMaximum
Likelihoodanalyses.Membersofthiscladeshareafewmorphologicalcharacters,suchastheabsenceofstromatic
tissue or clypeus, similar anatomies of the 2-layered ascomatal walls, thin-walled unitunicate asci with a distinct,
non-amyloidapicalannulus,symmetrical,transverselyseptateascosporesandholoblasticconidiogenesis.They
represent the only fungi in the Hypocreomycetidaewithapicallyfree,filiform tocylindrical,persistentorpartially
disintegrating paraphyses. The systematic placement of two other dematiaceous hyphomycetes was resolved
basedonDNAsequences;Phragmocephala stemphylioides is a member of the Pleurotheciales and Triadelphia
uniseptata is within the Savoryellales.
Article infoReceived:15July2015;Accepted:9October2015;Published:20October2015.
5 InkeepingwiththetenetsofthenewInternationalCodeontheNomencla-
ture of algae, fungi and plants, we hereafter routinely use the oldest generic
name for holomorphs; in some cases, this was originally the name of an
asexualmorph.

58 Persoonia–Volume37,2016
diogenous cells producing brown, thick-walled, transversely
septate or cheiroid, dictyoseptate macroconidia, rare characters
in the Hypocreomycetidae.Althoughtheasexualmorphsof
Savoryellaareunknown(Boonyuenetal.2011),darkbrown,
3–5-septate macroconidiawere obtained in living cultures
derived from ascospore isolates of two of our specimens of
S. limnetica(Changetal.1998)collectedonwoodsubmerged
infreshwaterinFrance.Identicalconidiawerealsoobserved
scatteredamongascomataonthehost.
Previousphylogeniesinferredfromsequencesofthesmalland
largesubunitofnuclearribosomalDNA(nuc18Sandnuc28S
rDNA)and thesecondlargestsubunitof RNApolymeraseII
(rpb2)revealedacloserelationship among members of the
Savoryellales and several terrestrial and freshwater genera
ofuncertaintaxonomicstatusformingtwoclades,i.e.Conio-
scypha and a clade comprising Phaeoisaria, Pleurotheciella,
Pleurothecium and Sterigmatobotrys (Réblová et al. 2012).
However,relationshipsamongthesegeneraremainedlargely
unresolved.Theyarecharacterisedbynon-stromatic,semi-im-
mersedtosuperficial,brown,subhyalinetopaleorangeperithe-
cial ascomata, paraphyses, unitunicate asci with a non-amyloid
apical annulus and ellipsoidal to fusiform, hyaline to subhyaline,
septateascospores(Fernándezetal.1999,Réblová&Seifert
2004,2011,Réblová etal.2012).Theirasexualmorphs are
hyphomycetes with dematiaceous or hyaline conidiophores,
holoblastic, sympodial conidiogenous cells and conidia that are
oftenformedonashortrachisondenticles.Theconidiogenesis
of Conioscyphaisunique;brown,non-septateconidiaareborn
in cyathiform to doliiform blastic conidiogenous cells surrounded
by hyaline, cup-like collarettes with a multilamellar structure
(Shearer&Motta1973).
PreliminaryanalysisofDNAsequencesofnuclearribosomaland
protein-coding loci of four undescribed ascomycetes revealed
their close relationship with members of the Savoryellales and
the clade mentioned above centred around Pleurothecium.
Threeoftheseunidentifiedfungiareperithecialascomycetes
that share with members of the Pleurothecium clade characters
ofascomata,asci,paraphysesandascospores.Fivespecimens
ofthefirstundescribedfunguswerefoundonstronglydecaying
wood of Quercus cerrisintheCzechRepublic.Althoughnoconi-
diophores were formed on the host, cultures derived from asco-
spore isolates yielded identical asexual morphs with oval to
bean-shaped, 1-septate, brown conidia formed holoblastically
ona shortdenticleonalmosttriangularconidiogenouscells.
Thesecondunidentifiedascomycetewascollectedondecay-
ing wood of Fagus sylvaticain theCzechRepublic.Asingle
collection of the third undescribed ascomycete was made on
decaying wood of Fraxinus excelsior submerged in freshwater
insouthernFrance.Culturesofbothfungiwerederivedfrom
isolatedascospores.Noconidiophoreswereobservedonthe
host and none were formed in vitro; only brown, ellipsoidal to
globose cells were formed blastically directly on vegetative
hyphaeinaxenicculture.Basedonthesimpleandnondescript
sexual morphological characteristics, we could not conclusively
attributeanyofthesethreefungitoaknownascomycetegenus.
Two morphologically similar specimens of a dematiaceous hy-
phomycetepreliminary identified as Phaeoisaria sp. were
made on decaying deciduous wood in Canada and the Czech
Republic.Theyrepresentafungusmorphologicallysimilarto
Ph. clematidis, the type species of the genus, in producing non-
septate, obovoid conidia holoblastically on short denticles on
sympodially proliferating conidiogenous cells, but differ in the
absence of well-developed synnemata on the host and in vitro
(VonHöhnel1909,Deighton1974).Inbothstrains,theconidio-
phoreswerearrangedinfasciclesandlackedadistinctstipe.
The aim of this study is to investigate phylogenetic relationships
ofthethreeunidentifiedperithecialascomycetes,Phaeoi saria
sp.,andalsoDactylaria uniseptata and S. limnetica, with mem-
bers of the Savoryellales and Pleurotheciumclade.Theaffinities
of two dematiaceous hyphomycetes Phragmocephala stem-
phylioides and Triadelphia uniseptata, coincidentally discovered
toberelatedtothisclade,arealsodocumented. We also inves-
tigate the relationships of taxa characterised by the presence of
paraphyses in the subclass Hypocreomycetidae.Althoughnot
a part of the presentation of new taxa, our re-examination of
this subclass allows further consideration of the Conioscypha
clade,presentlyconsideredincertaesedis(Réblová&Seifert
2004,Zelskietal.2014).Inordertofurtherclarifythesystematic
positions of the Conioscypha and Pleurothecium clades, we
utilisedDNAsequencecharactersfromthenucrDNAinternal
transcribedspacerbarcode(ITS1-5.8S-ITS2),threeprotein-
codingandtworibosomalnuclearloci.
MATERIALS AND METHODS
Herbarium material and fungal strains
Dryascomatawere rehydrated withwater;materialwas ex-
aminedwithanOlympus SZX12 dissectingmicroscopeand
hand-sectionedcentrummaterial(includingasci,ascospores
andparaphyses)wasmountedinMelzer’sreagent,Lugol,90%
lacticacid,aqueous cotton-blue(1mg/mL),Pelikanink and
blueorblackWatermanink.Handsectionsoftheascomatal
wallwerestudiedin3%KOHorheatedchloral-lactophenol.
All measurements were made in Melzer’s reagent. Means
±standarddeviation(SD)basedon20–25measurementsare
given for dimensions of asci and ascospores. Images were
capturedbydifferentialinterference (DIC) orphasecontrast
(PC)microscopyusinganOlympusDP70cameraoperatedby
ImagingSoftwareCellonanOlympusBX51compoundmicro-
scope.Conidiaandconidiogenous cells werephotographed
inthe living state using an FEI Quanta 200 Environmental
ScanningElectronMicroscope(ESEM).Ac. 2 × 2 mm cube of
agar with mycelium was observed at 20 kV after the sample
chamberachievedlocalthermodynamicequilibrium:chamber
pressure200Pa,sampletemperaturefrom-15°Cto-16°C.
AGaseous Secondary Electron Detector (GSED) was used
forsignaldetection.Coolingofthespecimeninthechamber
was achieved using a PC controlled Peltier cooling stage with
externalwaterchiller(madebyJTManufacturing,USA).
Multi-ascosporeandmulti-conidialisolateswereobtainedfrom
freshmaterialwiththeaidofasporeisolator(Meopta,Prague,
CzechRepublic).Ascosporesandasciwerespreadonwater
agar,ascosporesandconidiagerminatedwithin48h.Germina-
ting ascospores were transferred and isolates were grown on
wateragar,CMA(Difco), potatodextroseagar(PDA,Oxoid)
andpotato-carrotagar(PCA,Gamsetal.1998).Colonieswere
examinedafter7,21and30dincubatedat25°Cinthedark.
Ex-type and other cultures are maintained at the CBS-KNAW
FungalBiodiversityCentre,Utrecht, The Netherlands(CBS)
and Canadian Collection of Fungal Cultures, Agriculture and
Agri-FoodCanada,Ottawa,Canada(DAOMC).Typeandother
herbariummaterialisdepositedintheMycologicalHerbarium
intheNationalMuseuminPrague,CzechRepublic(PRM)and
CanadianNationalMycologicalHerbarium,Ottawa,Canada
(DAOM).TheOnlineAuctionColourChart (2004) wasused
asthecolourstandard.
DNA extraction, amplification and sequence alignment
CulturesusedforDNAisolationsweregrownaspreviouslyde-
scribedbyRéblováetal.(2011)andDNAwasextractedfollow-
ingtheprotocolsofLee&Taylor(1990).Proceduresforam-
plifyingandsequencingtheinternaltranscribedspacerrDNA
(ITSrDNA),small and largesubunitnuclearribosomal DNA
(nuc18S rDNA, nuc28S rDNA), second largest subunit of

59
M.Réblováetal.:ThenewordersConioscyphales and Pleurotheciales
RNApolymeraseII(rpb2)andDNAreplicationlicensingfactor
(mcm7) wereperformedasdescribedinRéblováetal.(2011,
2013).Afragment ofthe5’-end oftheβ-tubulingene region
(exons 3 to 6) was amplified and sequenced using primers
Bt2a/ benA1andBt2b(Glass&Donaldson1995,Geiseretal.
1998).SequenceswereeditedusingSequencherv.5.0(Gene
CodesCorp.,AnnArbor,MI,USA).
GenBankaccessionnumbersfornewlysequenced taxaand
otherhomologoussequencesofmembersoftheSavoryellales
and two new orders described in this study retrieved from Gen-
BankarelistedinTable1.Fordetailedinvestigationofphylo-
genetic relationships within the Sordariomycetes,sequences
ofthethreelocinuc28S,nuc18Sandrpb2includedinRéblová
etal.(2015)weredownloadedfromGenBankandcombined
withthosegeneratedduringthepresentstudy.
Taxon Source ex-type GenBank accession numbers
  ITS nuc28S nuc18S RPB2 MCM7 TUB2
Adelosphaeria catenata CBS138679 T KT278721 KT278707 KT278692 KT278743 KT278733 KT278754
Ascotaiwania lignicola NIL00005  – HQ446364 HQ446284 HQ446419 – –
Ascotaiwania limnetica CBS126576  – – KT278689 – KT278731 –
CBS126792  – – KT278690 – KT278732 –
Ascotaiwania mitriformis HKUCC3706  – AF132324 – – – –
Ascotaiwania sawadae SS00051  – HQ446363 HQ446283 HQ446418 – –
Ascotaiwania persoonii A57-14C T – AY094190 – – – –
A57-14C  – AY590295 – – – –
Bactrodesmiastrum obovatum FMR6482  – FR870266 – – – –
Bactrodesmiastrum pyriforme FMR10747  – FR870265 – – – –
FMR11931  – HE646637 – – – –
Brachysporiella setosa HKUCC3713  – AF132334 – – – –
Canalisporium caribense SS03683  – GQ390269 GQ390254 – – –
Canalisporium elegans SS00895  – GQ390271 GQ390256 HQ446425 – –
Canalisporium exiguum SS00809  – GQ390281 GQ390266 HQ446436 – –
Canalisporium grenadoideum BCC20507 T – GQ390267 GQ390252 HQ446420 – –
Canalisporium pulchrum SS03982  – GQ390277 GQ390262 HQ446431 – –
Conioscypha japonica CBS387.84 T – AY484514 JQ437438 JQ437438 – –
Conioscypha lignicola CBS335.93 T – AY484513 JQ437439 JQ429260 – –
Conioscypha minutispora CBS137253 T – KF924559 – – – –
Conioscypha peruviana ILL41202 T – KF781539 – – – –
Conioscypha varia CBS113653  – AY484512 AY484511 JQ429261 – –
Flammispora bioteca BCC13367 T – – AY722100 – – –
Helicoön farinosum DAOM241947  JQ429145 JQ429230 – – – –
ILLS53605  – AY094189 – – – –
ILLS53605  – AY316357 – – – –
Magnisphaera stevemossago CBS139776  – KT278704 KT278691 KT278740 – –
Melanotrigonum ovale CBS138742  KT278723 KT278708 KT278695 KT278744 – KT278756
CBS138743 T KT278724 KT278709 KT278696 KT278745 – KT278757
CBS138744  KT278725 KT278710 KT278697 KT278746 – –
CBS138815  KT278722 KT278711 KT278698 KT278747 – KT278755
M.R.3685  KT278726 KT278712 – KT278748 – KT278758
Phaeoisaria clematidis CBS113340  EU552148 – – – – –
DAOM226789  JQ429155 JQ429231 JQ429243 JQ429262 – –
Phaeoisaria fasciculata CBS127885 T KT278719 KT278705 KT278693 KT278741 – KT278752
DAOM230055  KT278720 KT278706 KT278694 KT278742 – KT278753
Phaeoisaria sedimenticola CGMCC3.14949 T JQ074237 JQ031561 – – – –
Phaeoisaria sparsa FMR11939  – HF677185 – – – –
Phaeoisaria sp. unknown  – nuc28S* – – – –
Phragmocephala stemphylioides DAOM673211  KT278730 KT278717 – – – –
Pisorisporium cymbiforme CBS127887  – – KT278699 KT278750 – –
CBS127888  – – KT278700 KT278751 – –
Plagiascoma frondosum CBS139031 T – KT278713 KT278701 KT278749 KT278734 –
Pleurotheciella centenaria DAOM229631 T JQ429151 JQ429234 JQ429246 JQ429265 – –
Pleurotheciella rivularia CBS125238 T JQ429160 JQ429232 JQ429244 JQ429263 KT278735 KT278759
CBS125237  JQ429161 JQ429233 JQ429245 JQ429264 KT278736 KT278760
Pleurotheciella uniseptata DAOM673210 T KT278729 KT278716 – – – –
Pleurothecium obovoideum CBS209.95 T EU041784 EU041841 – – – –
Pleurothecium recurvatum CBS101581  JQ429148 AF261070 JQ429248 JQ429266 – –
CBS138747  KT278728 KT278714 KT278703 – – –
CBS138686  KT278727 KT278715 KT278702 – KT278737 –
CBS131646  JQ429150 JQ429236 JQ429250 – – –
CBS131272  JQ429149 JQ429237 JQ429251 JQ429268 – –
Pleurothecium semifecundum CBS131271 T JQ429159 JQ429240 JQ429254 JQ429270 – –
CBS131482  JQ429158 JQ429239 JQ429253 – – –
Savoryella appendiculata NF00206  – – HQ446293 HQ446442 – –
Savoryella aquatica SS03801  – HQ446372 HQ446290 HQ446441 – –
Savoryella lignicola NF00204  – HQ446378 HQ446299 – – –
Savoryella longispora SAT00322  – HQ446380 HQ446302 HQ446450 – –
Savoryella paucispora SAT00866  – HQ446381 HQ446303 HQ446451 – –
Savoryella verrucosa SS00052  – HQ446374 HQ446298 HQ446445 – –
Sterigmatobotrys macrocarpa PRM915682  JQ429153 GU017317 JQ429255 – KT278739 KT278762
DAOM230059  JQ429154 GU017316 – JQ429271 KT278738 KT278761
Sterigmatobotrys uniseptata FMR11937  HF677178 – – – – –
Taeniolella rudis DAOM229838  JQ429152 JQ429241 JQ429256 JQ429272 – –
Triadelphia uniseptata DAOMC250376  – KT278718 – – – –
Table 1 A list of members of the Conioscyphales, Pleurotheciales, Savoryellalesandotherfungi,theirisolateinformationandnewsequencesdetermined
forthisstudyandthoseretrievedfromGenBank.SequenceswithGenBankaccessionnumbersinbold weregeneratedforthisstudy.Sequencenuc28S*
publishedinChewetal.(2010).

