1
Pleu oph agmium pa ispo um (Ascomyco a): One name, se en
s o ies – a case highligh ing he need o e i ica ion o s ains
om public cul u e collec ions
Ma ina Réblo á1, Jana Nek indo á2, Lucie Baucho á1, Ma ga i a He nández-Res epo3
1 CzechAcademyo Sciences,Ins i u eo Bo any,Depa men o Taxonomy,25243P ůhonice,CzechRepublic
2 Ins i u eo ClinicalBiochemis yandDiagnos ics,Uni e si yHospi alH adecK álo é,50005H adecK álo é,CzechRepublic
3 Wes e dijkFungalBiodi e si yIns i u e,3584CTU ech ,Ne he lands
Co espondingau ho :Ma inaRéblo á([email p o ec ed])
Copy igh : © Ma ina Réblo á e al.
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
Public eposi o ies o li ing ungal s ains p o ide essen ial e e ence poin s and suppo
di e se scien i ic ou comes. Cu en bes p ac ices o p ese ing ungal s ains empha-
sise he gene a ion o DNA ba codes and he managemen o comp ehensi e me ada a.
Howe e , challenges a ise when ype ma e ial o au hen ic e e ence s ains a e lacking, as
his p e en s di ec compa ison o DNA ba codes and o ces iden i ica ions o ely solely
on mo phology. This p oblem is pa icula ly p onounced o s ains deposi ed du ing he
p e-molecula e a, especially hose belonging o species wi h simple o con e gen mo phol-
ogies. In his s udy, we e-examined se en s ains deposi ed in a public cul u e collec ion un-
de he name Pleu oph agmium pa ispo um, including synonymous designa ions. Ou ap-
p oach combined cul i a ion expe imen s, compa a i e mo phological analyses, mul i-locus
phylogene ic econs uc ion o six nuclea ma ke s, and biogeog aphic assessmen s. Ou
analyses e ealed ha hese s ains a e sca e ed ac oss ou dis inc amilies o o de s in
h ee classes. Two s ains belong o Thysano ea (Chae o hy iales, Eu o iomyce es): T. ac o-
pleu ogena sp. no . and a s e ile s ain iden i ied as he al eady known T. melanica. Two o h-
e s ains we e esol ed wi hin Wongia (Papulosaceae ince ae sedis, So da iomyce es) and
in oduced as W. pallidopola is sp. no . and W. hachidopho a sp. no . Finally, wo s ains
ep esen no el axa wi hin he Tubeu iales (Do hideomyce es), desc ibed he e as Zaaneno-
myces hili e sp. no . and Skoliomycella la a gen. e sp. no . O he se en examined s ains,
only one con o med o he species concep o P. pa ispo um and is he e ega ded as i s e -
e ence s ain. The phylogene ic analyses esol ed P. pa ispo um wi hin Neomy mec idium
(My mec idiales, So da iomyce es). Consequen ly, Neomy mec idium was educed o syn-
onymy o Pleu oph agmium, leading o he p oposal o 11 new combina ions (P. asia icum
comb. no ., P. asymme icum comb. no ., P. usi o me comb. no ., P. gaoligongense comb.
no ., P. guizhouense comb. no ., P. luguense comb. no ., P. na icula e comb. no ., P. p e ido-
phy ophilum comb. no ., P. sep a um comb. no ., P. sichuanense comb. no ., and P. so bicola
comb. no .), and wo new names (P. lu iale nom. no . and P. jiulongheense nom. no .). In
addi ion, h ee species o me ly placed in Uncispo a a e ans e ed o Thysano ea, wi h new
combina ions p oposed based on cong uen mo phology and mul i-locus phylogene ic e i-
dence: T. hainanensis comb. no ., T. sinensis comb. no ., and T. wuzhishanensis comb. no .
This s udy e ines he gene ic limi s o Pleu oph agmium and mo phologically simila gene a
and e eals se e al p e iously un ecognised lineages. I highligh s how misin e p e a ion o
sub le mo phological ea u es may lead o s ains being misiden i ied and deposi ed unde
inco ec names in public collec ions, whe e hey isk pe pe ua ing axonomic e o s.
Academic edi o :
Sajeewa Maha achchikumbu a
Recei ed:
24 Sep embe 2025
Accep ed:
13 No embe 2025
Published:
25 No embe 2025
Ci a ion: Réblo á M, Nek indo á J,
Baucho á L, He nández-Res epo M
(2025) Pleu oph agmium pa ispo um
(Ascomyco a): One name, se en
s o ies – a case highligh ing he need
o e i ica ion o s ains om public
cul u e collec ions. IMA Fungus 16:
e173033. h ps://doi.o g/10.3897/
ima ungus.16.173033
IMA Fungus 16: e173033 (2025)
DOI: 10.3897/ima ungus.16.173033
2
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Key wo ds: Ascomyco a, Dac yla ia, DNA ba code, holoblas ic-den icula e conidiogenesis,
new axa, phylogene ics, eposi o y, sap obic, sys ema ics
In oduc ion
Public cul u e collec ions play a c i ical ole in mycological esea ch by
se ing as sou ces o e e ence ma e ial o axonomy, e olu iona y s ud-
ies, ecology, bio echnology and pa hology. They p o ide essen ial e e ence
poin s ha o e eliable in o ma ion ha is c i ical o da a-d i en esea ch.
As Hawkswo h (2004) emphasised, such collec ions a e mo e accu a ely
e med gene ic esou ce collec ions, e lec ing hei unc ion as eposi o-
ies o he i able biological di e si y.
Fungi a e one o he mos species- ich ye unde s udied g oups o o ganisms
on Ea h. Recen es ima es place global ungal di e si y be ween 1.5 o 3.2
million species, wi h a wo king mean o app oxima ely 2.53 million (Niskanen
e al. 2023). In con as , only abou 204 040 species ha e been scien i ically
documen ed, ep esen ing oughly 8% o he es ima ed mean (Index Fungo-
um, accessed on 10 Sep embe 2025). The disc epancy be ween known and
es ima ed di e si y is e en mo e p onounced in ex si u conse a ion e o s,
such as cul u e collec ions, c yobanks, and ge mplasm s o ages. Some public
collec ions a e skewed owa d a limi ed numbe o economically o medically
impo an axa, while as lineages, especially o non-lichenised, mic o ungal
lineages, emain poo ly ep esen ed o absen . Lo g en and S ajich (2021) high-
ligh ed he need o add ess he ‘in isible’ ungal biodi e si y c isis and posi-
ioned cul u e collec ions as key ools in ungal conse a ion and sys ema ics.
The Wo ld Da a Cen e o Mic oo ganisms (gcm.wdcm.o g, accessed on 10
Sep embe 2025) cu en ly lis s 588 984 s ains o mic oo ganisms, including
ungi, yeas s, and bac e ia, ep esen ing 57 219 species p ese ed ac oss 158
cul u e collec ions wo ldwide. The p opo ion o ungal species main ained in
cul u e collec ions is widely ecognised o be low. Ea lie es ima es sugges
ha ewe han 20 000 ungal species a e p ese ed ex si u (Hawkswo h
2004). Among exis ing eposi o ies, he Wes e dijk Fungal Biodi e si y Ins i u e
in U ech , he Ne he lands, main ains he mos comp ehensi e and globally
ecognised collec ion o e 130 000 s ains o mic oo ganisms, o which mo e
han 100 000 a e ungi, including yeas s, co esponding o app oxima ely 25
000 species. This ex ensi e a chi e se es as he p incipal eposi o y o no-
mencla u al ypes and au hen ic e e ence s ains. I p o ides essen ial ma e i-
al o axonomic, ecological, and bio echnological esea ch.
The e ec i eness and eliabili y o he cul u e collec ions depend hea ily on
accu a e species iden i ica ion and comp ehensi e me ada a. These collec-
ions a e commonly used unde he assump ion ha deposi ed s ains a e co -
ec ly iden i ied, especially when hey a e assigned o known species. Be o e
molecula ools became a ailable, s ain iden i ica ion elied mainly on mo -
phology. Ve i ica ion p ac ices a ied be ween ins i u ions, and many names
we e accep ed wi hou e-examina ion. Wi h he inc easing accessibili y o
DNA sequencing, howe e , ou ine gene a ion o DNA ba codes o newly de-
posi ed s ains has become easible and is now being implemen ed in a ious
3
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
collec ions (Vu e al. 2016, 2019). In pa allel, ad ances in da a managemen
sys ems (Reime and Yu ko 2022) a e imp o ing he eliabili y and aceabili-
y o e e ence ma e ials. Mo eo e , comp ehensi e monog aphic s udies a e
being conduc ed o achie e be e species esolu ion, pa icula ly in gene a o
medical, indus ial, o ag icul u al impo ance, such as Ac emonium, Al e na ia,
Aspe gillus, Ce a ocys is, Fusa ium, Penicillium, Phoma, and o he s (de G uy e
e al. 2013; Woudenbe g e al. 2013, 2015; de Bee e al. 2014; Chen e al. 2017;
Houb aken e al. 2020; Sando al-Denis e al. 2025).
Howe e , e-iden i ica ion o his o ical s ains in public collec ions, especial-
ly sap obic ungi, is s ill no ou inely pe o med. Some s ains deposi ed be o e
he molecula e a emain in collec ions unde un e i ied names, and hese may
conceal conside able phylogene ic di e si y, as demons a ed in he p esen
s udy. Misiden i ica ions, especially o mo phologically de ined isola es, can
ha e cascading e ec s in sys ema ics, phylogene ics, bio echnology, and ecol-
ogy. In esponse, he mycological communi y ad oca es o s anda diza ion,
molecula e i ica ion, and p ope ouche deposi ion as essen ial pilla s o
main aining he in eg i y and ep oducibili y o ungal science (Hawkswo h
2004; Aime e al. 2021; Lo g en and S ajich 2021).
Ne e heless, e en he implemen a ion o ou ine DNA-based au hen ica ion
may p o e insu icien i e e ence o ex- ype s ains o he a ge species a e
una ailable, which is o en he case. Unde hese ci cums ances, he adi ional
app oach o ca e ully alida ing he mo phology o ob ained s ains, guided by
he p inciple o ‘ us , bu e i y’, emains essen ial o ensu ing he eliabili y
and success o any axonomic o sys ema ic esea ch.
In his s udy, we p esen speci ic cases encoun e ed du ing ou own esea ch,
which illus a e how un e i ied iden i ica ions o s ains in public collec ions
can obscu e ue phylogene ic ela ionships and hinde axonomic cla i y. A no-
able example is p o ided by he s udies o Shenoy e al. (2006, 2010), whe e
eliance on misiden i ied li ing s ains, wi hou mo phological e i ica ion by
he au ho s, igge ed a cascade o e oneous conclusions, ul ima ely a ec ing
he ou comes o se e al subsequen sys ema ic in es iga ions. Using pa ial
nuclea la ge subuni (LSU) DNA sequences, Shenoy e al. (2010) p oposed
ha he genus Spadicoides was polyphyle ic. The genus, ypi ied by S. bina, was
es ablished by Hughes (1958) o accommoda e sap obic, p ima ily lignicolous,
dema iaceous hyphomyce es. To suppo hei hypo hesis o polyphyly, She-
noy e al. (2010) analysed ou a ailable s ains: Spadicoides a a CBS 489.77,
S. bina CBS 113708, S. e ucosa ex- ype CBS 128.86, and S. xylogena CBS
310.31. Al hough S. a a (GenBank accession: EF204506) and S. bina (GenBank
accession: EF204507) a e mo phologically simila , phylogene ic analysis did
no suppo hei congene ic placemen . The s ain o S. a a o med a sis e
ela ionship o h ee species o Len omi ella (Réblo á e al. 2018), whe eas S.
bina was placed wi hin he Po osphae ella clade (Mülle and Samuels 1982).
He nández-Res epo e al. (2017) accep ed he conclusions o Shenoy e al.
(2010), ecognising Spadicoides as a membe o he Co danaceae, whe e Po o-
sphae ella and i s asexual mo ph Co dana belong, and seg ega ing S. a a om
Spadicoides in o a newly es ablished genus, Xenospadicoides. In addi ion, a
new o de , Xenospadicoidales, was in oduced o accommoda e Len omi ella
and o he spadicoides-like axa. Using eshly collec ed ma e ial, Réblo á e al.
(2018) ob ained S. bina CBS 137794 in axenic cul u e by isola ing ascospo es o
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IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
an uniden i ied len omi ella-like species. Thei mul i-locus phylogene ic analy-
ses con i med ha S. bina, S. a a, and o he Spadicoides species a e congene -
ic. Based on his e idence, he genus Spadicoides was accep ed as a membe
o he Xenospadicoidales, and Xenospadicoides was educed o i s synonymy.
Howe e , he iden i y o he s ain S. bina CBS 113708, p e iously placed wi hin
he Co danaceae by Shenoy e al. (2010), emained un esol ed.