60 Persoonia–Volume37,2016
Sequences were manually aligned in BioEdit v.7.1.8(Hall
1999).Nuclearribosomal loci werealignedaccordingto the
secondary structure of Saccharomyces cerevisiae to improve
the decisions on homologous characters and introduction of
gaps(Gutell1993,Gutellet al. 1993, www.rna.ccbb.utexas.
edu).Theseproceduresand alignment of the sequences of
protein-codinggeneswereperformedasdescribedinRéblová
&Réblová(2013).
The single-locus datasets were examined for topological incon-
gruenceamongloci(ITS:26sequencesand616characters;
β-tubulin: 11sequences and 500 characters; nuc28S: 126
sequencesand 1947 characters; nuc18S: 104 sequences
and1792characters;rpb2segments5–7:77sequencesand
1 216 characters; mcm7:eightsequencesand659characters).
TheITSandβ-tubulinlociweregeneratedonlyformembers
of the new order Pleurotheciales.Becauseonlyafewmcm7
sequencesweregenerated,theywerenottestedfortopological
conflicts among clades at familial or ordinal rank in the Sordario-
mycetes.Foreachindividualpartition,500bootstrapreplicates
weregeneratedwithRAxML-HPCv.7.0.3 (Stamatakisetal.
2005,Stamatakis2006)andcomparedvisuallyfortopological
conflictamongsupportedcladesinphylogenetictrees.Acon-
flict between two loci was assumed to occur when a clade ap-
pearedmonophyleticwithbootstrapsupportof≥75%inone
tree,butwasnotsupportedasmonophyleticinanother(Mason-
Gamer & Kellogg 1996). Individual, conflict-free alignments
wereconcatenatedtocombinesequencesfortwosubsequent
phylogeneticanalyses.Themultiplesequencealignmentsare
depositedinTreeBASE(Studyno.18187).
Phylogenetic analyses
Phylogeneticrelationshipsoftheunidentifiedfungiandother
ascomyceteswereresolvedbytwocombinedanalysesofITS,
nuc18S,nuc28S,β-tubulin,mcm7 and rpb2sequencesofrep-
resentatives of the Sordariomycetes.Weanalysedthewhole
ITSrDNAbarcode,thefirst2 /3of the5’halfofthenuc28S,
theentirenuc18S,partialmcm7, exons3– 6ofβ-tubulin and
segments5 –7ofrpb2.Bases1–155ofthenuc18S,1–85of
the nuc28S and 1– 58 of the rpb2 alignments at the 5’-end
and1470 –1947ofthe nuc28Salignmentatthe3’-endwere
excluded from analysis because of incompleteness of the ma-
jorityoftheavailablesequences.Thecodingregions(exons)
3,4,5andpartly6oftheβ-tubulinwithatotallengthof291
nucleotideswereanalysed,non-codingregionswereexcluded.
The combined datasets were partitioned into several subsets
ofnucleotidesites,i.e.ITS,nuc28S,nuc18S,andfirst,second
andthirdcodon positionsofβ-tubulin, mcm7 and rpb2.Two
members of the Leotiomycetes, Leotia lubrica and Microglos-
sum rufum,wereusedtorootthetwomultilocusphylogenies.
TheprogramMrModeltest2v.2.3(Nylander2008)wasusedto
infertheappropriatesubstitutionmodelthatwouldbestfitthe
modelofDNAevolutionforeachsequencedatasetandeach
partitionofthecombineddatasets.Maximumlikelihood(ML)
andBayesianinference(BI)analyseswereusedtoestimate
phylogenetic relationships. ML analysis was performed with
RAxML-HPCv.7.0.3withaGTRCATmodelofevolution.Nodal
supportwasdeterminedbynon-parametricbootstrapping(BS)
with1000replicates.
Bayesian inference analysis was performed in a likelihood
frameworkasimplementedinMrBayesv.3.0b4toreconstruct
phylogenetictrees(Huelsenbeck&Ronquist2001).Forthe
ITS,nuc18S,nuc28S,andrpb2 dataset, we used for each par-
titiontheGTR+G+Isubstitutionmodel.Forβ-tubulinweused
HKY+G,F81andSYM+Gforthefirst,secondandthirdcodon
position, and for mcm7 weused HKY+G, GTR+GandGTR
forthefirst, second andthirdcodonposition.TwoBayesian
searcheswereperformedusingthedefaultparameters.Analy-
ses were run for 10 million generations, with trees sampled
every1000generations.Tracerv.1.6.0.(Rambautetal.2013)
wasusedtoconfirm convergence of treesandburn-in.The
first50000trees,whichrepresentedtheburn-inphaseofthe
analysis,werediscarded.Theremainingtreeswereusedfor
calculatingposteriorprobabilities(PP)ofrecoveredbranches
(Larget&Simon1999).
PHYLOGENETIC RESULTS
Inthefirstanalysis,134combinednuc18S,nuc28Sandrpb2
sequenceswereassessedfor120speciesin20ordersinthe
Sordariomycetes.Thealignmenthad2767distinctalignment
patterns(MLanalysis).IntheMLtreeshowninFig.1,astrongly
supportedmonophyleticcladewasresolved(100MLBS/1.0
PP)intheHypocreomycetidaewiththreenestedclades.The
Savoryellales(100/1.0)wasassociatedwiththeConioscypha
clade(100/1.0)includingfivespeciesandPleurothecium clade
(98/1.0)comprisingeightgeneraandthreeotherundescribed
ascomycetes.Theyrepresenttwonewlineagesoffreshwater
and terrestrial fungi and are introduced below as the orders
Conioscyphales and Pleurotheciales. Astrongly supported
monophyleticlineage(100/1.0)containingAscotaiwania per-
soonii, two species of the dematiaceous hyphomycetous genus
Bactrodesmiastrumandoneunidentifiedascomyceteisposi-
tioned as a sister to a clade containing Conioscyphales, Pleuro-
theciales and Savoryellales.Togethertheyformarobustmono-
phylum(100/1.0)intheHypocreomycetidae, including Flammi-
spora bioteca(BCC13367)inabasalposition.
Two undescribed perithecial fungi were nested within the
Pleurotheciales and are described here as new genera, Mela-
notrigonum and Adelosphaeria.TwostrainsofPhaeoisaria sp.
with fasciculate conidiophores were positioned in the strongly
supported Phaeoisaria clade(100 /1.0)ofthePleurotheciales
as the sister taxon to Phaeoisaria sparsa.Theyareintroduced
asanewspecies.Thethirdunidentifiedperithecialascomycete
from freshwater habitat was nested within the Bactrodesmia-
strum clade on a separate branch and it is described as a new
monotypic genus Plagiascoma.
Two strains of Savoryella limnetica and Triadelphia uniseptata
were positioned in the Ascotaiwania clade (79 /0.91) in the
Savoryellales.ThegenusAscotaiwania is polyphyletic in our
phylogeny.Helicoön farinosum, the asexual state of A. hughesii
is grouped within the Pleurotheciales.Ascotaiwania lignicola,
the type species, and three other species are members of the
Savoryellales, while A. persoonii is nested in the Bactrodesmia-
strumclade.Twodematiaceoushyphomycetes,Phragmoce-
phala stemphylioides and Dactylaria uniseptata, were grouped
among members of the Pleurotheciales; the latter is transferred
to Pleurotheciella below.
Inthesecondphylogeneticanalysis(Fig.2),thecombinedITS,
nuc18S,nuc28S,β-tubulin,mcm7 and rpb2 dataset consisted of
60sequencesrepresenting18speciesofthePleurotheciales,
fiveoftheConioscyphales, 15 species of the Savoryellales and
four species of the Bactrodesmiastrum clade.Thealignment
had2370distinctalignmentpatterns(MLanalysis).Therobust
clade containing the three orders and the Bactrodesmiastrum
clade(100/1.0)hasidenticaltopologiesinthethree-andsix-
genephylogenies.Sixterminal clades were identifiedinthe
PleurothecialesandarelabelledasCladeI to VI on Fig. 2.
CladesI,VandVIarestronglysupportedmonophyleticlineages
representing genera Melanotrigonum (100/1.0), Phaeoisaria
(100/1.0)andPleurotheciums.str.(100/1.0).CladeII(72/0.97)
is morphologically heterogeneous containing Brachysporiella
setosa, Helicoön farinosum, Phragmocephala stemphylioides