Spadicoides bina and Co dana paucisep a a (P euss 1851) a e mo phologi-
cally s ikingly simila , bo h p oduce 1-sep a e, b own, ellipsoidal, ac opleu og-
enous conidia o compa able size. Howe e , hey di e undamen ally in hei
mode o conidiogenesis. In Spadicoides, conidiogenous cells a e e ic, whe e-
as in Co dana, conidia a e bo ne on den icles on holoblas ic conidiogenous
cells om e minal and in e cala y (nodal) swellings. De ailed examina ion and
compa ison o he o iginal ouche specimen housed in he Uppsala he ba -
ium, he o iginal d ied cul u e and he de i ed li ing s ain CBS 113708, bo h
deposi ed in he CBS cul u e collec ion, con i med ha he ungus in ques ion
is, in ac , C. paucisep a a (Réblo á e al. 2018). I is e iden ha he s ain was
ini ially misiden i ied, ye he deposi ion o all ela ed ma e ial in public collec-
ions ul ima ely enabled accu a e iden i ica ion.
Ano he example conce ns he misiden i ica ion o a s ain o Bahusu abee-
ja dwaya (MTCC 9680), he ype species o he genus, by Shenoy e al. (2010). I
was placed wi hin he Bo yosphae iales (Do hideomyce es). The au ho s we e
unable o accoun o his placemen mo phologically and discussed he ap-
pa en dispa i y be ween he phylogene ic and mo phological da a. Based on
he ex- ype s ain, he species is cu en ly assigned o Codinaea wi hin he Cha-
e osphae iales (Réblo á e al. 2021). A simila case in ol es he misiden i ica-
ion o a non- ype s ain o Linkosia mul isep a um (HKUCC 10825), epo ed by
Shenoy e al. (2006) as a membe o he Rhampho iales. Mo e ecen molecula
da a om he ex- ype s ain o L. mul isep a um (CGMCC 3.20786; Wu and Diao
2022) ha e placed Linkosia wi hin he Chae osphae iales, oge he wi h o he
mo phologically simila spo idesmium-like ungi.
This s udy is ocused on he case o Pleu oph agmium pa ispo um and
e-e alua es he s ains deposi ed unde his name o cla i y he species’ iden-
i y and assess he b oade implica ions o s ain misiden i ica ion in public
collec ions. Pleu oph agmium (Cos an in 1888), ypi ied by P. bicolo (= P. pa -
ispo um, Holubo á-Jecho á 1972), was in oduced o dema iaceous hypho-
myce es. I is cha ac e ised by ans e sely sep a e, hyaline conidia ha be-
come subhyaline o pale b own a ma u i y, bo ne e minally and la e ally on
den icles on holoblas ic conidiogenous cells, a ising om simple, mac onema-
ous conidiopho es. De Hoog and on A x (1974) ans e ed Pleu oph agmium
o Dac yla ia (Sacca do 1880) based on o e lapping conidial mo phology, and
de Hoog (1985) la e accommoda ed i as Dac yla ia sec . Pleu oph agmium.
The concep o his species was p ima ily de i ed om obse a ions o wo
s ains, CBS 531.73 and CBS 770.83, which we e used o desc ibe bo h spe-
cies-le el ai s and colony mo phology (de Hoog 1985).
Pleu oph agmium pa ispo um is ep esen ed by se en s ains in he
CBS cul u e collec ion. The only a ailable molecula da a o P. pa ispo um
consis o a pa ial LSU sequence om he non- ype s ain CBS 531.73
(GenBank accession: EU107296; Bhilabu a e al., unpublished), which was
gene a ed wi hou p io mo phological e i ica ion. In a phylogene ic analysis
5
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
by Réblo á (2009), he placemen o Rhodo e onaea and mo phologically
simila gene a wi h holoblas ic-den icula e conidiogenesis was examined
using homologous LSU sequences ep esen ing species om 21 o de s
and amilies ac oss h ee ungal classes. Pleu oph agmium pa ispo um
CBS 531.73 g ouped wi h Papulosa ame ospo a AFTOL-ID 748 (Spa a o a
e al. 2006) in he So da iomyce es. Based on hese esul s, Réblo á (2009)
excluded Pleu oph agmium om Dac yla ia and en a i ely e-e alua ed i s
placemen wi hin he Papulosaceae. Howe e , he au ho no ed ha his
decision was made in he absence o e i ied o iginal ma e ial.
In addi ion o Dac yla ia, Pleu oph agmium also shows s iking mo pholog-
ical simila i y o Neomy mec idium (C ous e al. 2018a). Neomy mec idium,
based on N. sep a um, cu en ly encompassing 13 species, is cha ac e ised
by dema iaceous, mac onema ous, e ec , unb anched conidiopho es, holoblas-
ic-den icula e conidiogenous cells, and soli a y, usoid-ellipsoidal, obo oid o
na icula e, ans e sely sep a e conidia ha a e ini ially hyaline and become
pale b own wi h age, and ha e a mucoid shea h. Ne e heless, he close mo -
phological simila i y be ween he wo gene a has la gely been o e looked in
s udies p oposing new Neomy mec idium species (e.g. C ous e al. 2018a, b;
Se ano e al. 2020; Xu e al. 2023; Zhang e al. 2024, 2025; Cao e al. 2025).
Du ing a e ision o he genus Pleu oph agmium, we aimed o cla i y he sys-
ema ic posi ion o P. pa ispo um, a mo phologically dis inc i e ye poo ly un-
de s ood axon. Al hough i has been epo ed om empe a e egions o Asia
(Ma sushima 1975), No h Ame ica (Wang 2010), and Eu ope (Ellis 1968, 1971;
Holubo á-Jecho á 1972), we we e unable o ecollec i in na u e. Mo eo e , ou
ecen examina ion o he s ain CBS 531.73 aised suspicion ha i may be mis-
iden i ied. In esponse, we acqui ed se en s ains o his species a ailable in he
CBS cul u e collec ion. These isola es we e iden i ied based on mo phological
cha ac e is ics a he ime o hei deposi ion. Al hough ITS ba codes a e now
a ailable o hese s ains, he absence o ex- ype o e i ied s ain o P. pa ispo-
um makes accu a e iden i ica ion impossible wi hou manual alida ion.
In his s udy, we p esen a comp ehensi e in es iga ion o P. pa ispo um,
wi h pa icula emphasis on cla i ying he iden i y o se en s ains ob ained
om he CBS cul u e collec ion. Ou app oach combined cul i a ion expe i-
men s, compa a i e mo phological analyses, mul i-locus phylogene ic econ-
s uc ion using six nuclea ma ke s and biogeog aphic assessmen s. The e-
sul s e ealed unexpec ed phylogene ic di e si y among hese s ains, which
we e shown o ep esen no el and p e iously known species, dis ibu ed
ac oss se e al gene a, spanning h ee ungal classes.
Ma e ials and me hods
Isola es and mo phological s udies
Li ing s ains we e sou ced om he Wes e dijk Fungal Biodi e si y Ins i u e
(CBS) a U ech , he Ne he lands. S ains CBS 122759, CBS 440.70, and CBS
862.68 we e o iginally deposi ed as Pleu oph agmium simplex (a synonym o
P. pa ispo um) and la e eassigned o Dac yla ia pa ispo a o Dac yla ia sp.
S ain CBS 531.73 was deposi ed as P. pa ispo um and subsequen ly placed
unde D. pa ispo a, while s ains CBS 215.96, CBS 770.83, and CBS 113561 we e
6
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
deposi ed di ec ly as D. pa ispo a. D ied cul u es we e deposi ed in he CBS Fun-
ga ium, while addi ional he ba ium specimens o P. pa ispo um we e deposi ed
in he He ba ium o he Ins i u e o Bo any (PRA), Czech Academy o Sciences a
P ůhonice, Czech Republic. Biogeog aphic assessmen s we e conduc ed using
en i onmen al ITS da ase s e ie ed om he GlobalFungi da abase (Vě o ský
e al. 2020). Newly desc ibed axa we e egis e ed in MycoBank (C ous e al.
2004). De ails o he s ains examined, including hei o igin and GenBank acces-
sion numbe s o he gene a ed sequences, a e summa ised in Table 1.
Colony mac omo phology was examined using an Olympus SZX12 dis-
sec ing mic oscope (Olympus Ame ica, Inc., Mel ille, NY, USA). Mic oscopic
p epa a ions we e moun ed in 90% lac ic acid, wa e , o Melze ’s eagen , wi h
measu emen s aken om specimens in Melze ’s eagen . Conidial dimensions
a e p esen ed as mean ± s anda d de ia ion (SD) based on 20–25 measu e-
men s. Mic omo phological ai s we e s udied wi h an Olympus BX51 ligh
mic oscope, and mic og aphs we e cap u ed using an Olympus DP75 came a
ope a ed ia Olympus cellSens Dimension so wa e . 4.3. Colony images we e
aken wi h a Canon EOS 77D digi al came a equipped wi h a Canon EF 100 mm
/2.8L Mac o IS USM lens (Canon Eu ope L d., Middlesex, UK), illumina ed wi h
5500K 16W LED ligh s. All images we e p ocessed in Adobe Pho oshop CS6
(Adobe Sys ems, San Jose, CA, USA).
To examine colony mo phology, pigmen p oduc ion, and g ow h a es, s ains
we e cul i a ed on ou media: co nmeal dex ose aga (CMD) (co nmeal aga ,
Oxoid Limi ed, Basings oke, UK, supplemen ed wi h 2% w/ dex ose), mal ex-
ac aga (MEA) (Oxoid), Modi ied Leonian’s aga (MLA) (Malloch 1981), oa -
meal aga (OA), and po a o-ca o aga (PCA) (C ous e al. 2019). To p omo e
spo ula ion, s ains we e also g own on co nmeal aga (CMA) (C ous e al.
2019) supplemen ed wi h d ied s ems o U ica dioica and incuba ed unde al-
e na ing 12-h cycles o UV ligh and da kness. Colony ea u es we e eco ded
om 4-wk-old cul u es incuba ed a 23 °C in da kness.
Scien i ic names o ungal gene a epo ed in his s udy a e abb e ia ed as
ollows: Bo yosphae ia (B.), Campo esiomyces (Ca.), Cancellidium (Cn.) Co dana
(C.), Cyphellopho a (Cy.), Helicoma (H.), Minimelanolocus (M.), Neomy mec idium
(N.), Pleu oph agmium (P. ), Pseudospi opes (Ps.), Skoliomycella (S.), Thysano ea
(T.), Tubeu ia (Tu.), Uncispo a (U.), Wongia (W.), and Zaanenomyces (Z.).
Molecula me hods
Phylogene ic ela ionships we e e alua ed using six gene ma ke s: in e nal an-
sc ibed space ITS1–5.8S–ITS2 (ITS) o he nuclea DNA cis on (ITS ba code)
(Schoch e al. 2012), nuclea la ge subuni (LSU) DNA gene, he nuclea small
subuni (SSU) DNA gene, and h ee coding ma ke s, i.e. he second la ges sub-
uni o RNA polyme ase II (DNA-di ec ed RNA polyme ase) ( pb2), he in e media e
sec ion o he ansla ion elonga ion ac o 1-α ( e 1), and β- ubulin ( ub2) ma ked
by exons 3−6. The LSU and SSU ma ke s p o ide eliable phylogene ic esolu ion
a he gene ic and highe axonomic le els in ungi (e.g. Zhang e al. 2007; Schoch
e al. 2009). The coding gene ma ke s a e ecognised o hei e ec i eness in
esol ing in e speci ic ela ionships (S ielow e al. 2015; Meye e al. 2019).
7
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
P o ocols o DNA ex ac ion om 2–4-d old cul u es and PCR ampli ica-
ion o ci ed gene ma ke s we e conduc ed ollowing he me hods desc ibed
by Réblo á e al. (2022) and Réblo á and Nek indo á (2023). Au oma ed
sequencing was ca ied ou by Eu o ins Genomics Eu ope Sequencing Se -
ice (Cologne, Ge many). Analyses o aw sequence da a and assembly o
sequence con igs we e pe o med using Sequenche . 5.4.6 (Gene Codes
Co p., Ann A bo , MI, USA).
Phylogene ic analyses
P elimina y simila i y sea ches we e pe o med using he BLASTn and
megaBLAST algo i hms wi h ITS and LSU sequences o all s udied s ains o
iden i y hei closes ela i es. Co esponding ITS, LSU, SSU, pb2, e 1, and
ub2 sequences o ela ed axa we e hen e ie ed om GenBank sequence
da abase a NCBI (Saye s e al. 2022) and used in subsequen analyses. Se-
quences o hese species, oge he wi h hei GenBank accession numbe s
and e e ences, a e lis ed in Table 2.
Sequence alignmen s we e gene a ed wi h MAFFT . 7.487 (Ka oh and
S andley 2013) ia he CIPRES Science Ga eway . 3.3 (Mille e al. 2010)
and adjus ed manually in BioEdi . 7.1.8 (Hall 1999) when necessa y. The
bes nucleo ide subs i u ion models o each pa i ion (ITS, LSU, SSU, pb2,
e 1, ub2) we e selec ed unde he Akaike In o ma ion C i e ion using M -
Model es . 2.4 (Nylande 2004). Phylogene ic econs uc ions we e pe -
o med wi h Maximum Likelihood (ML) and Bayesian In e ence (BI) me h-
ods implemen ed in he CIPRES Science Ga eway.
Table 1. Species, isola e in o ma ion and new sequences de e mined o his s udy (in bold) and addi ional sequences
e ie ed om GenBank.
O ganism S ain S a us* Hos Subs a e Coun y GenBank accessions Re e ence
ITS LSU SSU pb2 e 1 ub2
Pleu oph agmium
simplex
CBS
770.83
uniden i ied dead
wigs
Japan PX283736 PX283746 PX283743 PX310214 PX310206 –This s udy
Skoliomycella
la a
CBS
122759
Tuniden i ied plan
deb is
Po ugal PX283737 PX283747 –PX310215 PX310207 –This s udy
Thysano ea
ac opleu ogena
CBS
215.96
Tuniden i ied wood Papua New
Guinea
PX283738 PX283748 – – – PX310220 This s udy
Thysano ea
melanica
CBS
862.68
n/a whea
ield soil
Ne he lands PX283739 PX283749 – – – PX310221 This s udy
Wongia
pallidopola is
CBS
440.70
Tn/a sandy soil Ne he lands PX283740 PX283750 PX283744 PX310216 PX310208 –This s udy
Wongia
hachidopho a
CBS
531.73
TBambusa
sp.
dead lea India PX283741 PX283751 PX283745 PX310217 PX310209 –This s udy
Zaanenomyces
hili e
CBS
113561
TCa ex sp. li e I an PX283742 PX283752 –PX310218 PX310210 –This s udy
Zaanenomyces
mode a icis-
academiae
CBS
148315
TJuncus
in lexus
dead culm Ne he lands OK664723 OK663762 –OK651167 PX310211 –C ous e
al. 2021
Zaanenomyces
quad ipa is
CBS
148310
TJuncus
e usus
dead culm Ne he lands OK664721 OK663760 –PX310219 PX310212 –C ous e
al. 2021
Zaanenomyces
e sa ilis
CBS
148312
TJuncus
in lexus
dead culm Ne he lands OK664730 OK663769 – – PX310213 –C ous e
al. 2021
*T deno es ex- ype cul u e.
8
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Table 2. Species, isola e in o ma ion and sequences e ie ed om GenBank.
Taxon S ain S a us GenBank Accession numbe s Re e ence
ITS LSU SSU pb2 e 1 ub2
Acan hohelicospo a
pinicola
MFLUCC 10-0116 T KF301526 KF301534 – – KF301555 – Boonmee e al. 2014
Acan hos igma
chiangmaiense
MFLUCC 10-0125 T JN865209 JN865197 – – KF301560 – Boonmee e al. 2011,
2014
Acan hos igma
pe pusillum
UAMH 7237 AY916492 AY856892 – – – – Tsui e al. 2006
Aciculomyces es ic us FMR 18994 T ON009870 ON009950 – – – ON667802 To es-Ga cia e al. 2023
Aculea a aqua ica MFLUCC 11-0529 T MG922571 MG922575 – – – – Dong e al. 2018
A okylind iopsis se ulosa HMAS 245592 T KP337330 KP337329 – – – – Ma e al. 2015
Be kleasmium aqua icum MFLUCC 17-0049 T KY790444 KY790432 –MF535268 KY792608 – Lu e al. 2017c, 2018b
Be kleasmium usi o me MFLUCC 17-1978 T MH558693 MH558820 –MH551007 MH550884 – Lu e al. 2018a
Be lesiella nige ima MUCL 39954 AF050251 AF050251 – – – – Un e eine and Na eau
1999
Boe lagiomyces
mac ospo us
MFLUCC 12-0388 KU144927 KU764712 – – KU872750 – Doilom e al. 2017
Bo yosphae ia aga es MFLUCC 10-0051 JX646790 JX646807 – – JX646855 – Liu e al. 2012
Bo yosphae ia do hidea CBS 115476 EKF766151 DQ678051 –DQ677944 DQ767637 – Schoch e al. 2006;
Slippe s e al. 2013
Bo yosphae ia
wangensis
HGUP 190007 MZ541933 MZ540051 –OP321271 – – Zhang e al. 2021, 2022
B unneospo ella aqua ica HKUCC 3708 AF177154 AF132326 – – – Ranghoo e al. 1999
Campo esiomyces bha ii GMBCC 1120 T PQ763360 PQ842543 –PV388888 PV388894 – Han e al. 2025
Ca. bha ii GMBCC 1125 PQ763361 PQ842544 –PV388889 PV388895 – Han e al. 2025
Campo esiomyces
co eae
GMBCC 1130 T PQ763358 PQ842545 –PV388890 PV388896 – Han e al. 2025
Ca. co eae GMBCC 1131 PQ763359 PQ842546 –PV388891 PV388897 – Han e al. 2025
Campo esiomyces mali KUMCC 19-0216 T MN792813 MN792811 – – MN794018 – Hyde e al. 2020
Campo esiomyces
pa agoniensis
BBB MVB 573 T JN127358 JN127359 – – – – Sánchez e al. 2012
Campo esiomyces
pue ensis
GMBCC 1113 T PQ763356 PQ842541 –PV388886 PV388892 – Han e al. 2025
Ca. pue ensis GMBCC 1114 PQ763357 PQ842542 –PV388887 PV388893 – Han e al. 2025
Campo esiomyces
accinii
CBS 216.90 T AY916486 AY856879 – – – – Tsui e al. 2006
Cancellidium
a ob unneum
MFLUCC 20-0100 T MT422724 MT422740 MT422726 –MT436438 – Hyde e al. 2021
Cancellidium cine eum MFLUCC 18-0424 T MT370353 MT370363 MT370351 MT370486 MT370488 – Hyde e al. 2021
Cancellidium
g iseonig um
MFLUCC 17-2117 T MT370354 MT370364 MT370352 MT370487 – – Hyde e al. 2021
Cap onia camelliae-
yunnanensis
CGMCC 3.19061 T MH807377 MH807378 – – – – Phookamsak e al. 2019
Cap onia kleinmondensis CBS 122671 T MH863226 MH874753 – – – – Vu e al. 2019
Cap onia leucadend i CBS 122672 T MH863227 MH874754 – – – – Vu e al. 2019
Cap onia lijiangensis CGMCC 3.20501 T OK487581 OK487580 – – – – Phukhamsakda e al.
2022
Cap onia pilosella AFTOL-ID 657 DQ826737 DQ823099 – – – – James e al. 2006
Cladophialopho a boppii CBS 126.86 T EU103997 NG_058762 – – – – Gueidan e al. 2008
Cladophialopho a
ca ionii
CBS 114393 KF928452 KF928516 – – – EU137151 de Hoog e al. 2007;
A ili-Angelis e al. 2014
Cl. ca ionii CBS 160.54 T AB109177 LC192080 – – – EU137201 Abliz e al. 2004; de
Hoog e al. 2007; Kiyuna
e al. 2018
Cladophialopho a
lo idana
NRRL 66282 T AB986343 AB986343 – – – – Obase e al. 2016
Cladophialopho a
chae ospi a
CBS 114747 EU035403 KF928514 – – – KF928578 C ous e al. 2007; A ili-
Angelis e al. 2014
Cladophialopho a
ma sushimae
MFC-1P384 T FN549916 FN400758 – – – – Koukol 2010
Cladophialopho a
myce oma is
CBS 122637 T FJ385276 LC192076 – – – – Badali e al. 2008;
Kiyuna e al. 2018
Cladophialopho a yeg esii CBS 114405 T EU137322 KX822323 – – – EU137209 de Hoog e al. 2007;
Vasse e al. 2017
Cyphellopho a aes i a aCBS 228.86 T KC455244 KC455257 – – – KC455227 Réblo á e al. 2013
Cyphellopho a lacinia a CBS 190.61 T EU035416 FJ358239 – – – JQ766329 C ous e al. 2007;
Gueidan e al. 2008;
Feng e al. 2014
9
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Taxon S ain S a us GenBank Accession numbe s Re e ence
ITS LSU SSU pb2 e 1 ub2
Cyphellopho a su onii CBS 449.91 T KC455243 KC455256 – – – KC455226 Réblo á e al. 2013
Dema iohelicomyces
helicospo us
MFLUCC 16-0213 T KX454169 KX454170 –MF535258 KY117035 – Hyde e al. 2016; Lu e
al. 2017b, 2018b
Dema iohelicospo um
gu ula um
MFLUCC 17-2011 T MH558705 MH558833 –MH551021 MH550896 – Lu e al. 2018a
Dema io ubeu ia
chiang aiensis
MFLUCC 10-0115 T JN865200 JN865188 – – KF301551 – Boonmee e al. 2011,
2014
Dic yospo a hailandica MFLUCC 16-0001 T KY873627 KY873622 –MH551023 KY873286 – B ahamanage e al.
2017; Lu e al. 2018a
Exophiala angulospo a CBS 482.92 T JF747046 MH874033 – – – JN112426 de Hoog e al. 2011; Vu
e al. 2019
Exophiala cas ellani CBS 158.58 T KF928458 MH869272 – – – KF928586 A ili-Angelis e al. 2014;
Vu e al. 2019
Exophiala de ma i idis CBS 207.35 T AF050269 KF928508 – – – KF928572 Un e eine and Na eau
1999; A ili-Angelis e
al. 2014
Exophiala equina FMR 18335 ON009852 ON009932 – – – ON491590 To es-Ga cia e al. 2023
Exophiala he e omo pha CBS 232.33 T AY857524 MH866871 – – – – P ena e a-Boldú e al.