61
M.Réblováetal.:ThenewordersConioscyphales and Pleurotheciales
Fig. 1Multilocusphylogeneticanalysisofthenuc18S-nuc28S-rpb2sequencesoftheSordariomycetesshowingmajorityoftherecognizedordinallineages.
PhylogramwasinferredfromtheMLanalysiswithRAxMLusingaGTRCATmodelofevolution.Onlyhighbranchsupportisshownatthenodes,maximum
likelihoodbootstrapsupport(MLBS)≥70%andBayesianposteriorprobability(PP)≥0.95.Symbol●indicatesnodeswith100%MLBSand1.0PP.Taxa
written in boldrepresenttaxonomicnovelties.
Savoryella appendiculata NF 00206
Phaeoisaria fasciculata DAOM 230055
Conioscypha japonica CBS 387.84
Conioscypha varia CBS 113653
Helicoön farinosum ILLS 53605
Pleurothecium recurvatum CBS 138747
Melanotrigonum ovale CBS 138742
Canalisporium pulchrum SS 03982
Bactrodesmiastrum pyriforme FMR 10747
Canalisporium elegans SS 00895
Sterigmatobotrys macrocarpa PRM 915682
Ascotaiwania sawadae SS 00051
Helicoön farinosum DAOM 241947
Pleurothecium semifecundum CBS 131271
Conioscypha lignicola CBS 335.93
Pleurotheciella centenaria DAOM 229631
Melanotrigonum ovale CBS 138815
Phaeoisaria fasciculata CBS 127885
Bactrodesmiastrum pyriforme FMR 11931
Taeniolella rudis DAOM 229838
Phaeoisaria sedimenticola CGMCC 3.14949
Ascotaiwania limnetica CBS 126576
Melanotrigonum ovale CBS 138744
Conioscypha peruviana ILL 41202
Melanotrigonum ovale CBS 138743
Savoryella verrucosa SS 00052
Conioscypha minutispora CBS 137253
Pleurothecium recurvatum CBS 138686
Sterigmatobotrys macrocarpa CBS 113468
Phaeoisaria clematidis CBS 113340
Ascotaiwania limnetica CBS 126792
Triadelphia uniseptata DAOMC 250376
Pleurotheciella rivularia CBS 125238
Pleurotheciella rivularia CBS 125237
Pleurotheciella uniseptata DAOM 673210
Savoryella paucispora SAT 00866
Brachysporiella setosa HKUCC 3713
Ascotaiwania persoonii A57-14C
Flammispora bioteca BCC 13367
Savoryella lignicola NF 00204
Savoryella longispora SAT 00322
Ascotaiwania mitriformis HKUCC 3706
Savoryella aquatica SS 03801
Ascotaiwania lignicola NIL 00005
Phaeoisaria sp.
Plagiascoma frondosum CBS 127885
Canalisporium exiguum SS 00809
Phaeoisaria sparsa FMR 11939
Bactrodesmiastrum obovatum FMR 6482
Phaeoisaria clematidis DAOM 226789
Ascotaiwania persoonii A57-14C
Pleurothecium obovoideum CBS 209.95
Adelosphaeria catenata CBS 138679
Sterigmatobotrys uniseptata FMR 11937
Pleurothecium recurvatum CBS 131272
Pleurothecium semifecundum CBS 131482
Canalisporium grenadoideum BCC 20507
Phragmocephala stemphylioides DAOM 673211
Canalisporium caribense SS 03683
Pleurothecium recurvatum CBS 101581
Helicoön farinosum ILLS 53605
Melanotrigonum ovale M.R. 3685
79/0.91
87/1.0
84/1.0
90/1.0
78/1.0
95/0.99
91/0.99
72/1.0
88/1.0
97/1.0
98/1.0
98/1.0
96/1.0
97/1.0
97/1.0
96/1.0
98/1.0
84/1.0
94/1.0
99/1.0
84/1.0
97/1.0
73/1.0 80/1.0
99/1.0
90/1.0
0.1
Xylariales
Leo$omycetes(outgroup)
Koralionastetales
Lulworthiales
Pisorisporiales
Sordariales
Chaetosphaeriales
Boliniales
100/1.0
Magnaporthales
100/1.0
Diaporthales
Calosphaeriales
Falcocladiales
Coronophorales
Melanosporales
Torpedosporales
Hypocreales
Glomerellales
Microascales
HYPOCREOMYCETIDAE
SORDARIOMYCETIDAE
XYLARIOMYCETIDAE
Pleurotheciales Savoryellales Conioscyphales
incertae
sedis

62 Persoonia–Volume37,2016
Fig. 2MultilocusphylogeneticanalysisoftheITS-nuc18S-nuc28S-β-tubulin-rpb2-mcm7sequencesoftheCPS(Conioscyphales, Pleurotheciales, Savoryel-
lales)andBactrodesmiastrumclades.PhylograminferredfromtheMLanalysiswithRAxMLusingaGTRCATmodelofevolution.Onlyhighbranchsupportis
shownatthenodes,MLBS≥70%,PP≥0.95.Taxawritteninboldrepresenttaxonomicnovelties.
0.2
Ascotaiwania persoonii A57-14C
Savoryella longispora SAT 00322
Conioscypha peruviana ILL 41202 T
Ascotaiwania persoonii A57-14C
Pleurothecium recurvatum CBS 101581
Ascotaiwania lignicola NIL 00005
Conioscypha minutispora CBS 137253 T
Pleurothecium recurvatum CBS 138686
Savoryella verrucosa SS 00052
Ascotaiwania mitriformis HKUCC 3706
Pleurothecium semifecundum CBS 131271 T
Ascotaiwania limnetica CBS 126792
Conioscypha varia CBS 113653
Canalisporium elegans SS 00895
Conioscypha japonica CBS 387.84 T
Sterigmatobotrys macrocarpa PRM 915682
Phaeoisaria clematidis CBS 113340
Pleurothecium recurvatum CBS 131272
Melanotrigonum ovale CBS 138743 T
Phaeoisaria fasciculata DAOM 230055
Pleurothecium semifecundum CBS 131482
Phaeoisaria sparsa FMR 11939
Ascotaiwania limnetica CBS 126576
Sterigmatobotrys macrocarpa CBS 113468
Savoryella aquatica SS 03801
Phaeoisaria sedimenticola CGMCC 3.14949 T
Pleurothecium recurvatum CBS 138747
Phaeoisaria fasciculata CBS 127885 T
Pleurotheciella rivularia CBS 125238 T
Savoryella lignicola NF 00204
Flammispora bioteca BCC 13367 T
Triadelphia uniseptata DAOMC 250376
Pleurotheciella centenaria DAOM 229631 T
Taeniolella rudis DAOM 229838
Phaeoisaria sp.
Adelosphaeria catenata CBS 138679 T
Phaeoisaria clematidis DAOM 226789
Bactrodesmiastrum pyriforme FMR 10747
Ascotaiwania sawadae SS 00051
Bactrodesmiastrum pyriforme FMR 11931
Pleurothecium obovoideum CBS 209.95 T
Pleurotheciella rivularia CBS 125237
Helicoön farinosum DAOM 241947
Leotia lubrica AFTOL-1
Canalisporium pulchrum SS 03982
Canalisporium caribense SS 03683
Melanotrigonum ovale CBS 138744
Brachysporiella setosa HKUCC 3713
Bactrodesmiastrum obovatum FMR 6482
Savoryella paucispora SAT 00866
Melanotrigonum ovale CBS 138742
Canalisporium exiguum SS 00809
Pleurotheciella uniseptata DAOM 673210
Plagiascoma frondosum CBS 127885 T
Microglossum rufum OSC 100641
Melanotrigonum ovale M.R. 3685
Phragmocephala stemphylioides DAOM 673211
Canalisporium grenadoideum BCC 20507 T
Conioscypha lignicola CBS 335.93 T
Melanotrigonum ovale CBS 138815
100/1.0
99/1.0
98/0.96
97/1.0
99/1.0
99/1.0
95/0.97
87/0.96
72/0.97
91/1.0
100/1.0
99/1.0
90/1.0
98/1.0
98/1.0
96/1.0
99/1.0
99/1.0
92/1.0
97/0.96
98/1.0
93/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
100/1.0
Pleurotheciales Savoryellales
Conioscyphales
CladeVI
CladeV
CladeIV
CladeIII
CladeII
CladeI
CPSclade
Bactrodesmiatrum

clade
62/1.0

63
M.Réblováetal.:ThenewordersConioscyphales and Pleurotheciales
and Pleurothecium obovoideum. CladeIII(100/1.0)includes
Sterigmatobotrys and Taeniolella rudis.CladeIV(92/1.0)in-
cludes the monophyletic Pleurotheciella (100/1.0)andAdelo-
sphaeria catenata.
TAXONOMY
ConioscyphalesRéblová&Seifert,ord. nov.—MycoBank
 MB813226
Type family.ConioscyphaceaeRéblová&Seifert.
Ascomataperithecial,non-stromatic.Ostioleperiphysate.Hama-
theciumof paraphyses.Asci unitunicate, with a non-amyloid
apicalannulus.Ascosporeshyaline,transverselymultiseptate.
Conidiophores micronematous, mononematous. Conidioge-
nous cells blastic,percurrentlyregenerating.Conidia brown, va-
riableinshape;secessionschizolytic.Saprobiconwood.
ConioscyphaceaeRéblová&Seifert,fam. nov.—MycoBank
MB813227
Type genus.Conioscypha Höhn.,Ann. Mycol.2:58.1904, emend.
Shearer,Mycologia65:128.1973.
= Conioscyphascus Réblová&Seifert,Stud.Mycol.50:100.2004.
Ascomataperithecial,immersedtosuperficial,papillateorwith
elongated neck. Ascomatal wall leathery, waxy, comprising
twolayers.Paraphysesfiliform, unbranched,longer thanthe
asci.Asciunitunicate,persistent,8-spored,withapronounced
non-amyloidapicalannulus,cylindrical-clavate,stipitate.Asco-
spores fusiform to fusiform-navicular, hyaline, transversely multi-
septate,lackingamucilaginoussheathorappendages.Coni-
diophores micronematous,mononematous,hyaline.Conidio-
genous cells blastic,cyathiformtodoliiform.Conidia brown,
non-septate, often with a basal pore, formed singly and suc-
cessively by percurrent regeneration of the apex of the conidio-
genous cell, liberating by apical rupture of the outer wall of the
conidiogenouscell.
PleurothecialesRéblová&Seifert,ord. nov.—MycoBank
 MB813228
Type family.PleurotheciaceaeRéblová&Seifert.
Ascomataperithecial,non-stromatic.Ostioleperiphysate.Ha-
matheciumofparaphyses.Asci unitunicate, with a non-amyloid
apicalring.Ascospores hyaline or versicolorous with polar cells
hyalineandmiddlecellsbrown,transverselymultiseptate.Co-
nidiophores macronematous or semi-macronematous, loosely
fasciculateoraggregatedinindeterminatesynnemata.Conidio-
genous cells producing conidia holoblastically, monoblastic
or with sympodial extension, conidial secession rhexolytic or
schizolytic.Conidia hyaline or brown or versicolorous, septate
ornon-septate.Saprobicon wood, rarelyhumanpathogens
causingkeratomycosis.
PleurotheciaceaeRéblová&Seifert,fam. nov.—MycoBank
 MB813229
Type genus.Pleurothecium Höhn.,Ber.Deutsch.Bot.Ges.37:154.1919.
= Carpoligna F.A.Fernández&Huhndorf,Mycologia9:253.1999.
Ascomata perithecial, immersed to semi-immersed to super-
ficial,papillateorwithacentralrarelyeccentricneck.Ostiole
periphysate.Ascomatal wall leathery to fragile, carbonaceous,
brown,comprisingtwolayers.Paraphyses abundant, sparsely
branched,partiallydisintegrating,cylindrical.Asci unitunicate,
8-spored,withapronouncednon-amyloidapicalannulus,cylin-
dricalorcylindrical-clavate.Ascospores ellipsoidal to fusiform,
hyaline or versicolorous with polar cells hyaline and middle cells
brown, transversely multiseptate, lacking a mucilaginous sheath
orappendages.Conidiomata present or absent, when present
indeterminatesynnemataorloosefascicles.Conidiophores ma-
cronematous or semi-macronematous, sometimes regenerating
percurrently.Conidiogenous cells producing conidia holoblasti-
cally, conidial secession rhexolytic on short denticles or rachis
on sympodially extending polyblastic conidiogenous cells, or
schizolytic on monoblastic or solitary thallic conidiogenous
cells.Conidia hyaline, sometimes with protracted maturation of
the middle cells, which turn brown, or brown or versicolorous,
septateornon-septate.
Adelosphaeria Réblová, gen. nov.—MycoBankMB813230
Type species.Adelosphaeria catenataRéblová.
Etymology.Adelo-(Gk),meaningunclear,referringtothedifficultyofrecog-
nising this taxon among other morphologically similar fungi; sphaera(L)
meaningglobe,referringtoascoma.
Ascomata perithecial, non-stromatic, semi-immersed becoming
superficial, subglobose, dark brown, papillate. Ostiole peri-
physate.Ascomatal wallleatherytofragile,2-layered.Para physes
abundant, persistent, septate. Asci unitunicate, cylindrical-
clavate,stipitate, 8-spored, apex with a non-amyloid apical
annulus.Ascospores ellipsoidal, slightly curved, hyaline, trans-
verselyseptate.Asexual morph unknown.
Adelosphaeria catenataRéblová, sp. nov.—MycoBank
 MB813231;Fig.3,4
Etymology.Cateniformis (L),meaningchain-shaped,referringtothedark
brown cells arranged in chains formed on vegetative hyphae in the axenic
culture.
Ascomata perithecial, non-stromatic, semi-immersed, becom-
ingsuperficial,solitaryorinsmallgroups;venter200–280µm
diam,300 – 360µmhigh, subglobose,darkbrown, glabrous,
papillate,opening by a rounded pore. Ostiole periphysate.
Ascomatal wallleatherytofragile,20– 30µmthick,2-layered;
outer layer consisting of brown, polyhedral cells of textura pris-
maticawithopaquewalls,innerlayerconsistingofseveralrows
ofthin-walled,hyaline,flattenedcells.Paraphyses abundant,
persistent, septate, hyaline, sparsely branched, anastomosing,
c.3.5–5.0µmwide,taperingtoc.2.5µm.Asci(85 –)93–105
µmlonginthesporiferouspart,12.5 –14.5µmwide(mean±
SD=199.7±5.4×12.5±1.2µm),withastipe20 –35(– 50)
µmlong; cylindrical-clavate, broadly rounded apically to
obtuse,8-spored, apex with a flattened, non-amyloid apical
annulus 3.0 –3.5 µm wide, about 2.0 µm high. Ascospores
16.5–19.5(–20)×5.0– 5.5(–5.8)µm(mean±SD=17.8±1.3
×5.4±0.2µm),ellipsoidal,straightorslightlycurved,hyaline,
smooth,3-septate,non-constrictedatthesepta,arranged1–2-
seriatelyinthesporiferouspart.
 Culturecharacteristics—Colonies slowgrowingreaching
12–15mmdiamonPDAafter21dat25°C.Aerialmycelium
darkbrown(aoc735),palerbrown(aoc723)towardsthemargin,
mainlyflat,felty,reversebrown(aoc734),withamarginalzone
ofdarkbrown(aoc734)submergedmycelium.Aerial and sub-
mergedhyphae1.5– 2.0µmwide,smooth,subhyalinetopale
brown,thin-walled,unbranchedorsparselybranched.Sporula-
tionabsent.Onvegetativehyphaeareformedbrown,globose
toellipsoidalcells5.0–14.5µmdiam(mean±SD=10.3±2.9
µm),withthick,oftenopaquewalls,arrangedinchainsoraris-
inglaterallyonanothercell.
Specimen examined.CzeCh RepubliC,SouthernBohemia,Novohradské
horyMts,Dobrávoda,HojnávodaNationalnaturemonument,decorticated
wood of a trunk of Fagus sylvatica, 4Oct.2012, M. Réblová M.R. 3755
(holotypePRM933853,cultureex-typeCBS138679).

64 Persoonia–Volume37,2016
Fig. 3 Adelosphaeria catenata.a,b.Ascomata;c.verticalsectionoftheascomatalwall;d–f.asciwithascospores;g,h.apicalannulus;i.paraphyses(a– i.
PRM933853holotype);d– f,h:DIC;g,i:PC.—Scalebars:a,b=250µm;c=30µm;d –f,i=10µm;g,h=5µm.

65
M.Réblováetal.:ThenewordersConioscyphales and Pleurotheciales
 Notes—Adelosphaeria catenata resembles species of Pleu-
rothecium and Pleurotheciellabecauseofitshyaline,3-septate
ascospores, cylindrical-clavate asci and brown semi-immersed
ascomata.Pleurothecium recurvatum is easily distinguishable
from A. catenata by its slender ascospores, pronounced apical
annulus, setose ascomata and macronematous conidiophores
bearing hyaline, polyblastic, denticulate conidiogenous cells
elongatinginasympodialpatternand3-septate,hyalineconidia
withprotractedmaturationofthemiddlecells,whichturnbrown.It
ismoredifficulttoseparateAdelosphaeria from Pleurotheciella,
because species of both genera share similar morphological
charactersofascospores,asciandascomata.Theonlyconspic-
uous difference lies in morphology of their asexual states; Pleuro-
theciella can be easily distinguished by Dactylaria-like, hyaline
tosubhyalineconidiophoresandconidia.
Melanotrigonum Réblová,gen.nov.—MycoBankMB813232
Type species.Melanotrigonum ovaleRéblová.
Etymology.Melas-(Gk),meaningdark, referringtothebrownconidia;
Trigonon(Gk)meaningtriangle,referringtoconspicuoustriangle-likeconi-
diogenouscellsoftheasexualmorph.
Ascomata perithecial, non-stromatic, immersed to semi-im mers-
ed,subglobosetobroadlyconical,darkbrown,papillate.Ostiole
periphysate.Ascomatal wallleatherytofragile,2-layered.Para-
physesabundant,persistent,septate.Asci unitunicate, cylindri-
cal,stipitate,8-spored,apexwithaconspicuous,non-amyloid
apicalannulus.Ascospores ellipsoidal, hyaline, transversely
septate.Asexual morph a dematiaceous hyphomycete, conidio-
phores semi-micronematous, conidio genous cells producing
brown,1-septateconidiaholoblasticallyonshortdenticles.
Melanotrigonum ovaleRéblová, sp. nov.—MycoBank
 MB813233;Fig.5,6
Etymology.Ovalis (L),referringtotheovalshapeofconidia.
Ascomata perithecial, non-stromatic, immersed to semi-im-
mersed, gregarious, occurring in small to large groups; venter
320–480µmdiam,400 – 500µmhigh,subglobosetobroadly
conical,brown,glabrous,papillate,openingbyaroundedpore.
Ostioleperiphysate.Ascomatal wallleatherytofragile,23– 30
µmthick,2-layered;outerlayerconsistingofbrown,polyhedral
cells of textura prismaticawithopaquewalls;towardstheexte-
rior grading into polyhedral to angular cells of textura angularis;
towardstheinteriorgradingintopale-brown,elongatedcells.
Innerlayerconsistingofseveralrowsofthin-walled,hyaline,
flattened cells. Paraphyses abundant, persistent, septate,
anastomosing,hyaline,sparselybranched,c.3.0–4.5µmwide,
taperingto c.3.0µm,longerthantheasci.Asci(105 –)115–
128(–142)µmlonginthesporiferouspart,(8.5–)9.0 –11.5µm
wide(mean±SD=122.8±7.4×11.0±5.2µm)withastipe
32 –50 µm long; cylindrical, obtuse apically, 8-spored, apex
withalarge,conspicuousnon-amyloidapicalannulus4.5– 5.0
µmwide,3.5– 4.0µmhigh.Ascospores(17–)18–21(–21.5)×
5.0–6.0(–6.5)µm(mean±SD=19.4±1.5×5.8±0.3µm),el-
lipsoidal,straighttoslightlycurved,hyaline,smooth,3-septate,
non-constricted at the septa, arranged obliquely uniseriate,
sometimes2-seriateonlyintheupperpartoftheascus.
 Culturecharacteristics—Coloniesslowgrowing,reaching
8–10mmdiamon PDAafter21dat25°C.Aerialmycelium
beige in the centre of the colony and on the inoculum block,
white towards the margin, felty, centre elevated, later with a
moistappearance,bent intodeepfolds,reverse darkbeige.
Fig. 4 Asexual morph of Adelosphaeria catenata.a.ColonyonPCA;b– f.browncellsarisingblasticallyfromvegetativemyceliumonPCA(a– f.CBS138679,
21d,25°C);b– f:DIC.—Scalebars:a=4mm;b –f=20µm.

66 Persoonia–Volume37,2016
Fig. 5 Melanotrigonum ovale.a.Ascomata;b,c.verticalsectionsoftheascomatalwall;d–f.asciwithascospores;g.apicalannulus;h.ascospores;i.para-
physes(a– i.PRM933852holotype);b –h:DIC;i:PC.—Scalebars:a=250µm;b,c=25µm;d – f,i=20µm;g,h=10µm.

67
M.Réblováetal.:ThenewordersConioscyphales and Pleurotheciales
Fig. 6 Asexual morph of Melanotrigonum ovale.a,b.ColonyonPDA(5mo,25°C);c,d.colonyonPDA(1mo,25°C);e,f,h,i.conidiogenouscellsonPCA;
g,j.conidiogenouscellswithconidiaborneonadenticleonPCA;k–n.conidia(a,b,g– i,k –n. M.R.3685;c, d– f,j. CBS138742;e – j.21d,25°C); e– n:
DIC.—Scalebars:a=2.5mm;b=5mm;c,d=10mm;e– n=5µm.

68 Persoonia–Volume37,2016
Fig. 7 Plagiascoma frondosum.a.Ascomata;b.verticalsectionoftheascomatalwall;c,d.germinatingascospores;e.ascospores;f.paraphyses;g.asci
infreshlycollectedmaterialinPelikanink;h,i.ascifromairdriedherbariummaterial,arrowindicatesapicalannulus;j.apicalannulus;k,l.pigmentedcells
formedinvitroonvegetativehyphaeonPCA(a–j.PRM933854holotype;k,l.CBS139031,21d,25°C);c–e,h– l:DIC;f,g:PC.—Scalebars:a=500µm;
b=250µm;c– f,j=10µm;g=25µm;h,i,k,l=20µm.

69
M.Réblováetal.:ThenewordersConioscyphales and Pleurotheciales
Aerialhyphae2.0–3.0µmwide,smooth,hyaline,thin-walled,
sparsely branched. Submerged hyphae 2.0 –2.5 µm wid e,
smooth,hyaline. Sporulation appears later on the youngest
aerialhyphaeatthemarginofthecolony.Conidiophoressemi-
micronematous, reduced to a conidiogenous cell, arising verti-
cally from hyphae, unbranched, smooth, tapering towards the
apex.Conidiogenouscell4.5–8.0(–10.0)µmlong,2.0– 3.0µm
wideinthebroadestpoint(mean±SD=6.9±1.1×2.8±0.4
µm),integrated, intercalary,almosttriangularto ampulliform,
tapering towards the apex, pale brown, with a single, rarely two,
palebrowntosubhyalinedenticle1.0 –2.0µmlong.Conidia
(10–)11.5–13.5(–14)×5.0–6.0µm(mean±SD=12.3±0.6×
5.4±0.5µm),1-septate,ovaltobean-shaped,straightorslightly
curved, leaving a pore when detached, smooth, rounded at both
ends or slightly tapering towards the base, brown, darker at the
septum,non-constricted,sometimescellsasymmetrical.
Specimens examined.CzeCh RepubliC,SouthernMoravia,Břeclavdistr.,
Valtice,RendezvousValticeNationalnaturemonument,decayingwoodofa
trunk of Quercus cerris, 17Nov.2012,M. Réblová M.R. 3698(holotypePRM
933852,cultureex-typeCBS138743);ibid.,M.R. 3685, M.R. 3688A (culture
CBS138742),M.R. 3688B (cultureCBS138744),M.R. 3699(cultureCBS
138815).
 Notes—Fivestrains of M. ovale were collected on soft,
strongly decaying wood of several fallen trunks of Quercus
cerris, the remains of old growth trees that were more than
hundredyearsold. SpecimenM.R. 3685 has asci shorter in
thesporiferouspart(100–105µmlong)andgenerallysmaller
ascospores,(15–)16–17.5(–19.5)×4.5–5.5µm.
Melanotrigonum ovale is similar to Pleurotheciella rivularia
in characters of ascospores, asci and ascomata, but differs
from it by the ascal apex with a conspicuous and larger apical
annulus,4.5 –5.0×3.5– 4.0µm(w×h)vs2.5 – 3.5×1.5µm
in P. rivularia. Both  species produce conidia on denticulate
conidiogenouscells. In Melanotrigonum, the conidiogenous
cells are almost triangular to ampulliform, tapering towards the
apex with a single or rarely two denticles, while conidiogenous
cells of Pleurotheciella are cylindrical, elongate sympodially
withonetoseveraldenticles.
Plagiascoma Réblová&J.Fourn.,gen. nov.—MycoBank
 MB813234
Type species. Plagiascoma frondosumRéblová&J.Fourn.
Etymology.Plágios(Gk),meaningslanting,oblique,sideways,referring
totheflattenedascomataarrangedhorizontallytothehost.
Ascomata perithecial, non-stromatic, immersed gradually erum-
penttosemi-immersed, conical, dark brown, lying obliquely
tohorizontally,papillateorwitha neck. Ostiole periphysate.
Ascomatal wallfragile,2-layered.Paraphyses abundant, per-
sistent,septate. Asci unitunicate, cylindrical to cylindrical-
fusiform,stipitate,8-spored,apexwith a non-amyloid apical
annulus.Ascosporesfusiform, hyaline,transverselyseptate.
Asexual morphunknown.
Plagiascoma frondosumRéblová&J.Fourn.,sp. nov.—Myco-
BankMB813235;Fig.7
Etymology.Frondosus(L),meaningleaf-bearing,referringtoadeciduous
treeasahost.
Ascomata non-stromatic, immersed, gradually erumpent to
semi-immersed, solitary or in small groups or in rows; venter
200–280 µm diam, 450 –550µmhigh,conical, dark brown,
glabrous,slightlypinchedlaterally,lyingobliquelytohorizon-
tally,papillateorwithabeak30–120µmhigh,conical,lateral,
openingbyaroundedpore. Ostiole periphysate. Ascomatal
wallfragile,24 –30µmthick,becomingthickerinthebeakupto
c.35µm,2-layered;outerlayerconsistingofbrown,polyhedral,
flattened cells of textura prismaticawithopaquewalls.Inner
layer consisting of several rows of thin-walled, hyaline, flattened
cells.Paraphysesabundant,persistent,septate,hyaline,c.4.0–
6.0(–7.0)µmwide,taperingtoc.3.5µm.Asci in fresh material
225–240µmlonginthesporiferouspart,13–15µmwide,with
astipe30–52µmlong,cylindrical,withascosporesarranged
obliquelyuniseriate;upondryingasci100–160µmlonginthe
sporiferouspart,15–20µmwidewithastipe53 –73µmlong,
cylindrical-fusiform, with ascospores arranged 2-seriately;
obtusetobroadlyroundedapically,8-spored;apexwithanon-
amyloidapicalannulus4.5– 5.5 µm wide,1.5–2.5 µm high.
Ascospores(28.5–)30– 34.5(–36)×7.5– 8.5(–9.0)µm(mean
±SD=32.8±1.9×8.4±0.6µm),fusiform,taperingtowards
theends,hyaline,smooth,3 – 5-septate,non-constrictedatthe
septa,withalargeguttuleineachcell.
 Culturecharacteristics—Coloniesslowgrowing,10–15mm
diamonPDAafter21dat25°C.Aerialmyceliumbrownnear
the centre of the colony and on the inoculum block, pale brown
tobeige (oac800) towards the margin, felty,reversebrown
(oac734).Aerial and submerged hyphae 1.5– 2.5 µm wide,
smooth,subhyaline,thin-walled,sparselybranched.Sporula-
tionabsent.Onaerialhyphaeariseellipsoidalcells5.5–9.0µm
diam(mean±SD=7.0±1.0µm),palebrowntosubhyaline,
thick-walled,intercalar,terminalorarrangedinashortchain.
Specimen examined. FRanCe,Midi-Pyrénées,Ariège, Rimont,valleyof
LaMaille brook,c.550 masl,submerged decorticatedwood ofFraxinus
excelsior, 9May2014,J. Fournier J.F. 14044(holotypePRM933854,culture
ex-typeCBS139031).
 Notes—TheexaminationoffreshmaterialofP. frondosum
revealedasciover200µmlongand13–15µmwide,withunise-
riateascosporesarrangedobliquely(Fig.7g).Upondrying,the
arrangement of ascospores changes and they became biseriate
withintheascus.Theasciindryherbariummaterialareshorter
inthesporiferouspart,100–160µmlong,andwider15–20µm
withalmosttwicethestipelength(Fig.7h,i).Nosheathorap-
pendageswereobservedonimmatureormatureascospores.
Freshwater perithecial ascomycetes often have ascospores
enclosed in a hyaline sheath or have appendages to facilitate
theirattachmentonmoistwoodysubstrates.Interestingly,this
is largely true for species from Asia, America and Australia but
not in Europe, where many of the most widespread freshwater
specieslackthesestructures.
Thefusiform,hyaline,3–5-septateascosporesofP. frondosum
resemble multiseptate ascospores of some species of Annu-
latascus,e.g.A. nilensis (Abdel-Wahabetal.2011)andA. tro-
picalis (Tsuietal.2002).Inourmultilocusphylogeny,P. frondo-
sum is positioned in the strongly supported Bactro desmiastrum
clade.
Phaeoisaria fasciculataRéblová & Seifert, sp. nov.—Myco-
BankMB813236;Fig.8
Etymology.Fasciculus (L),meaningfascicleorbundle,referringtoconi-
diophoresarrangedinfasciclesandlackingadistinctstipe.
Colonies in vivo effuse, dark grey, whitish to beige when sporu-
lating.Sexual morphnotobserved.Synnemata absent, conidio-
phoresformingfascicles. Conidiophores 25–65×3.0–3.5µm
(mean±SD=41.7±14.2×3.3±0.3µm),macronematous,
arising from brown, thick-walled cells, cylindrical, pale brown,
subhyalinetowardstheapex,unbranched,smooth-walled.Coni-
diogenous cells10–29(– 36)×2.5 – 3.5µm(mean±SD=20.2
±6.7×3.1±0.5µm),integrated,terminal,cylindrical,taper-
ing towards tip, pale brown to subhyaline near base, hyaline
towards apex, smooth-walled, polyblastic, forming conidia sym-
podiallyonconspicuousdenticles1.0–1.5µmlong,about0.5
µmwide,scatteredorclusteredintheapicalregion.Conidia