2006; Vu e al. 2019
Exophiala jeanselmei CBS 507.90 T MH862234 MH873915 – – – – Vu e al. 2019
Exophiala mesophila CBS 402.95 T JF747111 KX712349 – – – JN112476 de Hoog e al. 2011;
Teixei a e al. 2017
Exophiala nig a CBS 535.94 T MH86248 KX712353 – – – – Teixei a e al. 2017; Vu
e al. 2019
Exophiala nishimu ae CBS 101538 T AY163560 KX712351 – – – JX482552 de Hoog e al. 2003;
Woo e al. 2013; Teixei a
e al. 2017
Exophiala pisciphila CBS 537.73 T NR_121269 MH872483 – – – JN112493 de Hoog e al. 2011;
Schoch e al. 2014; Vu e
al. 2019
Exophiala p o o opha CBS 534.94 OR371992 – – – – – Unpublished
Exophiala adicis FMR 18645 ON009857 ON009937 – – – ON667789 To es-Ga cia e al. 2023
Exophiala salmonis CBS 157.67 T MH858932 MH870616 – – – JN112499 de Hoog e al. 2011; Vu
e al. 2019
Fluminicola aqua ica MFLUCC 15-0962 T MF374357 MF374366 MF374374 –MF370960 – Zhang e al. 2017
Fluminicola sap ophy ica MFLUCC 15-0976 T NR_153493 MF374367 MF374375 MF370954 MF370956 – Zhang e al. 2017
Fluminicola s ia a MFLUCC 18-0990 T MW286496 MW287770 – – – – Dong e al. 2021
Fonsecaea monopho a CBS 289.93 AY366925 – – – – EU938554 de Hoog e al. 2004;
Naja zadeh e al. 2009
Fonsecaea nubica CBS 269.64 T EU938592 – – – – EU938574 Naja zadeh e al. 2009
Fonsecaea ped osoi CBS 271.37 NAB114127 KJ930166 – – – EU938559 de Aze edo e al. 2015;
Naja zadeh e al. 2009
Fonsecaea pugnacius CBS 139214 T KR706553 KR706549 – – – KR706547 de Aze edo e al. 2015
Helicangiospo a lignicola MFLUCC 11-0378 T KF301523 KF301531 – – KF301552 – Boonmee e al. 2014
Helicoa c a us aqua icus MFLUCC 17-1996 T MH558707 MH558835 –MH551024 MH550898 – Lu e al. 2018a
Helicodochium
aqua icum
MFLUCC 17-2016 T MH558709 MH558837 –MH551026 MH550900 – Lu e al. 2018a
Helicohyalinum
aqua icum
MFLUCC 16-0014 MH558711 MH558839 –MH551028 MH550902 – Lu e al. 2018a
Helicohyalinum
in undibulum
MFLUCC 16-1133 T MH558712 MH558840 –MH551029 MH550903 – Lu e al. 2018a
Helicoma b unneispo um MFLUCC 17-1983 T MH558714 MH558842 –MH551031 MH550905 – Lu e al. 2018a
Helicoma longispo um MFLUCC 17-1997 T MH558720 MH558846 –MH551037 MH550911 – Lu e al. 2018a
Helicomyces hyalospo us MFLUCC 17-0051 T MH558731 MH558857 –MH551047 MH550922 – Lu e al. 2018a
Helicomyces o qua us MFLUCC 16-0217 MH558732 MH558858 –MH551048 MH550923 – Lu e al. 2018a
Helicospo ium aqua icum MFLUCC 17-2008 T MH558733 MH558859 –MH551049 MH550924 – Lu e al. 2018a
Helicospo ium
la ispo um
MFLUCC 17-2020 T MH558734 MH558860 –MH551050 MH550925 – Lu e al. 2018a
Helicospo ium
lu eospo um
MFLUCC 16-0226 T KY321324 KY321327 –MH551056 KY792601 – Lu e al. 2017a; 2018a
Helicospo ium se i e um MFLUCC 17-1994 T MH558735 MH558861 –MH551051 MH550926 – Lu e al. 2018a
Helicospo ium ege um CBS 254.75 – DQ470982 –DQ470934 DQ471105 – Spa a o a e al. 2006
Helicospo ium esica ium MFLUCC 17-1795 T MH558739 MH558864 –MH551055 MH550930 – Lu e al. 2018a
Helico ubeu ia
guangxiensis
MFLUCC 17-0040 T MH290018 MH290023 –MH290033 MH290028 – Liu e al. 2018b
Helico ubeu ia hydei MFLUCC 17-1980 T MH290021 MH290026 –MH290036 MH290031 – Liu e al. 2018b
Chlamydo ubeu ia
cylind ica
MFLUCC 16-1130 T MH558702 MH558830 –MH551018 MH550893 – Lu e al. 2018a
16
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
CBS 449.91 T Cyphellopho a su onii
CBS 190.61 T Cyphellopho a lacinia a
CBS 228.86 T aCyphellopho a aes i a
99/1 BSS 376
T171 Tm12
CBS 269.64 T Fonsecaeanubica
CBS 139214 T Fonsecaeapugnacius
CBS 289.93 Fonsecaea monopho a
CBS 271.37 T Fonsecaeaped osoi
81/–
MFC 1P384 T Cladophialopho a ma sushimae
CBS 122637 T Cladophialopho a myce oma is
CBS 126.86 T Cladophialopho a boppii
NRRL 66282 T Cladophialopho a lo idana
CBS 114747 Cladophialopho a chae ospi a
CBS 114405 T Cladophialopho a yeg esii
CBS 160.54 T
CBS 862.96
UAMH 10875 T Phialopho a ame icana
CBS 140325 E Phialopho a e ucosa
BMU 01890 T Phialopho a chinensis
91/0.99
87/1
92/1
JAUCC M0840-1 T Neo e onaea sinensis
FMR 18994 T Aciculomyces es ic us
HMAS 245592 T A okylind iopsis se ulosa
90/1 CBS 482.92 T Exophiala angulospo a
MFLUCC 120389 T Melanoc ona ec onae
MUCL 39954 Be lesiella nige ima
CGMCC 3.19061 T Cap onia camelliae yunnanensis
CGMCC 3.20501 T Cap onia lijiangensis
AFTOL ID-657 Cap onia pilosella
MFLUCC 16-1449 T Ma inophialopho a ga e hjonesii
CBS 534.94 Exophiala p o o opha
CBS 232.33 T Exophiala he e omo pha
CBS 207.35 T Exophiala de ma i idis
87/–
99/1
CBS 181.65 IN Rhinocladiellaanceps
CBS 496.78 T Rhinocladiellaphaeopho a
CBS 313.73 T Rhinocladiellaaquaspe sa
MFLUCC 11-0529 T Aculea a aqua ica
CBS 101538 T Exophiala nishimu ae
CBS 507.90 T Exophiala jeanselmei
CBS 535.94 T Exophiala nig a
CBS 101597 Phaeoannellomyces elegans
CBS 264.49 T Melanchlenus eume abolus
CBS 317.33 AS Rhinocladiella a o i ens
CBS 122672 Cap onia leucadend i
CBS 122671 Cap onia kleinmondensis
93/1
CBS 147585 T Neohe po ichiella juglandicola
CBS 537.73 T Exophiala pisciphila
CBS 157.67 T Exophiala salmonis
FMR 18335 Exophiala equina
FMR 18645 Exophiala adicis
93/1CBS 572.90 T e
CBS 254.57 T
CBS 268.75 T Ve onaea compac a
CBS 776.83 T Ve onaea japonica
88/1
99/1
76/0.95
CBS 158.58 Exophiala cas ellanii
CBS 402.95 T Exophiala mesophila
CBS 145909 T Thysano ea can elliae
CBS 145910 T Thysano ea sei e ii
CBS 215.96 T Thysano ea ac opleu ogena sp. no .
YMF 1.03683 T Thysano ea sinensis comb. no .
YMF 1.04038 T Thysano ea hainanensis comb. no .
YMF 1.04080 T Thysano ea wuzhishanensis comb. no .
99/1
MFLUCC 15-0416 Thysano ea obscu a
MFLUCC 17-2378 T Thysano ea non amosa
CBS 212.96 T
MFLUCC 15-0966 T d
MFLUCC 15-0971 T Thysano ea hailandensis
CBS 862.68
MFLUCC 15-0415 T
MFLUCC 15-0237 T Thysano ea asia ica
MFLUCC 15-0259 T Thysano ea cu a a
CBS 126086 Thysano ea ousseliana
KUMCC 15-0206 T Thysano ea subme sa
MFLUCC 15-0414 T Thysano ea yunnanensis
86/1
82/–
86/0.99
77/0.99
91/1
88/1
–/0.99
–/0.98
–/0.98
–/0.97
–/0.95
–/1
–/0.97
100/–
–/1
–/0.95
0.1
He po ichiellaceae (Chae o hy iales)
Thysano ea melanica
Thysano ea papuana
ou g oup
Cyphellopho aceae
Thysano ea
Ve onaeabo yosa
Cladophialopho a ca ionii
Valen iella maceioensis
Figu e 2. Maximum likelihood phylogene ic ee o he He po ichiellaceae based on ITS–LSU– ub2 DNA sequences.
Names in bold indica e axonomic no el ies; s ains wi h hei accession numbe s in bold and highligh ed in iole colou
we e sequenced in his s udy; a ex- ype o Cyphellopho a e mispo a, d ex- ype o Thysano ea aqua ica, e ex- ype o Ve o-
naea cons ic a. T, E, IN and AS deno e ex- ype, ex-epi ype, ex-isoneo ype, and au hen ic s ains, espec i ely. Thickened
b anches indica e suppo ML BS = 100% and PP alues = 1.0. B anch suppo o nodes ≥ 70% ML and ≥ 0.95 PP is indi-
ca ed abo e o below b anches. A hyphen (–) indica es alues lowe han 75% ML BS o 0.95 PP.
17
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
(YMF1.03683), and U. wuzhishanensis (YMF1.04080), we e nes ed wi hin he
Thysano ea clade, co obo a ing hei ans e o Thysano ea. These species
oge he o med a sis e lineage (71/95) o T. ac opleu ogena. The sys ema ic
posi ion o U. ha oldiae, he ype species o Uncispo a, emains unknown.
The hi d phylogene ic analysis (Fig. 3), based on he ITS–LSU–SSU– pb2–
e 1 da ase , included 27 ing oup s ains ep esen ing 21 species in ou gene a
o he Papulosaceae and ela ed gene a (So da iomyce es ince ae sedis). A o al
o 258 nucleo ides we e excluded om he 5′ and/o 3′ ends o LSU, SSU, pb2,
and e 1 da ase s due o incomple e sequences in mos s ains. O he 4 472 cha -
ac e s (including gaps), 1 335 we e unique si es iden i ied by RAxML: 328 in ITS,
190 in LSU, 87 in SSU, 481 in pb2, and 249 in e 1. Th ee species o Cancellidium
(Cancellidiales), i.e. Cn. a ob unneum MFLUCC 20-0100, Cn. cine eum MFLUCC
18-0424, and Cn. g iseo-nig um MFLUCC 17-2117, we e used as ou g oup axa.
The bes - i models o nucleo ide subs i u ion selec ed we e: SYM+I+G (ITS),
GTR+I+G (LSU, pb2, e 1), and GTR+I (SSU). Papulosaceae we e eco e ed as
a s ongly suppo ed lineage (100/1.0) comp i sing B unneospo ella aqua ica,
Fluminicola (96/90), Papulosa ame ospo a and Wongia (100/1.0), oge he wi h
he closely ela ed gene a Pseudos anjehughesia (100/1.0) and Pla y achelon.
Bo h s ains, CBS 440.70 and CBS 531.73, clus e ed wi hin he well-suppo ed
Wongia subclade. The s ain CBS 440.70 g ouped wi h W. aqua ica in a basal
posi ion and is he e desc ibed as a new species, W. pallidopola is. S ain CBS
531.73 was esol ed as a new lineage, W. hachidopho a, which was nes ed
wi hin a s ongly suppo ed subclade (100/1.0). I includes mo phological-
ly simila species such as W. bambusae, W. bandungensis, W. usi o mis, and
W. suae ha likely ep esen a species complex.
To assess he ela ionships o W. hachidopho a wi h hese species, h ee
loci commonly employed o species-le el delimi a ion, ITS, e 1, and pb2,
we e compa ed. Pai wise sequence iden i ies indica ed ha W. hachidopho-
a is mos closely ela ed o W. bandungensis (ITS: 99.4%, e 1: 99.0%, pb2:
99.4%) and W. suae (ITS: 98.4%, e 1: 98.7%, pb2: 99.6%), while showing lowe
simila i y o W. bambusae (ITS: 97.1–97.8%, e 1: 98.5%, pb2: 98.7%), and W.
usi o mis (ITS: 95.3%, e 1: 98.5%, pb2: 96.9%).
The ITS egion shows mode a e di e gence (0.6–1.6%). The ITS sequences
show ha W. hachidopho a is closes o W. bandungensis (99.4%). Howe e ,
i s ill shows small di e ences ha may o may no exceed in aspeci ic a i-
a ion h esholds. The e is also high simila i y among W. hachidopho a and
W. suae/W. bambusae (97–98%), bu below ypical species h eshold (≥98.5–
99%) (Vu e al. 2019; Lücking e al. 2020). In he e 1 da ase , all alues ho e
a ound 98.5–99%, showing s ong ela edness ac oss species; W. hachidopho a
is again mos simila o W. bandungensis. The pb2 sequence o W. hachidopho-
a is sho e 755 bp s 900–1000 bp in o he s, which can in la e iden i y alues
sligh ly because less a iable egions a e o en e ained. Apa om W. usi o m-
is, all pb2 iden i ies a e e y high (≥98.7%). The highes sequence iden i y is o
W. suae and o W. bandungensis. Toge he wi h mo phological dis inc i eness
(see no es o W. hachidopho a), he molecula da a suppo W. hachidopho-
a as a dis inc species wi hin he species complex. In addi ion, he sequence
compa ison o W. bandungensis and W. suae indica e ha ITS iden i y (98.6%) is
igh a he species bounda y in ungi. Thei e 1 and pb2, howe e , show nea ly
comple e iden i y (≥99.9%), s ongly indica ing hey a e he same species.
18
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
MFLUCC 18-0424 T Cancellidium cine eum
MFLUCC 20-0100 T Cancellidium a ob unneum
MFLUCC 17-2117 T Cancellidium g iseonig um
CBS 125235 Pla y achelonabie is
MFLUCC 16-0569 T Pseudos anjehughesia aqui opica
MFLUCC 15-0352 T Pseudos anjehughesia lignicola
AFTOL ID-748 Papulosaame ospo a
HKUCC 3708 B unneospo ellaaqua ica
MFLUCC 18-0990 T Fluminicola s ia a
MFLUCC 15-0962 Fluminicola aqua ica
MFLUCC 15-0976 T Fluminicola sap ophy ica
99/1
96/–
MFLUCC 18-1607 T Wongia aqua ica
CBS 440.70 T Wongia pallidopola issp. no .
KUNCC 24-17692 T Wongia gu ula a
KUNCC 23-13715 T Wongia la a
BCRC FU32062 T Wongia miscan hi
BRIP 69019 T Wongia iche ai
DAR 79637 T Wongia ga e ii
DAR 80512 T
BRIP 60377
DLUCC 1767
KUNCC 23-16632
MFLUCC 21-0032 T
MFLUCC 21-0028
82/–KUNCC 24-17699
CGMCC 3.24360 T
CBS 531.73 T Wongia hachidopho a sp. no .