70 Persoonia–Volume37,2016
Fig. 8 Phaeoisaria fasciculata.a.ColonyonPCA;b.conidiophoresinvivo;c.conidiainvivo;d,f,g –i.conidiophoresonPCA;e.conidiaonPCA(a,d–f.
CBS127885;b,c.PRM933855holotype;g– i.DAOM230055;a,d– i.21d,25°C);b–f:DIC;g –i:ESEM.—Scalebars:a=50µm;b=250µm;b,d,f=20
µm;c,e=10µm;g– i=10µm.

71
M.Réblováetal.:ThenewordersConioscyphales and Pleurotheciales
6.0–8.0(–9.0)µmlong,about2.0µmwide(mean±SD=7.3
±1.2×2.0±0.1µm),ellipsoidaltoobovoid,straight,rounded
at the apex, obtuse and tapering towards base, hyaline, non-
septate,smooth-walled.
 Culturecharacteristics—Coloniesreaching12 –18mmdiam
onPCAafter21dat25°C.Aerialmyceliumbeigetopalebrown
(oac662),atfirstsmooth,latercottony,reversebrown(oac640).
Aerial and submergedhyphae2.0 – 3.0µmwide,hyalinetopale
brown,sparselybranched,smooth-walled.Sporulationappears
firstinthecentreofthecolony,laterpresentover thewhole
colonyorinisolatedpatches;sporulatingcolonybeige(oac809)
withapowderyappearance.Synnemata absent.Conidiophores
20–75× 2.5 –3.5µm(mean ±SD=45.3±17.0 ×3.0±0.3
µm),macronematoustosemi-macronematous reduced to a
single conidiogenous cell, arising from aerial hyphae, cylindri-
cal, slightly tapering towards the apex, pale brown, subhyaline
towardstheapex,unbranched,smooth-walled.Conidiogenous
cell10– 29(–36)×2.5– 3.5(–4.0)µm(mean±SD=20.2±6.7
×3.1±0.5µm),integrated,terminalandintercalar,cylindrical,
tapering towards tip, pale brown to subhyaline, hyaline towards
apex, smooth-walled, polyblastic, with numerous conspicuous
denticles1.0–1.5µmlong,c.0.5µmwide,scatteredalongthe
whole length of intercalar conidiogenous cell and clustered in
theapicalregion.Conidia5.5–7.5(–8.5)×2.0 – 2.5(– 3.0)µm
(mean±SD=6.7±0.9×2.5±0.3µm),ellipsoidaltoobovoid,
straight, rounded at the apex, obtuse and tapering towards
base,hyaline,non-septate,smooth-walled.
Specimens examined.Canada, Ontario,GoulbournTwp.,Stittsville,bark
onbranchonground, 8 Oct. 2001, Keith A. Seifert K.A.S. 1433(DAOM
230055). – CzeCh RepubliC, Southern Moravia, Břeclav distr., Milovice,
MilovickástráňNatureReserve,northslopesofMtŠpičák,293masl,de-
corticated wood of Sambucus nigra, 18Nov.2009,M. Réblová M.R. 3084
(holotypePRM933855,cultureex-typeCBS127885).
 Notes—Phaeoisaria fasciculata is easily distinguished from
other species of the genus by its conidiophores, which grow in
fascicles on the host, while typical indeterminate synnemata are
notformed.Theellipsoidaltoobovoid,non-septateconidiaof
Ph. fasciculata resemble those of Ph. caffera, the Ph. clematidis
species complex and Ph. magnifica. Phaeoisaria caffera differs
fromthenewspeciesbylonger,paleyellowishbrownconidia.
The conidia of Ph. clematidis and Ph. magnifica are subhyaline
toverydiluteolivaceousandgenerallywider.
Ascotaiwania limnetica(H.S.Chang&S.Y.Hsieh)Réblová
&J.Fourn.,comb. nov.—MycoBankMB813237;Fig.9,10
Basionym.Savoryella limneticaH.S.Chang& S.Y. Hsieh,Mycol.Res.
102:715.1998.
Ascomata perithecial, non-stromatic, semi-immersed, gradually
erumpenttoalmostsuperficial,scatteredorclusteredinsmall
groupsof2 –3,upright,obliquelyorientedorlyinghorizontallyon
thehost;venter210– 260µmdiam,220– 250(–300)µmhigh,
black, subglobose with a flattened base and a broadly conical
apex, often laterally flattened, flask-shaped when lying horizon-
tally, with a papilla or short neck, broadly conical or cylindrical,
apically truncate, central, eccentric or lateral, oriented upwards
whenascomataliehorizontally. Ostioleperiphysate.Ascomatal
wallfragile, 9–15µm thick,thickerattheapexupto 20µm,
2-layered; outer layer consisting of dark brown, polyhedral, flat-
tened cells of textura prismaticawithopaquewallsandsparse
pores, outwards grading into small protruding cells, inner layer
consistingofseveralrowsofthin-walled,hyaline,flattenedcells.
Asci125 –150×11–14µm(mean±SD=137±9.4×12.6±
1.2µm),cylindrical,short-stipitate,broadlyroundedapicallyto
obtuse,withanon-amyloid,discoidapicalannulus4.5–5.5µm
wide,1.0–2.0µmhigh.Paraphyses sparse, partially disintegra-
tingatmaturity,septate,branching,anastomosing,4.0– 9.5µm
wide.Ascospores(17.5 –)19.5–23.5(–24)×(6.3–)7.0–8.5µm,
(mean±SD=21.4±1.4×7.7±0.5µm),ellipsoidal,equilateral,
straight, versicolorous, middle cells olivaceous brown to brown,
containing numerous small guttules, polar cells smaller, hya-
line,smooth,unequally 3-septate, slightly constricted at the
septa,withoutsheathorappendages,arrangedobliquelyuni-
seriatelyintheascus.Colonies in vivo diffuse, visible only as
single scattered macroconidia arising from short, hyaline conidio-
genouscellsonvegetativemyceliumnearascomata.Conidia
(30–)33–41×15 –17.5µm,ellipsoidal,broadlyroundedatthe
apex,tapering basally,darkbrown,opaque,basalcell sub-
hyalinetopalebrown,(3 –)5– 6-septate,septaobscuredbya
darkerband.
 Culturecharacteristics—Coloniesslowgrowing,reaching
c.8–10mmdiamonPDAafter21dat25°C.Aerialmycelium
brown(oac639),palebrown(aoc661)inthecentreofthecolony
andoninoculumblock,velvety,reversebrown(oac733).Aerial
hyphae smooth, thin-walled, sparsely branched, hyaline to sub-
hyaline1.5–2.0µm,submergedhyphaesometimespalebrown
2.0 –3.0 µm wide. Conidiophores reduced to a monoblastic
conidiogenouscell.Conidiogenouscells4.5–7.0×5.0– 8.0µm,
usually with several subtending cells, integrated, hyaline to sub-
hyalinewithasingleconidiogenouslocus.Conidia32–36(–39)
×(14.5–)16–17.5(–18.5)µm(mean±SD=34.5±2.2×17.0
±1.0µm),ellipsoidaltoobovoid,straightorslightlycurved,
smooth,darkbrown,3– 5-septate,withdarkerbandsobscuring
the septa, non-constricted at the septa, basal cell subhyaline
3.0–4.5µmwidetaperingto2.5 – 3.0µm.
Specimens examined.FRanCe,Midi-Pyrénées,Ariège,Rimont,valleyof
thePeyraubrook,c.400masl,23Feb.2008,onsubmergedwood,J. Four-
nier J.F. 08011(PRM933849, culture CBS126792);ibid., 22May2009,
submerged wood of Alnus glutinosa, J. Fournier J.F. 09127(PRM933851,
cultureCBS126576);ibid.,19Apr.2010,submergedwoodofFraxinus ex-
celsior, J. Fournier J.F. 10014;ibid.,Vernajoul,Vernajoulbrook,PontFagé,
c.350masl,onunidentifiedsubmergedwood,2July2007,J. Fournier J.F.
07123(PRM933850).
 Notes—Savoryella limnetica was originally collected on de-
caying wood submerged in freshwater in Taiwan and assigned
tothegenusbasedonits3-septateascosporesandflattened
apicalapparatus(Changetal.1998).Thisspecieswasrecently
repeatedly collected on submerged deciduous wood in southern
France.Twolivingculturesweresuccessfully obtained from
isolatedascosporesfromfreshmaterial.
Savoryella and Ascotaiwania are closely related, morphologi-
cally similar genera and their delimitation is based primarily on
ascospore septation, morphology of the apical apparatus of
theascusandwidthoftheparaphyses(seeDiscussion).The
transfer of S. limnetica to Ascotaiwania is supported by molecu-
lardataandculturecharacters.ThemajorityofAsco taiwania
specieshave5 –7-septateascosporesandonlyfewarecharac-
terisedbyascosporeswiththreesepta,i.e.A. hughesii, A. pal-
micola and A. sawadae. Helicoön farinosum and its sexual
morph described as A. hughesii (Fallahetal.1999),isamem-
ber of the Pleurotheciales.Ascotaiwania palmicola differs from
A. limneticabyterrestrialhabitatandaffiliationtopalmwood,
asciwithaconspicuousapicalapparatus4×5µmandslender
ascospores,17.5– 20×5.0– 6.5µmwith polarmucilaginous
appendages(Hyde1995).Ascotaiwania sawadae can be com-
pared to A. limnetica by ascomatal morphology, but differs by
asciwithalessflattenedapicalapparatusandlargerandinequi-
lateralascospores25–30×7.5 –10µm(Sivichaietal.1998).
When observed in Congo red, the asci of A. limnetica revealed
a conspicuous flattened apical annulus that stains deep red
(Fig.9k).

72 Persoonia–Volume37,2016
Fig. 9 Ascotaiwania limnetica.a,b.Ascomata,arrowindicatesascosporesaggregatedatthetopoftheneck;c,d.verticalsectionsoftheascomatalwall;
e.asciwithascosporesinPelikanink;f.asciwithascospores;g– i.paraphyses;j.apicalannulus,arrowindicatesthetipofascalapex,whenascosporeis
releasedthroughtheannulus;k.asciwithapical annulusinCongored(a –e,k.PRM 933850;f,g,i,j.PRM933851;h. PRM933849);c– f,j,k:DIC;g –i:
PC.—Scalebars:a,b=150µm;c,j=10µm;d=100µm;c,e=10µm;e,f,k=50µm.

73
M.Réblováetal.:ThenewordersConioscyphales and Pleurotheciales
Fig. 10 Asexual morph of Ascotaiwania limnetica.a.Conidiainvivo;b.ascomawithmacroconidiascatteredonwoodsurface;c–h.conidiaandconidiogenous
cellsonPDA;i.colonyonPCA;j– m.conidiaonPCA(a,c– h.CBS126576;b.PRM933850;j–m.CBS126792;a,c –m.21d,25°C);a,c– h,j,k:DIC;l,m:
ESEM.—Scalebars:a,c– h,j,k,l=20µm;b=250µm;i=5mm;m=10µm.

74 Persoonia–Volume37,2016
Pleurotheciella uniseptata (Matsush.)Seifert,comb. nov.—
MycoBankMB813238;Fig.11
Basionym.Dactylaria uniseptata Matsush., Microfungiofthe Solomon
IslandsandPapua-NewGuinea:19.1971.
Colonies in vivo effuse,visibleassolitaryto4–5caespitose
darkbrownconidiophoreswithdry,whitishtogreyishconidia.
Sexual morphnotobserved.Conidiophores mostly100–150µm
tall,4.5–5.0µmwideatthebase,taperingto3.0–4.0µmwide,
macronematous, unbranched, straight or sinuous, dark brown
atthebase,withwallsupto1.0µmthicknearthebase,thinner
towards the apex, cylindrical, smooth-walled or slightly granular
or roughened, usually with a terminal node of denticles, but
rarely extending through the original node with a new exten-
sionoftheconidiophore.Conidiogenous cell15 –32µmlong,
2.5–3.5µmwideatthebase,2.0– 3.0µmwidebelowthefertile
zone, integrated, terminal, cylindrical or tapering towards tip,
pale brown to subhyaline near base, hyaline towards apex,
smooth-walled or slightly granular, polyblastic, forming conidia
sympodiallyonconspicuousdenticles1.0–2.0µmlong,about
0.5µmwide,sometimesslightlybroaderatbase,occluded,fer-
tilezoneatfirstjustafewdenticles,butcanexpandintoanode-
like zone that is cylindrical to ellipsoidal in outline, usually with
compactclustersof4–15denticlesbutsometimesextended,
rarelygeniculate,upto5.0– 9.0 × 3.0 µm, wide, or becon-
stricteddownto1.5µm,upto15denticlesseen.Conidia12.5–
16.5×2.0 –4.0µm(mean±SD=14.1±0.9×2.9±0.5µm),
fusoid or slightly clavate, straight, rounded at the apex, obtuse
and tapering towards base, hyaline, 1-septate with an incon-
spicuouscentralseptum,oftenwith1–2largeguttulesineach
cell, smooth-walled, remains of denticle sometimes attached
tosecededconidium.
 Culturecharacteristics—Coloniesreaching8–10mmdiam
onCMAafter21dat25°C.Aerialmyceliumabsent,colonyand
reverseinconspicuoustowhite.Submergedhyphae1.5 – 2.0
µmwide,hyaline,smooth-walled.Sporulationappearsfirston
the inoculum of the colony, and later is sparsely present on
the older parts of the new growth. Conidiophores 50 –85 ×
3.5–4.0µmwide,slightlyswollenatbasetoabout4µm,semi-
macronematous, pale brown, subhyaline towards the apex,
unbranched,smooth-walled.Conidiogenouscellsandconidia
similartothoseproducedinvivo.
Specimen examined.Canada,Ontario,Arnprior,MacNamaraTrail,onde-
cayingwetwood,12Oct.2011,K.A. Seifert & G. White K.A.S. 4459(DAOM
673210,cultureDAOMC250294).
 Notes—Pleurotheciella uniseptata is known only from its
asexual morph. Its occurrence on water saturated decayed
wood is consistent with the ecology of the other two species
nowclassifiedinthisgenus.Itsconidiaareofasimilarlength
Fig. 11 Pleurotheciella uniseptata.a,b.Conidiophoresinvivo;c.conidiainvivo;d.conidiophoresonCMA;e.colonyinvivo(a –c.DAOM673210,d.DAOMC
250294,21dat25°C);a:PC;b– d.DIC.—Scalebars:a –d=10µm;e=100µm.

75
M.Réblováetal.:ThenewordersConioscyphales and Pleurotheciales
and septation to those of P. rivularia, but narrower and more
uniformly fusiform rather than the often obovoidal shape of the
latterspecies.TheconidiaofP. centenaria are also fusoid, but
longerthantheothertwospeciesandconsistently3-septate.
OurspecimenfromCanadafitsthedescriptionandillustration
of Dactylaria uniseptata byMatsushima(1971)well,considering
that the protologue was based on a culture grown on banana
leafagar.WenotethatDeHoog (1985) failed to obtainthe
holotype of D. uniseptata and we did not attempt to obtain it
here.WehaveresistedthetemptationtoepitypifyaJapanese
specieswithaCanadianspecimenandculture.Lectotypification
with the drawings from the protologue would be a precondition
toepitypificationiftheholotypeistrulyunavailable.
ThisisthefirstspeciesofPleurotheciella for which the coni-
diophoreshavebeenobservedonthenaturalsubstrate.The
protologue of the genus suggested that the conidiophores were
dactylaria-like, but in P. uniseptata the conidiophores are macro-
nematous and much more similar to those of Pleurothecium
species. However,theconidiophoresof P. uniseptata produced
in culture lack dark basal cells and are rather similar to those
produced by P. rivularia and P. centenaria in vitro(Fig.11d).
ItseemspossiblethattheconidiophoresorallPleurotheciella
species would be macronematous in vivo. Morphologically,
there are few if any characters to distinguish between the
asexual morphs of Pleurotheciella uniseptata and some species
classifiedinthe hyphomycetegenusPleurophragmium. The
two genera are clearly phylogenetically distinct, with Pleuro-
phragmium parvisporum, thetypeofthatgenus,classifiedin
the Papulosaceae, SordariomycetesbyRéblová(2009).Agreat
morphologicaldiversityofspeciesareclassifiedinPleurophrag-
mium(seekeyinD’Souza&Bhat2012)anditisunlikelytobe
phylogeneticallyhomogeneous.
DISCUSSION
The CPS (Conioscyphales/ Pleurotheciales/Savoryellales)
clade
The combined three- and six-gene phylogenetic analyses of the
newly described genera Melanotrigonum and Adelosphaeria
with members of the Savoryellales and other taxa related to Co-
nioscypha and Pleurothecium revealed a robust monophylum in
the Hypocreomycetidae (Fig.1).Itcontainsthreenestedmono-
phyleticcladessignificantlysupportedbyBIandMLmethods,
namelyi)theSavoryellales;ii)acladecontainingfivespecies
of Conioscypha; andiii)anothercladethatcomprisesseveral
genera centred around Pleurothecium.Thetwolatterclades
represent distinct taxonomic groups at the ordinal systematic
level and are introduced as the Conioscyphales and Pleurothe-
ciales above.Asisterrelationshipwasrevealedbetweenthe
CPS(Conioscyphales/Pleurotheciales/Savoryellales) clade
and a monophyletic strongly supported lineage of uncertain sys-
tematic position containing Ascotaiwania persoonii, Bactro-
desmiastrum and the new genus Plagiascoma.
MembersoftheCPS and Bactrodesmiastrum clades share a
few morphological features such as the absence of stromatic
tissue or clypeus, similar anatomies of the ascomatal walls,
thin-walled unitunicate asci with a distinct, non-amyloid apical
annulus, paraphyses and symmetrical, transversely septate
ascospores.Theknown asexual morphs are dematiaceous
hyphomyceteswithholoblasticconidiogenesis.Although the
morphology of sexual morphs is more or less uniform and
rather nondescript within each order, the observed variability
in extension of conidiogenous cells and conidial morphology is
characteristicofeachorder.IntheCPSclade,pleomorphismis
commonlyobserved,i.e.theabilityoffungitoreproducesexu-
ally and asexually and form independent spore-stages in the life
cycle.Allknownlife-historiesdiscussedherewereestablished
experimentally,i.e.Ascotaiwania (Ranghoo&Hyde1998,Sivi-
chaietal.1998,thisstudy),Canalisporium (Sri-indrasutdhiet
al.2010),Conioscypha (Réblová&Seifert2004,Zelskietal.
2014),Helicoön farinosum (Fallahetal.1999),Pleurothecium
(Fernándezetal.1999),Pleurotheciella(Réblováetal.2012),
Sterigmatobotrys(Réblová&Seifert2011)andthenewgenera
describedinthisstudy.
At the base of the monophyletic clade with the nested CPS and
Bactrodesmiastrum clades, Flammispora bioteca is positioned
onaseparatebranch(Fig.1,2).Thisspecieswascollectedon
submerged leaves of the peat swamp palm Licuala longecaly-
cata and is characterised by non-stromatic, black, immersed
ascomata,clavatedeliquescentasciwithoutanapicalannulus,
subcylindrical to elongate-fusiform ascospores with a polar
appendageandabsenceofparaphyses(Pinruanetal.2004).
Itsasexualmorphisunknown.
The Bactrodesmiastrum clade
Bactrodesmiastrum, based on B. obscurum, was described by
Holubová-Jechová(1984) for dematiaceous hyphomycetes
characterised by schizolytic conidial secession and a forma-
tion of conidiogenous cells related to the maturation of brown,
septateconidia.Whentheconidiummaturesatthetipofthe
conidiogenous cell, a new monoblastic conidiogenous cell is
borne near the previous one on repent basal hyphae, followed
by formation of other conidiogenous cells in the same man-
ner.NoDNAsequencesareavailableofthetype speciesof
Bactrodesmiastrum.Thesexualstateof Bactrodesmiastrum
isunknown (Holubová-Jechová 1984, Hernández-Restrepo
etal.2013,2015)andnoconidiaorconidiogenouscellswere
observed on the type and other herbarium material of its closest
ascoma-producing sibling A. persoonii(Fallahetal.1999).We
prefer to avoid proposing a new genus for A. persoonii or its
new combination in Bactrodesmiastrum,basedoncurrentDNA
sequencedata,untilsimilaritiesinthelifehistoriesofthesetwo
taxaareprovenordisprovenexperimentally.
Conioscyphales
The Conioscyphales comprises a single genus Conioscypha
with12speciesfromfreshwaterandterrestrialhabitats.Conio-
scyphascus based on C. varius was originally proposed for
fungi with ConioscyphaasexualmorphsbyRéblová&Seifert
(2004).Conioscyphaexhibitsauniquemodeofconidiogenesis
with multiple, conspicuous collarettes forming a multilamel-
lar structure around the blastic conidiogenous locus of the
intercalaryconidiogenous cells(Shearer& Motta1973).Itis
characterised by inconspicuous perithecial ascomata that are
typically immersed to semi-immersed, hyaline, subhyaline to
pale orange with a papilla or long upright neck, coriaceous,
waxy ascomatal wall, cylindrical-clavate stipitate asci with a
pronouncednon-amyloidapical annulus,filiformparaphyses
andfusiformtofusiform-navicular,septate,hyalineascospores.
Ninespeciesareknownasapparentlyasexual(VonHöhnel
1904,Shearer1973,Matsushima1975,1993,1996,Kirk1984,
Udagawa&Toyazaki1983,Chen&Tzean2000,Crousetal.
2014)andonlytwohaveexperimentallyprovenlinkbetween
thesexualandasexualmorphs,i.e.C. varia(Réblová&Seifert
2004)andC. peruviana (Zelskietal. 2014).Athirdsexually
reproducing species, Conioscyphascus gracilis, was recently
transferred to Conioscypha(Zelskietal.2014).
Pleurotheciales
Six monophyletic clades that include species of eleven genera
were nested in the clade that we describe above as the Pleuro-
theciales(Fig.2).MembersofthePleurotheciales share dark,