CGMCC 3.24295 T Wongia suae
TBRC-BCC 95171 T
TBRC-BCC 95343 I
91/1
98/0.98
99/1
98/1
94/1
81/0.97
96/1
97/1
76/–
0.05
100/1
ou g oup
Cancellidiales
Papulosaceae
Wongia
So da iomyce idaeince aesedis
Wongia bandungensis
Wongia bambusae
Wongia usi o mis
Wongia g i inii
Figu e 3. Maximum likelihood phylogene ic ee o he Papulosaceae based on ITS–LSU–SSU– pb2– e 1 DNA sequenc-
es. Names in bold indica e axonomic no el ies; s ains wi h hei accession numbe s in bold and highligh ed in iole
colou we e sequenced in his s udy. T and I deno e ex- ype and ex-iso ype s ains, espec i ely. Thickened b anches
indica e suppo ML BS = 100% and PP alues = 1.0. B anch suppo o nodes ≥ 70% ML and ≥ 0.95 PP is indica ed abo e
o below b anches. A hyphen (–) indica es alues lowe han 75% ML BS o 0.95 PP.
The inal phylogene ic analysis (Fig. 4), conduc ed on he ITS–LSU– pb2–
e 1 da ase , included 71 ing oup s ains ep esen ing 65 species o he
Tubeu iaceae (Tubeu iales, Do hideomyce es). Nine y- i e nucleo ides we e
excluded om he 5′ end o he LSU da ase due o incomple e sequences
in mos s ains. O he o al o 3 512 cha ac e s (including gaps), 1 631 we e
unique si es iden i ied by RAxML: 442 in ITS, 271 in LSU, 538 in pb2, and
380 in e 1. Th ee membe s o Bo yosphae ia (Bo yosphae iales), i.e. B.
aga es MFLUCC 10-0051, B. wangensis HGUP190007, and B. do hidea CBS
115476, we e selec ed as ou g oup axa. The bes - i models o nucleo ide
19
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
CBS 115476 E Bo yosphae iado hidea
HGUP 190007 Bo yosphae ia wangensis
MFLUCC 10-0051 Bo yosphae iaaga es
MFLUCC 10-0124 E
MFLUCC 16-0024
MFLUCC 10-0125 T Acan hos igma chiangmaiense
UAMH 7237 Acan hos igma pe pusillum
CBS 122759 T Skoliomycella la agen. e sp.no .
CBS 148315 T Zaanenomyces mode a icis-academiae
CBS 149453
CBS 148312 T
CBS 113561 T Zaanenomyces hili e sp. no .
CBS 148310 T
CBS 148272
GZAAS 19-1751 T Neodic yospo a ka s i
MFLUCC 16-0226 T Helicospo ium lu eospo um
MFLUCC 17-1994 T Helicospo ium se i e um
MFLUCC 17-2008 T Helicospo iumaqua icum
CBS 254.75 Helicospo ium ege um
MFLUCC 17-2020 T Helicospo ium la ispo um
MFLUCC 17-1795 T Helicospo ium esica ium
88/1
KUMCC 19-0216 T Campo esiomyces mali
BBB MVB 573 Campo esiomyces pa agoniensis
CBS 216.90 T Campo esiomyces accinii
GMBCC 1125
GMBCC 1120 T
90/1GMBCC 1114
GMBCC 1113 T
GMBCC 1130 T
GMBCC 1131
92/1
85/0.99
99/1
83/1
MFLUCC 16-1125 T Neo ubeu ia k abiensis
MFLUCC 11-0378 T Helicangiospo a lignicola
MFLUCC 16-0001 T Dic yospo a hailandica
MFLUCC 17-2011 T Dema iohelicospo umgu ula um
MFLUCC 10-0116 T Acan hohelicospo a pinicola
MFLUCC 11-0510 T Neoacan hos igma usi o me
MFLUCC 17-0049 T Be kleasmium aqua icum
MFLUCC 17-1978 T Be kleasmium usi o me
MFLUCC 10-0115 T Dema io ubeu ia chiang aiensis
MFLUCC 11-0379 T Neohelicoma agacea um
MFLUCC 17-1997 T Helicoma longispo um
MFLUCC 17-1983 T Helicoma b unneispo um
CGMCC 3.23539 T Neomanoha acha iellaaqua ica
MFLUCC 17-1996 T Helicoa c a us aqua icus
MFLU 11-0228 T Kamalomyces bambusicola
MFLUCC 13-0233 T Kamalomyces hailandicus
86/1
93/1
77/1
MFLUCC 17-1982 T Pleu ohelicospo ium pa ispo um
GZCC 20-0489 T Pleu ohelicospo ium hyalinum
MFLUCC 16-1134 T Chlamydo ubeu ia k abiensis
MFLUCC 16-1130 T Chlamydo ubeu ia cylind ica
MFLUCC 16-0010 T Aquaphila albicans
MFLUCC 17-1980 T Helico ubeu ia hydei
MFLUCC 17-0040 T Helico ubeu ia guangxiensis
MFLUCC 16-0016 T Neochlamydo ubeu ia usi o mis
MFLUCC 10-0118 T Neochlamydo ubeu ia khunko nensis
MFLUCC 16-0213 T Dema iohelicomyces helicospo us
MFLUCC 17-2016 T Helicodochiumaqua icum
MFLUCC 16-1133 T Helicohyalinumin undibulum
MFLUCC 16-0014 Helicohyalinumaqua icum
MFLUCC 17-1523 T Neohelicospo iumpa ispo um
MFLUCC 17-1522 T Neohelicospo iumguangxiense
MFLUCC 12-0170 T Manoha acha iella ec onae
MFLUCC 12-0388 Boe lagiomyces mac ospo us
KUMCC 15-0276 Mu ipulch aaqua ica
MFLUCC 16-0993 T Neohelicomyces aqua icus
KUMCC 15-0470 T Neohelicomyces g andispo us
MFLUCC 17-0051 T Helicomyces hyalospo us
MFLUCC 16-0217 Helicomyces o qua us
MFLUCC 17-2021 Pa ahelicomyces albo ii
CGMCC 32-3535 Pa ahelicomyces hyalospo us
MFLUCC 17-0053 T Tubeu ia inaequalis
MFLUCC 17-1524 T Tubeu ia eccen ica
GZCC 22-2010 T Pseudo ubeu ia hyalospo a
GZCC 22-2011 T Pseudo ubeu ia laxispo a
94/1
86/1
83/1
99/1
81/–
92/1
0.1
–/0.98
–/0.98
–/0.99–/1
–/0.98
1x
ou g oup
Bo yosphae iales
Tubeu iaceae (Tubeu iales)
Campo esiomyces co eae
Campo esiomyces pue ensis
Campo esiomyces bha ii
Zaanenomyces quad ipa is
Zaanenomyces e sa ilis
Thax e iellopsis lignicola
Figu e 4. Maximum likelihood phylogene ic ee o he Tubeu iaceae based on ITS–LSU– pb2– e 1 DNA sequences.
Names in bold indica e axonomic no el ies; s ains wi h hei accession numbe s in bold and highligh ed in iole colou
we e sequenced in his s udy. T and E deno e ex- ype and ex-epi ype s ains, espec i ely. Thickened b anches indica e
suppo ML BS = 100% and PP alues = 1.0. B anch suppo o nodes ≥ 70% ML and ≥ 0.95 PP is indica ed abo e o below
b anches. A hyphen (–) indica es alues lowe han 75% ML BS o 0.95 PP.
20
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
subs i u ion selec ed we e: GTR+I+G (ITS, LSU, and e 1) and SYM+I+G
( pb2). S ains CBS 113561 and CBS 122759 we e esol ed as membe s
o he Tubeu iaceae. The esul ing opology eco e ed 37 well-suppo ed
gene ic lineages wi hin he amily. S ain CBS 113561 clus e ed wi hin
Zaanenomyces clade (83/1.0) closely ela ed o Z. quad ipa is, suppo ing
i s ecogni ion as a new species, Z. hili e . They o med a s ongly suppo ed
sis e lineage (100/1.0) o a subclade (100/1.0) comp ising Z. mode a icis-
academiae and Z. e sa ilis. In con as , s ain CBS 122759 o med a
dis inc lineage, jus i ying he es ablishmen o a new mono ypic genus,
Skoliomycella, wi h S. la a as i s ype species. Bo h Skoliomycella and
Zaanenomyces we e placed wi hin a obus subclade (92/1.0) oge he
wi h Acan hos igma (100/1.0), Campo esiomyces (99/1.0), Helicospo ium
(88/1.0), and Neodic yospo a ka s ii.
Mo phological s udies
Excep o CBS 113561 (see below) and CBS 862.68 (which emained s e ile),
all o he s ains could be eadily dis inguished om P. pa ispo um unde cul-
u e condi ions. Two s ains ep esen wo dis inc species o Thysano ea (A -
zanlou e al. 2007) in he He po ichiellaceae. Thysano ea ac opleu ogena (CBS
215.96) spo ula ed abundan ly, p oducing simple o apically loosely b anched
conidiopho es wi h e minal and in e cala y conidiogenous cells o ming an
elonga ed den icula e achis, and pale b own o pale oli aceous b own, p e-
dominan ly 3-sep a e conidia. In con as , T. melanica (CBS 862.68) (Liu e al.
2015) emained s e ile on all es ed media, and i s iden i ica ion elied exclu-
si ely on molecula da a.
Two o he s ains belong o he genus Wongia in he Papulosaceae, whe e
hey ep esen dis inc species lineages, he e desc ibed as W. pallidopola -
is (CBS 440.70) and W. hachidopho a (CBS 531.73). Bo h species p oduced
b own o da k b own conidia, bo ne on e minal conidiogenous cells wi h one
o se e al den icles on ela i ely sho , obus conidiopho es.
The wo las s ains, CBS 113561 and CBS 122759, a e membe s o
he Tubeu iaceae. S ain CBS 113561 p oduced hyaline, mos ly 3-sep a e
conidia, and simple conidiopho es wi h a sho den icula e achis and
some imes bea ing nodulose swellings. In cul u e, i s mo phology closely
esembled ha o P. pa ispo um, making i s dis inc ion challenging.
This s ain is desc ibed he e as he new species Zaanenomyces hili e . In
con as , s ain CBS 122759 is eadily dis inguished om P. pa ispo um
by lexuous o sinuous conidiopho es wi h a p onounced zig-zag pa e n,
bea ing soli a y den icles sca e ed along he en i e conidiopho e axis,
and by i s 3–7-sep a e, subhyaline o pale oli aceous-b own conidia. I
ep esen s a no el lineage in he Tubeu iaceae and is in oduced he e as he
new genus and species Skoliomycella la a.
Biogeog aphy assessmen
The biogeog aphic assessmen e ealed s iking con as s in dis ibu ion
pa e ns among he s udied axa (Fig. 5). Some species, such as T. melani-
ca, W. pallidopola is, and Z. hili e , a e clea ly cosmopoli an and ecologically
21
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
e sa ile, occu ing ac oss mul iple con inen s, biomes, and clima e zones.
Thei s ong ep esen a ion in soils, pa icula ly c oplands and g asslands,
poin s o an ecological p e e ence o e es ial subs a es and sugges s ha
human ac i i ies, especially ag icul u e, may ha e played a ole in hei global
dissemina ion. The high abundance o W. pallidopola is and T. melanica in
Eu opean c oplands is pa icula ly no able and suppo s he idea ha hese
axa h i e in dis u bed and an h opogenic habi a s.
By con as , o he species appea much mo e es ic ed. Pleu oph agmi-
um pa ispo um shows a concen a ion in Eas Asia wi h sca e ed Eu ope-
an eco ds, mainly om aqua ic habi a s, e lec ing a na owe ecological
niche. These indings p o ide a b oade pe spec i e and con as wi h p e-
iously published s udies, which ha e so a been based la gely on Eu ope-
an collec ions o his species (Cos an in 1888; Ellis 1968; Holubo á-Jecho á
1972) and a ew eco ds om Japan (Ma sushima 1975) and No h Ame ica
Figu e 5. Geog aphical dis ibu ion o he s udied species based on en i onmen al DNA eco ds om he GlobalFungi
da abase. Each do indica es a geo e e enced sample in which he species was de ec ed, wi h he numbe o samples
(n) gi en in pa en heses.
22
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
(Wang 2010). Skoliomycella la a is ep esen ed by a single en i onmen al
eco d in u ban ai om he Ibe ian Peninsula, while W. hachidopho a shows
a disjunc dis ibu ion be ween India and sub opical No h Ame ica, hin ing
a ei he ue a i y o unde -sampling.
Finally, T. ac opleu ogena emains known only om i s ype locali y, wi h no
ma ches in en i onmen al sequencing da ase s. These pa e ns collec i e-
ly illus a e how some lineages a e widesp ead ye c yp ic, while o he s a e
genuinely a e o o e looked, unde sco ing he impo ance o in eg a ing eDNA
su eys wi h adi ional axonomy o esol e hei ecology and biogeog aphy.
Taxonomy
Pleu oph agmium Cos an in, Les mucédinées simples: 100. 1888.
Synonyms. Dac yla ia sec . Pleu oph agmium (Cos an in) de Hoog, S ud.
Mycol. 26: 36. 1985.
Neomy mec idium C ous, Pe soonia 41: 287. 2018.
Type species. Pleu oph agmium pa ispo um (P euss) Hol.-Jech.
Desc ip ion. Sexual mo ph. No obse ed. Asexual mo ph. Colonies e use,
hai y, b own o black, subhyaline, beige o pale b own when spo ula ing; ege a i e
hyphae imme sed o semi-imme sed. Conidiopho es mac onema ous, mononem-
a ous, soli a y o in small g oups, e ec , s aigh o sligh ly lexuous, unb anched
o a ely b anched, some imes elonga ing pe cu en ly, b own, sep a e. Conidio-
genous cells in eg a ed, e minal and/o in e cala y esul ing om he o ma ion
o a sep um wi hin he o iginal cell du ing sympodial elonga ion, polyblas ic, oc-
casionally monoblas ic, sympodially p oli e a ing, bea ing one o se e al den icles
o o ming a achis wi h minu e, pimple-like den icles o p o usions sca e ed
o e he su ace; conidiogenesis holoblas ic-den icula e. Conidia soli a y, d y,
ac opleu ogenous, obo oid o subcla a e, usi o m, ellipsoidal, usually ape ing
owa ds he base, wi h a dis inc hilum, hyaline, subhyaline o pale b own, some-
imes wi h pale end cells, asep a e o ans e sely sep a e, wall smoo h o inely
o namen ed, wi h an epheme al mucoid shea h; conidial secession schizoly ic.
No es. Mo phological compa isons oge he wi h phylogene ic analyses o
P. pa ispo um CBS 770.83 suppo he iew ha gene a Neomy mec idium and
Pleu oph agmium a e congene ic, leading o he p oposal o 11 new combina-
ions and he in oduc ion o wo new names.
Gi en he subs an ial mo phological a iabili y among species cu en ly
placed in Pleu oph agmium, he genus is unlikely o ep esen a single e olu-
iona y lineage, ende ing he adi ional gene ic concep un enable. Acco ding-
ly, he p esen gene ic desc ip ion is based on he mo phological a iabili y o
species whose placemen is suppo ed by molecula da a and e e s o Pleu o-
ph agmium s. s . The ollowing lis o species is o ganised in o h ee ca ego-
ies: (i) species accep ed in Pleu oph agmium s. s .; (ii) species o unce ain
s a us ha emain in Pleu oph agmium s. la .; and (iii) species excluded om
Pleu oph agmium and ans e ed o o he gene a. Names in bold indica e he
cu en ly accep ed classi ica ion, accompanied by ull synonymy and, whe e
app op ia e, b ie explana o y no es. The in e speci ic a iabili y o species lis -
ed in he i s wo ca ego ies is summa ised in he synopsis able (Table 3). Key
o species o Pleu oph agmium was p o ided by D’Souza and Bha (2012).
23
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Table 3. Synopsis able o species o Pleu oph agmium.
Species accep ed in Pleu oph agmium s. s .
Taxon Subs a e* Size (µm) Conidia End cells O namen a ion Conidiogenous cellsConidiopho es Re e ence
Sep a Shape Shea h Colou Size (µm) Size (µm)
Pleu oph agmium
asia icum
C(13–)15–16(–17) ×
(3.5–)4.5(–5)
(2–)3 ellipsoid o obo oid, wi h unca e hilum
a he base
p esen pale b own smoo h-walled 5–35 × 4–7 50–100 × 3–5 C ous e al.
2018a
Pleu oph agmium
asymme icum
NS 12–15 × 2–3 1na owly cla a e o subcla a e absen yellowish o subhyaline smoo h-walled 8–35 × 3.5–5 40–210 × 3.5–9 Se ano e al.
2020
Pleu oph agmium
lu iale
NS 14–16 × 4–6 3obo oid, ounded a he apex, poin ed
a he base
absen subhyaline smoo h-walled n/a 211–308 × 5–7 Luo e al. 2019
Pleu oph agmium
usi o me
NS 29–34 × 4–6 (0–)1–3 na icula o ape ing, poin ed a bo h
ends
absen pale b own pale smoo h-walled n/a 214–285 × 4–5 Zhang e al.
2024
Pleu oph agmium
gaoligongense
NS 16–24 × 5–7 0–3 cla a e-cymbi o m, unca e a he base absen subhyaline o pale
b own
smoo h-walled 34–68 × 3–6 138–226 × 4–7 Xu e al. 2023
Pleu oph agmium
guizhouense
NS 8.9–12.7 × 2.8–4.8 (2–)3 usoid-ellipsoid, apex ob use o ape ing,
wi h a sub unca e hilum a base
absen subhyaline o pale
b own
smoo h-walled 2.2–4.3 (wid h) 75–140 × 2–4.5 Hyde e al. 2020
Pleu oph agmium
jiulongheense
NS 16–23 × 3.7–5.6 3cla a e o usi o m, unca e a he base p esen subhyaline o pale
b own
smoo h-walled n/a (144–)204–332 ×
3.4–4.8
Cao e al. 2025
Pleu oph agmium
luguense
NS 12–15 × 4–7 (0–)2–3 obo oid, ape ing a he base absen subhyaline o pale
b own
smoo h-walled 39–68 × 3–5 152–298 × 4–6 Xu e al. 2023
Pleu oph agmium
na icula e
NS 16–24 × 5.5–7.5 (1–) 3 na icula o usi o m, ape ing o a hilum
owa ds he base, ob use a he apex
p esen hyaline, becoming pale
b own
pale smoo h-walled n/a 100–200 × 4–5.6 Yang e al. 2023
Pleu oph agmium
pa ispo um
NS 10–18 × 3.5–6 (0–)3(–4) ellipsoid o subcla a e, ounded a he
apex, poin ed a he base
absen hyaline o e y pale
b own
c umpled 26–47(–51) ×
4.5–5.5
(80–)118–250(–
270) × (3.5–)4–5
This s udy
Pleu oph agmium
p e idophy ophilum
NS 8.5–11 × 3–4 0–1 obo oid absen hyaline o pale b own ough-walled 20–55.5 × 2.5–4(–
4.5)
150–224 ×
2.7–4.9
Zhang e al.
2025
Pleu oph agmium
sep a um
C(12–)14–16(–20) ×
(3.5–)4(–5)
(1–) 3 usoid-ellipsoid, apex ob use, ape ing in
lowe hi d o a unca e hilum
p esen hyaline, becoming pale
b own
smoo h-walled 30–40 × 4–5 40–70 × 4–5 C ous e al.
2018a
Pleu oph agmium
sichuanense
NS 9.5–14.5 × 4–4.5 0–1 usi o m o na owly obo oid absen subhyaline o pale
b own
inely e ucose n/a 93–130 × 4–5 Chen e al. 2025
Pleu oph agmium
so bicola
C(7–)8–10(–15) ×
4(–5)
(0–)1(–3) obo oid, ob use a he apex p esen hyaline, pale b own
wi h age
smoo h-walled 20–65 × 3–4 50–200 × 4–7 C ous e al.
2018b
Pleu oph agmium s. la .: species o unce ain s a us
Pleu oph agmium
angamosense
C24.5–40 × 5–8 3–7 mos ly 5 usi o m, cu ed o s aigh absen pale b own pale smoo h-walled 3–3.5 (wid h) 200 × 3.5–4.5 Ma sushima
1995
Pleu oph agmium
aqua icum
NS 25–30 × 6–6.5 3 usi o m o cla a e, some imes
na icula , unca ed a he base
absen b own, pale a ound
sep a
pale smoo h-walled n/a 200–390 × 5–11 He edia e al.
2007
Pleu oph agmium
bi unica um
C20–35 × 4.5–7 3(–5) usi o m, na owly unca ed a he base absen b own smoo h-walled n/a (50–) 100–220
× 3–5
Ma sushima
1975
Pleu oph agmium
cla a um
NS 9.5–16.5 × 3.5–4.5 0cla a e o usi o m, na owe a he
ends, ape ed o a poin a he base
absen yellowish b own e ucose n/a 75–160 × 3.3–4 Ma e al. 2014
Pleu oph agmium
ellipsoideum
NS 10–17 × 6–8.5 3ellipsoid o obo oid, ounded a he
apex, poin ed a he base
absen pale b own smoo h-walled n/a 150–260 × 3.5–6 Ma e al. 2014
24
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Species accep ed in Pleu oph agmium s. s .
Taxon Subs a e* Size (µm) Conidia End cells O namen a ion Conidiogenous cellsConidiopho es Re e ence
Sep a Shape Shea h Colou Size (µm) Size (µm)
Pleu oph agmium
ha unganae
NS 12–18 × 5–7 0 usoid, cu ed absen subhyaline o pale
oli aceous
smoo h-walled n/a 150 × 4–5 Hans o d 1946
Pleu oph agmium
indicum
NS 20–30 × 4.5–11 3ellipsoidal o obo oid, ounded a he
apex, poin ed a he base, s aigh o
sligh ly cu ed
absen da k b own pale smoo h-walled 15–23 × 4.5–9.5 100–160 × 7–22 D‘Souza and
Bha 2012
Pleu oph agmium
malaysianum
C(20–) 40–75 × 4–5 3–10
pseudosep a e
cylind ical o cla a e, ounded a he
apex, p o uding a he base
absen hyaline smoo h-walled n/a 25–50 (–100) ×
3.5–5
Ma sushima
1996
Pleu oph agmium
miniumbona um
NS 16–19 × 6–7 (2–) 3 obo oid, py i o m o b oadly cla a e,
some imes sligh ly cla a e, umbona e a
he apex, unca e a he base
absen da k b own o b own subhyaline
basal cell
smoo h-walled 55–95 × 3–3.5 90–150 × 5.5–7 Cas añeda 1999
Pleu oph agmium
na iculi o me
C18–32 × 6.5–10 1na icula , poin y a he apex, poin ed a
he base
absen hyaline smoo h-walled n/a 70–140 × 6–8 Ma sushima
1975
Pleu oph agmium
obcampanuloides
C(10–)11.5–15.5(–
18) × 5–6.5(–7)
(1–)2 obpy i o m o u bina e absen pale oli aceous,
oli aceous in mass
smoo h-walled n/a 100–150 × 3.8–5 Ma sushima
1995
Pleu oph agmium
pe uamazonicum a .
pe uamazonicum
C10–30 × 4–6 2cylind ical, ounded a he apex, poin ed
a he base
absen pale b own, uscous
in mass
subhyaline smoo h-walled n/a 100–300 × 3–4 Ma sushima
1993
Pleu oph agmium
pe uamazonicum a .
in la um
C13–20 × (5–)6–
9(–10)
(1–)2 o oid, ounded a he apex, na ow a
he base
absen pale soo y subhyaline
apical cell
smoo h-walled n/a (50–) 200–600 ×
4–5.5
Ma sushima
1993
Pleu oph agmium
sub usi o me
C21–38 × 5.8–9.2 3–7 usi o m, s aigh o some imes cu ed absen subhyaline, pale
uscous in mass
smoo h-walled n/a 200–500 × 4–5 Ma sushima
1975
Pleu oph agmium
aiwanense
C12–20 × 3–4.5 3cylind o-cla a e, wi h a sligh ly in la ed
apical cell
absen pale b own hyaline smoo h-walled n/a 70–200 (–250) ×
3.5–4.5
Ma sushima
1987
Pleu oph agmium
icolo
NS 17–18 × 4–5 2ellipsoidal wi h, ounded a he apex,
poin ed a he base
absen b own, ligh b own
basal cell
hyaline
apical cell
smoo h-walled n/a 196–200 × 5–7 Rambelli 2009
Pleu oph agmium
a iesep a um
NS 19–22 × 4–5 1–4 cylind ical, ounded a he apex, poin ed
a he base
absen pale oli aceous,
oli aceous-g ey in mass
smoo h n/a 70–125 × 3.5–5 Ma sushima
1975
Pleu oph agmium
e uculosum
C10–16.5 × 3.3–5 1–3(–3) subcla a e, cla a e, poin ed a he base absen subhyaline o pale
b own
e ucose n/a 8.2–26.4 (–15) ×
1.6–4.5 (–3)
Tiwa i e al. 1969
Pleu oph agmium
yunnanense
NS 10–15 × 6–7 (2–)3 b oadly usi o m, na owed a he ends,
ape ed o a poin a he base
absen pale b own smoo h n/a 185–270 ×
3.5–4.5
Ma e al. 2014
* na u al subs a e (NS); cul u e (C).