76 Persoonia–Volume37,2016
papillate, glabrous or sparsely setose perithecia, upright or
lying horizontally to the host, asci with a distinct non-amyloid
apicalannulus,filiformparaphysesthatdisintegratepartiallyat
maturityandfusiformtoellipsoidal,septate,hyalineascospores.
Only ascospores of the sexual morph of Helicoön farinosum are
versicolorouswithbrownmiddlecellsandhyalinepolarcells.
The variation in the details of holoblastic conidiogenesis corre-
lateswithcladesrecoveredwithintheorder.Rhexolyticconidial
secession either on short denticles or rachis on sympodially
proliferating conidiogenous cells occurs in Helicoön farinosum,
Phaeoisaria, Melanotrigonum, Pleurothecium, Pleurotheciella
and Sterigmatobotrys.Thistypeofconidiogenesisischarac-
teristicof CladesI,IV,V,VIandpartially occursinCladesII
andIII. Schizolytic conidial secession on a single locus on
percurrently regenerating conidiogenous cells is characteristic
of Brachysporiella sensuEllis(Ellis1959).Thesametypeof
secession but on monoblastic or solitary thallic conidiogenous
cells is typical of Phragmocephala. Both latter genera are
positionedinCladeII.InTaeniolella, a sister of Sterigmatobo-
trysinCladeIII,thedarkbrownmacroconidiaareformedon
monoblastic conidiogenous cells in dry, acropetal chains, while
the apex of the conidium may develop into a fertile penicillate
head with sympodially elongating conidiogenous cells similar
to Sterigmatobotrys(seefurtherunderTaeniolella).
The nondescript morphology of sexual characters of members
of the Pleurotheciales makes their correct placement in the
Sordariomycetesdifficultand significantlyhinderstheir iden-
tificationandevendistinctionfromeachother.Withoutcultiva-
tion and/or molecular data their correct systematic placement
ischallenging.Thepresenceofconspicuousasexualmorphs
inintimatejuxtapositiontoascomataonthenaturalsubstratum
helpsidentificationofseveral genera only.Somespecies of
Pleurotheciella do not form conidiophores in vivo, only reduced,
hyalinetosubhyalineconidiophoresintheaxenicculture.Gene-
ra like Adelosphaeria and Plagiascoma, the latter is positioned
in the Bactrodesmiastrum clade outside the Pleurotheciales,
donotevenformtypicalasexualmorphs.Theyproducebrown,
ellipsoidal to globose, non-septate cells arising blastically from
vegetativehyphaeorothercellsintheaxenicculture.
MembersofChaetosphaeria (Chaetosphaeriales)aremorpho-
logically similar to Pleurothecium, Pleurotheciella and Sterig-
matobotrys of the Pleurotheciales, especially species with
Menispora asexualmorphs,e.g.C. ciliata, C. ovoidea, C. pul-
viscula or C. tortuosa (Holubová-Jechová1973,Réblováetal.
2006,Réblová&Seifert2008).Theypossessbrown,upright,
papillateascomata,fusiform,3-septate,hyalineascosporesin
cylindrical-clavate asci with distinct apical annulus and their
phialidicasexualmorphsareoftenabsentonthehost.Several
freshwater genera such as Aquaticola, Annulatascus and An-
nulusmagnus(Hoetal.1999,Hyde1992a,Campbell&Shearer
2004)canbecomparedwithAdelosphaeria, Melanotrigonum,
Pleurothecium, Pleurotheciella and Sterigmatobotrys based on
morphologyof ascomata,asci,ascospores andparaphyses.
Species of Aquaticola have miniature, coriaceous ascomata
lying horizontally to the host, asci with inconspicuous non-
amyloid apical annulus and septate or non-septate, hyaline as-
cospores(Hoetal.1999,Tsuietal.2003).Annulatascus and
Annulusmagnus are easily distinguished by asci with a conspic-
uous, non-amyloid apical annulus and relatively large, septate,
fusiform ascospores with a sheath or appendages in the former
taxon,arranged1-seriatelyorobliquely1-seriatelyintheascus.
Their asexual morphs are unknown and when isolated from
ascospores, sterile mycelium, or in the case of Annulusmag-
nus triseptatus abundantfertileascomata(M.Réblová,pers.
obs.) are formed in vitro. Phomatospora, whose taxonomic
placement in the Sordariomycetidaeisuncertain(Lumbsch&
Huhndorf2010),is another perithecial ascomycete that can
be compared with genera of the Pleurotheciales.Its species
are distinguishable by occurrence primarily on submerged
herbaceous stems, rarely on wood in freshwater and marine
habitats, immersed ascomata with thickened wall surrounding
the ostiolum and hyaline, longitudinally striate non-septate
ascospores enclosed in mucilaginous sheath or with bipolar
appendages(e.g.Hyde1988,1992b,Fallah&Shearer1998,
Fournier&Lechat 2010). OnlyPhomatospora berkeleyi, the
type species, and P. arenaria produce sporothrix-like asexual
morphs with holoblastic denticulate conidiogenesis in axenic
culture(Rappaz1992).
Savoryellales
The Savoryellales was placed in the Hypocreomycetidae based
onDNAsequences of six ribosomal and protein-coding loci
(Boonyuenetal.2011).Itformsawell-supportedlineagethat
includes saprobic, lignicolous species from terrestrial, marine,
brackish and freshwater environments and water-cooling tow-
ers(e.g.Jones&Eaton1969,Minoura&Muroi 1978,Hyde
&Jones1988,Changetal.1998,Ranghoo&Hyde1998).
AlthoughRanghoo(1998)introducedthefamilySavoryellaceae
as a member of the HalosphaerialesinherPhDThesis,avalid
descriptionwasneverpublished.Thefamilywasformallyintro-
duced recently as Savoryellaceae(Jaklitsch&Réblová2015).
As now delimited, the Savoryellales comprises three genera,
Ascotaiwania, Canalisporium and Savoryella.Ascotaiwania
is polyphyletic in our analyses, although the genus appeared
monophyleticinthreeprevious studies(Campbell &Shearer
2004,Hernández-Restrepoetal.2013,2015).Thelatterresults
were inadvertently distorted by the inclusion of species of Asco-
taiwania that only represent the CPS and Bactrodesmiastrum
cladesonasmallscale. In our multilocus phylogenies (Fig.
1,2)thecoreofAscotaiwania in the Savoryellales is centred
around the type species A. lignicola(Sivanesan&Chang1992)
and three other species. Helicoön farinosum(asA. hughesii,
Fallahetal.1999) is nested in the Pleurotheciales, while A. per-
soonii (Fallahetal.1999)isinastronglysupportedmonophy-
letic clade with Bactrodesmiastrum and Plagiascoma basal to
theCPSclade.
Genera of the Savoryellales share a similar morphology of dark,
minute perithecial ascomata with elongated, dark or subhyaline
neck,oftenoblique orlyinghorizontally onthehostwith the
neck facing upwards, asci with a non-amyloid apical annulus,
partlydeliquescingparaphysesandellipsoidaltofusiform,trans-
verselyseptate,versicolorousascospores.Thegenericdelimi-
tation of Ascotaiwania and Savoryella is narrow and for two
decades was based predominantly on ascospore septation, and
themorphologiesofparaphysesandtheascalapex,i.e.size
and shape of the apical annulus and presence or absence of
apicalthickening.TheascalapexofSavoryella was variously
interpreted in different studies, by authors studying different
species.Intheprotologueof the type species S. lignicola, the
ascal apex was described as apically thickened with a pore
(Jones&Eaton1969).Sivanesan&Chang(1992)separated
Ascotaiwania from Savoryella by an unthickened ascal apex
with a distinct apical annulus and ascospores with more than
three septa, while delimiting Savoryella for species lacking an
apicalring and having 3-septate ascospores. Later, several
otherspecieswereintroducedtothegenus,e.g.S. aquatica
(Hyde1993)andS. limnetica (Changetal.1998),characterised
byathickenedascalapexcontainingapicalannuluswithapore.
Readetal.(1993)basedtheirdistinctionofAscotaiwania and
Savoryella on ultrastructural observations and used the term
‘apicalapparatus’todescribethecomplexstructureoftheascal
apexofthesefungi.They characterised species of Ascotai-
wania by ascal apical apparatus comprising an annulus with a
protrusion(pendant)andpluggedpore,whereasinspeciesof

77
M.Réblováetal.:ThenewordersConioscyphales and Pleurotheciales
Savoryella the ascus apex was described as thickened with a
pore,butlackingapendant-likeprotrusion.Changetal.(1998)
alsousedcharactersofparaphysestodelimitthegenera,i.e.
narrow,filiform,earlydeliquescingfilamentsupto2µmwide
in Ascotaiwaniavsfilamentsconsistingofbroad,partiallydis-
integratingcellsupto8µmwideinSavoryella.Sri-indrasutdhi
etal.(2010)introducedanothermorphologicallysimilargenus,
Ascothailandia, as the sexual state of Canalisporium and dis-
tinguished it from Savoryella byitsconspicuousapicalannulus.
Recently,Boonyuenetal.(2011)modifiedthegenericconcept
of Savoryellaandacceptedspecieswith3-septateascospores
andcomparativelyflattenedapicalring.
The transfer of S. limnetica to Ascotaiwania proposed above
is based on molecular evidence and an experimentally proven
lifehistory.ThemicromorphologicalcharactersofS. limnetica,
i.e.flattenedapicalannulus,cylindrical,septate,disintegrating
paraphyses4.0–9.5µmwideand3-septateascospores,donot
fitwellwiththelong-heldmorphology-basedconceptsofeither
genus.StabledelimitationofAscotaiwania and Savoryella will
requirere-evaluationofallsexualandasexualmorphological
characters and concentrated sampling filtered through the
opticsofmultigenephylogenetics.
Asexual morphs associated with the Savoryellales were de-
scribed for Canalisporium grenadoideum (asAscothailandia
grenadoideasexual morph, Sri-indrasutdhi et al. 2010) and
three species of Ascotaiwania were linked with Brachysporiella-
like dematiaceous hyphomycetes, A. mitriformis and A. sawa-
dae(asMonotosporella, Ranghoo& Hyde 1998,Sivichaiet
al.1998)andA. limnetica (thisstudy).Withsomereservations
Acarocybiopsis was suggested as another suitable genus for
asexual morphs of Ascotaiwania (Réblová & Seifert 2004).
They are characterised by semi-macronematous conidiophores
often reduced to conidiogenous cells with a single locus and
brownmacroconidia. Conidia are either cheiroid, dictyosep-
tate with pores between cells and conidiogenous cells arise
from sporodochia in Canalisporium.Theasexualmorphs of
Ascotaiwania produce aleuroconidium-like, transversely sep-
tate macroconidia with darker bands around septa and a few
rhizoids arising from subtending cells beneath the monoblastic
conidiogenouscell.
Inouranalysis,thedematiaceoushyphomyceteTriadelphia uni-
septata nested within the monophyletic Ascotaiwania clade
as a sister to A. mitriformis. Triadelphia, based on T. hetero-
spora, was introduced for fungi from freshwater and brackish
environments and characterised by conidiophores reduced to
subglobose, subhyaline to dematiaceous conidiogenous cells,
schizolytic conidial secession and conidia produced blastically
fromasinglelocus(Shearer&Crane1971).Conidiaofspe-
cies of Triadelphia are brown or versicolorous often with one
or two polar cells paler than the middle ones, septate, usually
withdarkerbandsobscuringseveralsepta.AlthoughT. hetero-
spora was described with two morphologically distinct types
ofconidia,currentlyeighttypesareknown(Constantinescu&
Samson1982),but these otherasexualmorphshave never
beenformallynamed.Thegregarioustocaespitose,globoseto
subglobose to ampulliform conidiogenous cells borne directly on
vegetative hyphae are the hallmark of Triadelphia.Theyarealso
remarkably similar to cylindrical to lageniform aggregated conid-
iogenous cells of Bactrodesmiastrum.Themorphologyoflarger,
broad to ellipsoidal, brown, septate conidia of T. he terospora
andtheirconidiogenesisillustratedintheprotologue(Shearer
&Crane1971:f.9c,f,g)resemblesconidiaandconidiogenesis
of A. limnetica (Fig.10c –h)thatweobservedinaxenicculture
onPDA.Theampulliformconidiogenouscellsareabsentand
conidia arise directly from mycelium or a small monoblastic
conidiogenouscellwithseveral supporting cells. Triadelphia
comprises17species,butphylogeneticplacementofitstype
speciesisunknown.TheonlyavailableITSandnuc28SrDNA
sequencesin the GenBank belong to T. pulvinata and they
showaffinitywithmembersoftheMicroascales(Edathoduetal.
2013).ThepositionofT. uniseptata in the Savoryellales shown
here demonstrates that the present concept of Triadelphia is
polyphyletic, and that the application of this generic name, and
theredispositionofitsspecies,requiresmuchimprovedsam-
pling.
The ascoma centrum in the Hypocreomycetidae
InmembersoftheHypocreomycetidae, the centrum consists of
severaltypesofinterthecialfilaments.Theothertwosubclasses
of the Sordariomycetes, Sordariomycetidae and Xylariomy-
cetidae include either only paraphyses and periphyses in the
ostiolumorparaphysesarelackingin some groups.Apical,
lateral and centripetal paraphyses occur in members of the
Hypocreales(e.g.Samuels1973,Mhasker&Rao1976,Jak-
litsch2009,Jaklitsch&Voglmayr2014).Filamentsconsisting
of wide, inflated, early disintegrating cells interspersed among
the asci occur in the Bertiaceae and Chaetosphaerellaceae of
the Coronophorales (Réblová1999,Huhndorfetal.2004).A
hamatheciumconsistingof catenophyses,i.e.pseudoparen-
chymatous cells that break up to form chains of large, thin-
walled, early dissolving cells interspersed among asci or the
pseudoparenchyma may completely disappear in mature as-
comata, is typical of members of the Halosphaeriaceae of the
Microascales (Spataforaetal.1998,Sakayarojetal.2011).A
pseudoparenchymatous centrum occurs in the Melanosporales
(Goh&Hanlin1994,Samuels&Blackwell2001).Areticulate
network offiliform,branchingandanastomosingfilamentsat-
tachedatthetopandbottomofthecavityuniquelycharacterises
the Reticulascaceae of the Glomerellales while in members
of other two families, Australiascaceae and Glomerellaceae,
sparseseptatefilamentsoccur(Samuels&Müller1978,Sivane-
san&Alcorn2002,Réblováetal.2011).Numerousunbranched
filamentsattachedtothetopandbottomoftheascomatalcavity
occur in members of the Torpedosporales except for Marinoku-
lati chaetosa,wherethefilamentsareapicallyfree(Jonesetal.
2014,2015).Insomegroups,ahamatheciumislacking,e.g.
in the Scortechiniaceae and Nitschkia of the Coronophorales
(Huhndorfetal.2004)orinsomememberswithcleistothecial
ascomata of the Microascales.Thepresenceofperiphysesin
genera of the Coronophorales is variable and depends on how
theapexofascomataisformed,whetheritcontainsaQuellkor-
per(Nannfeldt1975)andwhetheritisostiolateornon-ostiolate
(Huhndorfetal.2004).
MembersoftheCPScladerepresenttheonlythreeordersin
the Hypocreomycetidaedefined by the presence of apically
freeparaphysesintheascomatalcentrum.Thesesterile,fili-
form,septatefilamentsemergefromthesubhymeniumeitherin-
terspersedamongtheasci,e.g.inAscotaiwania, Conio scypha,
Melanotrigonum, Pleurotheciella, Savoryella and Sterigmato-
botrys, orformseparatetuft-likestructures,e.g.Pleuro thecium.
Paraphyses are usually longer than the asci and may disinte-
grate at maturity; for example in some species of Savoryella
or Ascotaiwaniatheydisintegraterapidlyand are difficult to
observe.
Recently,theneworderPisorisporiales was introduced for pre-
dominantlyaquaticfungi,whichmorphologicallymimicmem-
bers of the Annulatascaceae in ascoma and ascospore char-
acters, and the Amphisphaeriaceae in a conspicuous, amyloid
apicalannulus and non-stromatic ascomata (Réblová et al.
2015).Theorderisisolatedonaseparatebranchasasister
to the Hypocreomycetidaebutwithoutstatisticalsupport.The
Pisorisporiales represents another group related to this sub-
classandcharacterisedbyfiliform,septate,partlydisintegrating
paraphyses interspersed among asci, but densely branching