25
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Aquap e idospo a (Dis osep ispo ales, So da iomyce es) (Yang e al. 2015),
is a dema iaceous hyphomyce e genus ha wa an s compa ison wi h Pleu-
oph agmium. Bo h gene a sha e igid, e ec , da kly pigmen ed, simple conid-
iopho es on he na u al subs a e ha bea e minal and equen ly in e cala y
holoblas ic-den icula e conidiogenous cells ha p oli e a e sympodially, as
well as sep a e conidia ha may be su ounded by a mucoid shea h. Howe e ,
hey di e p ima ily in conidial pigmen a ion. In Pleu oph agmium s. s ., co-
nidia a e ypically hyaline o subhyaline, becoming pale b own a ma u i y o
when in mass, occasionally showing pale end cells, seldom wi h a mucoid
shea h, and ha e hin-walled sep a. Pleu oph agmium usi o me is an excep ion
in ha ing pale b own conidia wi h pale ends. Howe e , in Aquap e idospo a
conidia a e p ima ily pigmen ed, con inuously b own o da k b own, o en wi h
pale end cells, wi h hick-walled sep a and a e o en embedded in a mucoid
shea h. The excep ion is A. hyalina desc ibed wi h hyaline conidia ha become
subhyaline o pale b own a ma u i y (Ma e al. 2022). Based on molecula e i-
dence, Pleu oph agmium bambusinum (Dai e al. 2016) has been ans e ed o
Aquap e idospo a by Bao e al. (2021). In addi ion, se e al o he Pleu oph ag-
mium species o unce ain s a us (see below) and which lack molecula da a,
show close mo phological simila i y wi h membe s o his genus.
Pleu o heciella (Réblo á e al. 2012) is ano he genus wa an ing compa -
ison wi h Pleu oph agmium. Despi e hei close mo phological esemblance
in asexual cha ac e is ics, which makes hem di icul o dis inguish, he wo
gene a a e clea ly sepa a ed phylogene ically. Pleu o heciella belongs o he
Pleu o heciales, whe e i o ms a obus and species- ich lineage.
Species accep ed in Pleu oph agmium s. s .
He e we lis 14 species cu en ly accep ed in Pleu oph agmium s. s . whose
placemen is suppo ed by molecula da a. These species a e mo phologically
e y simila , wi h conidial cha ac e s p o iding he p ima y basis o dis inc-
ion. Se e al species o Pleu oph agmium a e known only om cul u e, wi h no
con i med wild ype ( he ypical o m o a species as i occu s in na u e). This
eliance on in i o obse a ions may complica e iden i ica ion and lead o dis-
o ion o key ai s, i.e. dis ibu ion o e ile egions on he conidiopho e (api-
cally o in nodulose swellings), conidiopho e appea ance, conidial dimensions
and sep a ion pa e ns, and e en he p esence o a mucoid shea h. In cul u e,
he mo phology o conidiopho es and conidia may de ia e om ha obse ed
in he wild ype (Réblo á e al. 2021, 2022). In addi ion, he conidial size o spe-
cies cu en ly e e ed o Pleu oph agmium s. s . show conside able o e lap,
making i di icul o dis inguish species based solely on conidial dimensions.
Pleu oph agmium asia icum (C ous) Réblo á & He n.-Res ., comb. no .
MycoBank No: 860799
Basionym. Neomy mec idium asia icum C ous, Pe soonia 41: 291. 2018.
Typus. THAILAND • Ra chabu i P o ince; on lea es o uniden i ied ine;
2008; P. W. C ous HPC 2252 (holo ype CBS H-23774, cul u e ex- ype CPC 34535
= CBS 145080).
32
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
8588 (PRA- 24034); • Olomouc Region, H ubý Jeseník M s., U Kříže o es
be ween Bělá pod P adědem and Vidly; on decaying wood o Fagus syl a ica;
8 Aug 1971; V. Holubo á-Jecho á 8584 (PRA-24035); • Ibid.; Bučina i gin
o es abo e F an iško a mysli na cabin nea Kou y nad Desnou; on decay-
ing wood o a b anch o Fagus syl a ica; 4 Aug 1971; V. Holubo á-Jecho á
8586 (PRA-24036). JAPAN • Kyo o, Dai okuji Tempel D; on uniden i ied dead
wigs; 28 Aug 1983; W. Gams & M. Tsuda (CBS H-3522, d ied cul u e CBS
H-3506, li ing cul u e CBS 770.83). SLOVAK REPUBLIC • Lúčanská Malá Fa-
a M s., Š ámko á Na ional Na u e Rese e, Kýče y; on decaying wood o
Fagus syl a ica; 26 Sep 1983; V. Holubo á-Jecho á 8556 (PRA-24037).
Habi a and geog aphical dis ibu ion. Sap obe on decaying wood and
he baceous s ems o a ious hos s, such as A c ium lappa, B assica ole acea,
Campanula medium, Ca pinus be ulus, Conium macula um, Epilobium hi su um,
Fagus syl a ica, Filipendula ulma ia, He acleum sphondylium, Li hoca pus edulis,
Polygonum sieboldii, Que cus sp., Sambucus ebulus, U ica dioica in he Belgium,
Czech Republic, Denma k, F ance, Ge many, Japan, Mexico, Ne he lands, New
Zealand, Slo ak Republic, UK and New Je sey and Washing on, USA (P euss
1852; Cos an in 1888; Be keley and B oome 1861; Ellis 1968; Holubo á-Jecho á
1972; Ma sushima 1975; Wang 2010; GBIF Sec e a ia 2023).
Acco ding o GlobalFungi da abase, P. pa ispo um was de ec ed in 40
samples. Mos eco ds o igina e om Asia, p ima ily om Japan and China,
wi h a smalle con ibu ion om Sou h Ko ea. Addi ional eco ds come om
Eu ope, all om Spain. The species is mos equen ly de ec ed in eshwa-
e aqua ic habi a s (45%), ollowed by an h opogenic/u ban si es (50%),
sh ubland (17.5%), o es (15%), and c opland (2.5%) biomes. I was de ec -
ed om wa e (45%), soil (35%), ai (17.5%), and dus (2.5%). These samples
we e collec ed ac oss a wide ele a ional ange, 26–948 m, sugges ing no
s ong es ic ion o high- o low-al i ude habi a s. Occu ences a e associ-
a ed wi h MAT ~15.4 °C and MAP ~1 059 mm/yea .
No es. Pleu oph agmium was o iginally desc ibed wi h a single species,
P. bicolo , wi h he p o ologue and accompanying illus a ion (Cos an in 1888:
ig. 70) ep oduced he e in Fig. 6. The species was ea ed in de ail unde
he synonymous names P. simplex by Ellis (1968), and P. pa ispo um by
Holubo á-Jecho á (1972) and Ma sushima (1975). These au ho s epo ed
a ela i ely b oad conidial leng h ange based on mul iple collec ions om
wood and he baceous s ems: 10–21 µm (mos ly 15.2 µm) × 3.5–6 µm (mos ly
4.5 µm) (Ellis 1968), and 12.5–22.5 (mos ly 15–18.5 µm) × 3.5–6.5 µm (mos ly
5 µm) (Holubo á-Jecho á 1972); o Ma sushima’s measu emen s, see
below. Collec ions om he Czech Republic and Japan examined in his s udy
a e consis en wi h hese accoun s, al hough he maximum conidial leng h
obse ed was sligh ly sho e han in he published anges. Ou obse a ions
con i m ha wi hin a single species, conidial dimensions can be a iable and
a e ully cong uen wi h p e ious desc ip ions.
In he Japanese ma e ial used in his s udy, conidial dimensions obse ed
bo h in cul u e (CBS 770.83: 9.5–14 × 3–4.5 µm) and on he na u al subs a e
(CBS H-3522: 10–13.5 × 3.5–4.5 µm) we e highly simila and all wi hin he ange
o a iabili y epo ed o his species om Eu ope (Ellis 1968; Holubo á-Jecho á
1972; his s udy). Al hough conidia om he Japanese ma e ial we e somewha
sho e han hose o he Czech specimens (10–18 × 3.5–6 µm on he na u al
33
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Figu e 8. Pleu oph agmium pa ispo um (PRA-24032). A Spo ula ing conidiopho es B–F conidiopho es wi h conidiog-
enous cells and conidia G Basal cell H–L conidia. Images: on na u al subs a e. Scale ba s: 300 µm (A); 20 µm (B, C);
10 µm (D–L).
34
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
subs a e, his s udy), hei o e all mo phology is indis inguishable, including he
sub le cha ac e s such as he o namen a ion o he ou e conidial wall due o
collapsing shea h, a ea u e no p e iously epo ed in he li e a u e bu clea ly
isible unde DIC and PHC mic oscopy (Figs 7, 8). No ably, Ma sushima (1975)
epo ed h ee Japanese collec ions o P. pa ispo um om decaying wood o
Li hoca pus edulis and Que cus sp., wi h a ying conidial sizes in cul u e: 14.5–
25 × 4.5–6.5 µm and 10–16 × 4–6 µm. We he e o e concluded ha CBS 770.83
and he Czech ma e ial o P. pa ispo um lis ed abo e a e conspeci ic and ep e-
sen a eliable e e ence o his species. A emp s o ex ac DNA om he six
he ba ium specimens om he Czech Republic we e unsuccess ul.
The ITS dis ibu ion da a de i ed om he GlobalFungi da abase sugges ha
P. pa ispo um has a b oad ecological ampli ude (Fig. 5), occu ing ac oss di e se
biomes and clima ic condi ions, bu wi h a no able concen a ion in aqua ic en i-
onmen s in Eas Asia (China, Japan). Reco ds om Spain con i m he dis ibu-
ion o he species in Eu ope, al hough based on published da a (see abo e) he
species has been epea edly epo ed om se e al Eu opean coun ies. Reco ds
om No h Ame ica, om whe e he species was also epo ed (Wang 2010), and
o he pa s o he wo ld (GBIF Sec e a ia 2023), a e absen in he GlobalFungi da-
abase. This b oad ecological and clima ic ange may indica e genuine ecological
e sa ili y, o al e na i ely, he exis ence o a species complex, as sugges ed by he
a iabili y in conidial leng h and occu ence on bo h he baceous and woody hos s.
To cla i y i s axonomy, molecula da a om addi ional s ains collec ed ac oss
di e se geog aphical egions a e essen ial o s abilising he species concep .
Among Pleu oph agmium species, P. pa ispo um is mos compa able o
P. lu iale, P. luguense, and P. sep a um in i s conidial mo phology. These
axa sha e obo oid o usoid-ellipsoid conidia ha a e b oadly ounded a
he apex, ape ing o a na ow base wi h a hilum, p edominan ly 3-sep a e,
and hyaline o subhyaline, becoming pale b own a ma u i y. In he absence
o molecula da a, dis inguishing among hese species would be highly
challenging. Fo de ailed compa ison, see Table 3.
In e es ingly, se e al biologically ac i e compounds, collec i ely known as
dac yl ungins, we e isola ed om P. pa ispo um (s ain D500 ob ained om
a dead lea in Japan) and we e epo ed o exhibi an i ungal ac i i y agains
Candida pseudo opicalis and o he ungi (Xaio e al. 1993). Howe e , hese
indings should be in e p e ed wi h cau ion due o he simple mo phology o he
species and he ecen ly ecognised axonomic ambigui y among p ese ed
s ains iden i ied as P. pa ispo um.
Pleu oph agmium p e idophy ophilum (Jing Y. Zhang, K.D. Hyde & Y.Z. Lu)
Réblo á & He n.-Res ., comb. no .
MycoBank No: 860808
Basionym. Neomy mec idium p e idophy ophilum Jing Y. Zhang, K.D. Hyde &
Y.Z. Lu, Fungal Di e si y 132: 375. 2025.
Typus. CHINA • Guizhou P o ince, Zunyi Ci y, Chishui Coun y, Hushi Town,
Chishui Alsophila Na u al Rese e; on dead ond s alks o P e idaceae sp. in
e es ial habi a s; 28 Jul 2022; J. Y. Zhang BL9 (holo ype GZAAS 23-0664, ex-
ype cul u e KUNCC 23-13858).
35
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Pleu oph agmium sep a um (C ous) Réblo á & He n.-Res ., comb. no .
MycoBank No: 860809
Basionym. Neomy mec idium sep a um C ous, Pe soonia 41: 287. 2018.
Typus. THAILAND • Ra chabu i P o ince; on lea es o uniden i ied ine;
2008, P. W. C ous HPC 2252 (holo ype CBS H-23768, cul u e ex- ype CPC 34585
= CBS 145073).
Pleu oph agmium sichuanense (Yan P. Chen & Maha achch.) Réblo á &
He n.-Res ., comb. no .
MycoBank No: 860810
Basionym. Neomy mec idium sichuanense Yan P. Chen & Maha achch., Phy o-
axa 701(2): 134. 2025.
Typus. CHINA • Sichuan P o ince, Chengdu Ci y, Dujiangyan Ci y, Longchi
Na ional Fo es Pa k; ele a ion 702 m a.s.l.; 31°00.18'N, 103°38.77'E; on decay-
ing b anches o an uniden i ied he baceous plan ; 5 Oc 2021; Y. P. Chen & W. H.