78 Persoonia–Volume37,2016
andanastomosingabovetheirapicesintheascomacavity.
Although the two species of Pisorisporium, P. cymbiforme and
P. glaucum, were described from wood submerged in fresh-
water, several recent collections of P. cymbiforme were made
interrestrialhabitatsintheCzechRepublic,suggestingthatthe
fungusmightbewidespread.Thenewnuc18SrDNAandrpb2
sequencesofterrestrialstrainsarelistedinTable1.
Pleurotheciales: The polyphyletic genera Helicoön,
Phaeoisaria, Pleurothecium and Taeniolella
Helicoön
SeveralgeneranowclassifiedinthePleurotheciales appear
polyphyleticbasedonmolecularphylogenies.Helicoön farino-
sum, which has hyaline, coiled, septate conidia formed holo-
blastically on short denticles, is the only representative with
helicosporous conidia in the Pleurotheciales and in the whole
CPSclade. Itwasexperimentallylinkedwithitssexual state
Ascotaiwania hughesii(Fallahetal.1999)andinourphylogeny
itisnestedinCladeIasasistertoBrachysporiella setosa.We
confirmedthephylogeneticposition of H. farinosum (DAOM
241947) with collections, cultures and sequences made in
Canada(Réblováetal.2012).Althoughthecorrectspeciesepi-
thet for this holomorphic fungus would be ‘farinosum’,whether
the generic assignment should be Helicoön is unclear pending
confirmationofthephylogeneticplacementandclassificationof
the type species H. sessile.ThegenusHelicoön sensu Goos et
al.(1986)wasshowntobepolyphyleticwithDNAsequences
oftwonucrDNAlocibyTsui&Berbee(2006),butH. sessile
wasnotincluded.TheonlyavailableITSrDNAsequenceofthis
species(U72605,Pfisteretal.1997)shows99%similaritywith
theITSsequenceofSarocladium kiliense of the Hypocreales
(KP132606,Irinyietal.2015),anunlikelyrelationshipsugges-
tiveofamislabelled or contaminated culture.Otherspecies
of Helicoön were placed in the Pleosporales, Tubeufiales and
Venturiales of the Dothideomycetes(Tsui&Berbee2006).
Phaeoisaria
Phaeoisaria is a dematiaceous hyphomycete genus with spe-
cies producing indeterminate synnemata with septate or non-
septate ellipsoidal, obovoidal, fusiform-cylindrical or falcate
conidia formed on a sympodially extending rachis, occurring on
decayingwood,plantdebrisorsoilsediments(e.g.Sutton1973,
Deighton1974,Castañedaetal.2002,Seifertetal.2011,Mel’nik
2012,Chengetal.2014,Crousetal.2015).Thegenuswas
proposedbyVonHöhnel(1909)withtheonlyspeciesPh. bam-
busae.ItwasoriginallydescribedasanasexualstateofNeo-
peckia bambusae, inferredfromthe intimatejuxtapositionof
synnemataandascomata.Basedonhisrevisionoftypeand
herbariummaterial,Deighton(1974)consideredPh. bambusae
a synonym of Ph. clematidis.Hecompiledanextensivesyno-
nymy of the latter species, distinguishing it from morphologically
similar Ph. magnifica,whichhasbroaderconidia.Deighton’s
concept of Ph. clematidis seems to represent a complex of
severalphylogeneticspecies.
Phaeoisarianowincludes19species,fiveofwhichwereanalys-
edinourstudy.Thesampledspeciesformastronglysupported
monophyletic clade in the Pleurotheciales that includes species
withsynnemataandconidiophoresformedinfascicles.Inour
analysis, P. clematidis is represented by two strains isolated
from bark and senescent flower heads of Protea.
Phaeoisaria curvata is the only described mononematous spe-
cies; it was isolated from leaves of Parinari capense and its wild
typeisunknown(DeHoog&Papendorf1976).Thenuc28S
sequenceoftheex-typestrainCBS153.72(sequenceinthe
CBSstraindatabase)showsaffinitywithtaxaoftheSordario-
mycetidae.
AlthoughthemajorityofPhaeoisaria species are asexual, in-
cluding all species in our analyses, several perithecial asco-
mycetes have been linked with Phaeoisaria-like asexual
states.IntheSordariomycetes, Lentomitella and Rhamphoria
produce sparsely branched, mononematous conidiophores with
aseptateconidiaborneon ashortrachisinculture(Müller&
Samuels1982,Réblová2006).TwogeneraoftheDiatrypaceae,
Eutypella(asPeroneutypella,Deighton1974)andPareutypella
(Ju&Rogers1995),werelinkedwithPhaeoisaria-like synnema-
tousasexualstates. Fortheseconnections, themorphologi-
cally similar synnematous genus Harpographium,typified by
the asexual state of Eutypella scoparia,shouldbeconsidered.
Although Phaeoisaria is usually considered non-pathogenic to
humanbeings,twocasesofinflammationoftheeye’scornea
called keratitis were attributed to Phaeoisaria sp. (Chew et
al.2010)andPh. clematidis (Guarroetal.2000).Theformer
pathogenic strain Phaeoisaria sp.wasincludedinouranalysis
and is a sister taxon to two saprobic strains of Ph. clematidis
withstrongbranchsupport.
Pleurothecium
Pleurothecium includes fungi with dematiaceous, macronema-
tous, unbranched conidiophores and holoblastic, hyaline to sub-
hyaline, sympodially extending conidiogenous cells with a con-
spicuousrachisofdenticlesandhyaline,septateconidia.The
sexual morph is known only for the very common P. recurvatum,
thetypespecies(asCarpoligna pleurothecii,Fernándezetal.
1999).Oftheeightspeciesassignedtothegenus,onlythree
havebeenstudiedwith DNAsequencedata. Pleurothecium
recurvatum and P. semifecundum represent the core of the
genus and form a strongly supported monophyletic clade in the
Pleurotheciales, while P. obovoideum is nested within another
clade and sister to Brachysporiella setosa.Theasexualmorph
of P. semifecundum lacks macronematous conidiophores in
culture and sporulates sparsely; whether its wild type would
better match the distinctive conidiogenous apparatus of P. re-
curvatum remainsunknown.
Pleurothecium obovoideum, originally described in Ramichlo-
ridium, is known only from culture and it was isolated from a
dead leaf of Pasania edulis(Matsushima1975).Itischaracter-
ised by reduced, septate conidiophores, sympodially proliferat-
ingconidiogenouscellswithashortrachisgivingriseto2–3
denticles and ellipsoidal to obovate, pale brown, non-septate
conidiaformedsinglyorinshort chains. The morphology is
rather nondescript and we prefer to avoid introducing a new
genus for this species, until either the wild type is collected or
relationshipwithothermorphologicallysimilartaxaisrevealed.
Based on its morphology, P. obovoideum is similar to Rhinocla-
diella mackenziei(Chaetothyriales),apathogencausingsevere
cerebralphaeohyphomycosisinhumans(Suttonetal.1998).
ItalsoresemblesmembersofSubramaniomyces(Xylariales,
Crouset al.2007)andPterygosporopsis (Kirk1983),whose
phylogeneticplacementisunknown.
Taeniolella
Taeniolella exilis, the type of the genus, is commonly found on
decaying wood and bark of Betula (Hughes1958,Ellis1971).
DuringarevisionofthetypematerialofT. exilisbyJonesetal.
(2002),apenicillatelybranchedconidiophorewasobservedas
anextensionoftheterminalmacroconidia.Asimilarpenicillate
conidiophore was observed in two other species, T. longis-
sima and T. rudis(Hughes1980,Jonesetal.2002).Thelatter
taxon was shown to be closely related to Sterigmatobotrys
macrocarpa of the Pleurotheciales, whose asexual state is
characterised by similar penicillate conidiophores with several
series of branches and metulae terminating macronematous
conidiophores(Réblová&Seifert2011).However,brown,sep-

79
M.Réblováetal.:ThenewordersConioscyphales and Pleurotheciales
tate Taeniolella macroconidia were not observed in axenic
cultures obtained from conidia or ascospores of S. macrocarpa.
Several other species of Taeniolella are positioned in distantly
relatedgroups.Taeniolella-like conidiawere obtained in a
culture derived from ascospores of the freshwater ascomycete
Chaetorostrum quincemilensis, tentatively placed in the Annu-
latascaceae(Zelskietal.2011).Sheareretal.(2009)showed
the strain of T. alta (CBS488.80)nestedinacladewithDia-
porthe angelicae and Phomopsis sp.,andtheex-typestrainof
T. typhoides (CCMF-10198)intheLingdomycetaceae of the
Pleosporales.Ataeniolella-like fungus was isolated from the
rhizosphere soil of strawberry, producing a phialophora-like
asexual state on vegetative hyphae or directly on macroconidia
in vitro, and described as T. phialosperma (Watanabe1989,
1992).TheITSsequencesoftwononex-typestrainsofT. phia-
losperma depositedinGenBank(KF703925,GU966521,un-
publ.)indicatearelationshipwithmembersoftheSordariales.
Finally, the sexual morph of a Taeniolellasp.withascolocular
ascomadevelopmentwasclassifiedasMytilinidion gemmige-
num(Mytilinidiales,Minter&Holubová-Jechová1981).These
inconsistencies suggest that the generic concept of Taeniolella
requiresincreasedtaxonsamplingandinvestigationwithmo-
lecularmethods.
A case of human subcutaneous phaeohyphomycosis caused
by T. exilisspecieswasreportedbyAlonsoetal.(1993).While
the pathogenic strain of T. exilis isolated from a human skin
lesion(strainIP2199.93)wasshowncloselyrelatedtoOchro-
cladosporium elatum (CBS 146.33) of the Pleosporales by
Masclauxetal.(1995),theplacementofthewood-inhabiting
strain of T. exilis resembling T. rudis(Pleurotheciales)hasyet
tobeconfirmedwithmolecularsequencedata.
AcknowledgementsThisstudywassupportedbytheProjectoftheNatio-
nalFoundationoftheCzechRepublic(GA506/12 /0038),andasalong-term
researchdevelopmentprojectoftheInstituteofBotany,AcademyofSciences
No.RVO67985939,andof theInstitute ofMicrobiology,AcademyofSci-
encesNo.RVO61388971.WethankWalterGamsforhelpfulsuggestions
onnamesofthenewtaxaproposedinourstudy.
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