Tian LC64 (holo ype HUEST 24.0068).
No es. The e is a disc epancy be ween he p o ologue and he accompany-
ing igu es o P. sichuanense (Chen e al. 2025: ig. 3). While he conidia we e
desc ibed as subhyaline o pale b own, 0–1-sep a e, and inely e ucose,
hese ea u es a e no clea in he o iginal image. The conidia appea hyaline,
smoo h, and 1–3-sep a e in he igu es. Species wi h o namen ed conidia
a e ela i ely a e in Pleu oph agmium and cu en ly include P. pa ispo um,
P. sichuanense along wi h wo o he species: P. cla a um (Ma e al. 2014) and
P. e uculosum (Tiwa i 1969).
Pleu oph agmium so bicola (C ous & R.K. Schumach.) Réblo á & He n.-
Res ., comb. no .
MycoBank No: 860811
Basionym. My mec idium so bicola C ous & R.K. Schumach., Fungal Sys . E ol.
1: 191. 2018.
Synonym. Neomy mec idium so bicola (C ous & R.K. Schumach.) C ous, Pe -
soonia 41: 287. 2018.
Typus. GERMANY • nea Be lin; on b anch o So bus aucupa ia; 17 Feb 2016; R.
K. Schumache (holo ype CBS H-23405, cul u e ex- ype CPC 30455 = CBS 143433).
Pleu oph agmium s. la .: species o unce ain s a us
Al oge he , 19 species and a ie ies a e ecognised as membe s o Pleu oph agmi-
um s. la . in his s udy. This sec ion comp ises a mo phologically he e ogeneous
assemblage o species his o ically desc ibed in Pleu oph agmium, bu o which
molecula da a a e lacking. While some species con o m o he gene ic concep
and emain plausible candida es o inclusion in Pleu oph agmium s. s ., o he s
di e ge conside ably om i . No ably, only h ee species p oduce hyaline o sub-
hyaline conidia, occasionally appea ing uscous in mass, whe eas he emaining
36
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
axa bea pale b own, yellowish b own, b own, o da k b own conidia, o en wi h
dis inc ly pale end cell(s). To cap u e his a iabili y and highligh inconsis encies
in mo phological cha ac e s, diagnos ic ea u es o Pleu oph agmium s. la . spe-
cies cu en ly e ained in he genus a e summa ised in Table 3. Howe e , he ue
a ini ies o hese axa canno be eliably assessed un il molecula da a become
a ailable, emphasising he need o a ge ed ecollec ion and sequencing.
We p o ide de ails on he holo ype o acili a e u u e ecollec ion o hese
species by ellow mycologis s.
Pleu oph agmium angamosense Ma sush., Ma shushima Mycol. Mem. 8: 30.
1995.
Typus. PERU • Colonia Angamos; on he decayed pe iole o palm; Jul 1994; T.
Ma sushima (holo ype MFC-4P738).
No es. The species di e s om he Pleu oph agmium concep p ima ily in he
mo phology o i s conidiogenous cells and conidia. Al hough he cells p oli e a e
sympodially, he conidiogenous loci a e conspicuous, appea ing as dis inc sca s
3–3.5 µm wide. Conidia a e usi o m, ans e sely sep a e, smoo h, and b own,
wi h he e minal cells no iceably pale , and wi h da k, hick-walled sep a. The
o iginal illus a ion by Ma sushima (1995) u he sugges s ha conidial seces-
sion may be hexoly ic, as emnan s o he ou e wall appea o emain a ached
bo h o he conidiogenous locus and o he basal pa o he conidium. These
ea u es clea ly con as wi h Pleu oph agmium s. s ., in which conidiogenous
loci a e educed o minu e den icles and conidial secession is schizoly ic.
Pleu oph agmium aqua icum R.F. Cas añeda, He edia & R.M. A ias,
Myco axon 101: 92. 2007.
Typus. MEXICO • Ve ac uz, “Los Tux las”; on decaying wood subme ged in a
s eam; 19 May 2002; R. M. A ias & J. Y. C. Elizondo (holo ype XAL CB743, iso-
ype: MUCL 45625).
No es. Conidia o med on holoblas ic-den icula e conidiogenous cells a e
usi o m o cla a e, occasionally na icula , and o en sub-umbona e a he
apex. They a e b own, becoming pale b own o subhyaline owa d he ends and
a ound he sep a. By hei shape and pigmen a ion, he conidia closely esem-
ble hose o P. miniumbona um (He edia e al. 2007). Pleu oph agmium aqua -
icum also esembles P. usi o me in ha ing usi o m, pigmen ed conidia wi h
pale ends, bu he la e species di e s in ha ing na owe and uni o mly pale
b own conidia excep he ends (wi hou he pale bands a he sep a).
Pleu oph agmium bi unica um Ma sush., Icon. mic o ung. Ma sush. lec .:
115. 1975.
Typus. JAPAN • Tokyo P e ec u e, Hachijo Island; on decaying lea es o uniden-
i ied dico yledon plan ; Feb 1969; T. Ma sushima (holo ype MFC-1669).
37
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
No es. The species is ema kably simila o membe s o Aquap e idospo a
(Yang e al. 2015). Ano he species, P. bambusinum, was ecen ly ans e ed o
Aquap e idospo a based on molecula da a (Bao e al. 2021).
Pleu oph agmium cla a um L.G. Ma & X.G. Zhang, Myco axon 127: 216. 2014.
Typus. CHINA • Yunnan P o ince, he Fo bidden Fo es o Banna; on dead
b anches o Beilschmiedia pe co iacea (Lau aceae); 31 Oc 2011; L. G. Ma (ho-
lo ype HSAUP H2090, iso ype HMAS 243416).
No es. Ma e al. (2014) desc ibed h ee mo phologically simila species,
P. cla a um, P. ellipsoideum, and P. yunnanense, cha ac e ised by yellow-
ish-b own o pale b own conidia and conidiopho es ha o en e mina e in a
den icula e achis and ha e dis inc nodulose swellings along he whole leng h.
These swellings, howe e , a e no ypical o Pleu oph agmium based on ob-
se a ions o species on na u al subs a es, whe e he e ile egion o he
conidiopho e is es ic ed o i s uppe pa and bea s e minal and in e cala-
y conidiogenous cells. In cul u e, by con as , we equen ly obse ed such
swellings in P. pa ispo um. Conidia o P. cla a um a e consis en ly asep a e,
whe eas in Pleu oph agmium conidia a e in a iably sep a e, wi h bo h asep a e
and mul i-sep a e o ms occu ing wi hin a single species. Molecula da a a e
he e o e needed o con i m he placemen o hese species in he genus.
Pleu oph agmium ellipsoideum L.G. Ma & X.G. Zhang, Myco axon 127: 214.
2014.
Typus. CHINA • Yunnan P o ince, he Fo bidden Fo es o Banna; on dead
b anches o Bauhinia acumina a L. (Caesalpiniaceae); 17 Oc 2008; L. G. Ma
(holo ype HSAUP H0042, iso ype HMAS 243411).
No es. See no es unde P. cla a um.
Pleu oph agmium indicum D’Souza & Bha , Myco axon 119: 477. 2012.
Typus. INDIA • Goa, Molem Wildli e Sanc ua y; on allen dead and decaying
lea es o Dend ocalamus s ic us (Poaceae); 11 Ma 1999; M. A. D’Souza
(holo ype GUBH 367).
No es. The species is cha ac e ised by dis inc ly e sicolo ous conidia, wi h
he middle cells da k b own, he end cells pale b own, and he sep a conspicu-
ously hick-walled and da k b own. In o e all mo phology, i bea s esemblance
o membe s o Aquap e idospo a (Yang e al. 2015).
Pleu oph agmium ha unganae Hans ., Mycol. Pap. 15: 211. 1946.
Typus. UGANDA • En ebbe Road, on so i o Hemileia on lea es o Ha ungana
madagasca iensis; C. G. Hans o d 2803, 3013.
38
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
No es. The species is pa asi ic on he so i o Hemileia (Pucciniales). I was
desc ibed as ha ing dema iaceous conidiopho es wi h a e minal den icula e
achis, accompanied by addi ional, disconnec ed e ile egions along he con-
idiopho e axis. Conidia a e asep a e, cu ed, usi o m, and subhyaline o pale
oli aceous. Based on hese ea u es, he species appea s mo e consis en wi h
mo phologically simila gene a such as Ramichlo idium o My mec idium, pend-
ing ecollec ion and he acquisi ion o molecula da a o con i m i s placemen .
Pleu oph agmium malaysianum Ma sush., Ma sushima Mycol. Mem. 9: 20.
1996.
Typus. MALAYSIA • Selango Da ul Ehsan, Ulu Gombak, The Uni e si y o Mala-
ya Field S usy Cen e; on decaying lea es o a deciduous ee; 12 Jun 1995; T.
Ma sushima (holo ype MFC-5T054).
No es. The species is cha ac e ised by cylind ical o cla a e, pseudosep a e
conidia (Ma sushima 1996), ea u es ha ha e no p e iously been obse ed in
membe s o Pleu oph agmium s. s .
Pleu oph agmium miniumbona um (R.F. Cas añeda, I u . & Gua o) R.F.
Cas añeda, Myco axon 101: 96. 2007.
Basionym. Co dana miniumbona a R.F. Cas añeda, I u . & Gua o, Myco axon
73: 5. 1999.
Typus. VENEZUELA • Es ado de A agua, Pa que Nacional “Hen y Pi ie ”,
Es ación Rancho G ande, Camino de In e p e ación de la Na u aleza “Andy
Fields”, in undis u bed ain o es ; on allen decaying lea es on an uniden i ied
plan ; 25 No 1997; R. F. Cas añeda & T. I u iaga (holo ype MUCL 40700).
No es. He edia e al. (2007) ans e ed C. miniumbona a o Pleu oph agmi-
um based on i s supe icial esemblance o P. aqua icum (in he same s udy)
and he p esence o conidiopho es wi h a e minal den icula e e ile egion,
a ea u e mo e consis en wi h Pleu oph agmium han wi h Co dana (P euss
1851; He nández-Res epo e al. 2014). The conidia o P. miniumbona um a e
da k b own, excep o he hyaline basal cell, and exhibi an umbona e apex
wi h a iable shapes anging om obo oid and py i o m o b oadly cla a e.
Howe e , such da k b own conidia a e a ypical o Pleu oph agmium unde i s
cu en ci cumsc ip ion.
Pleu oph agmium na iculi o me Ma sush., Icon. mic o ung. Ma sush. lec .:
115. 1975.
Typus. JAPAN • Okinawa P e ec u e, I iomo e island; on decaying lea es o an
uniden i ied deciduous ee; Feb 1972; T. Ma sushima (holo ype MFC-4355).
No es. The species is cha ac e ised by 1-sep a e, na icula conidia acu e
a he apex and emain hyaline, bo h indi idually and in mass. The e ile
39
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
egion o he conidiopho e is apical, ela i ely small, and o en appea s
in la ed and cu ed. Mo phologically, he species esembles P. na icula e
(Yang e al. 2023), bu he la e di e s in possessing a longe e ile egion
and smalle , e ucose conidia ha become pale b own wi h hyaline end
cells a ma u i y. Molecula da a a e needed o de e mine whe he P. na icu-
li o me ep esen s a species belonging o a dis inc genus o alls wi hin he
in e speci ic a iabili y o Pleu oph agmium.
Pleu oph agmium obcampanuloides Ma sush., Ma sushima Mycol. Mem. 8:
29. 1995.
Typus. JAPAN • Okinawa P e ec u e, I iomo e island; in li e on he o es loo ;
1992; T. Ma sushima (holo ype MFC-2J039).
No es. Due o i s unique obpy i o m o u bina e conidia, which a e sep a e,
pale oli aceous, and oli aceous in mass (Ma sushima 1995), his species is
eadily dis inguished om o he membe s o he genus.
Pleu oph agmium pe uamazonicum Ma sush., Ma sushima Mycol. Mem. 7:
61. 1993.
Typus. PERU • Lo e o, o es Rio Neg o; on decaying palm lea es; Jun 1992; T.
Ma sushima (holo ype MFC-2P079).
No es. The species is cha ac e ised by e sicolo ous, 3-celled conidia wi h
a b own median cell and hyaline end cells. I s conidiopho es bea an apical
e ile egion ha o en ex ends along he uppe hal o he conidiopho e axis,
wi h den icles spa sely dis ibu ed (Ma sushima 1993). Mo phologically, i is
s ikingly simila o P. icolo (nom. in al., A . 40.1; Rambelli 2009); o a
de ailed compa ison, see Table 3. While P. pe uamazonensis was desc ibed
om cul u e, measu emen s o P. icolo we e de i ed om na u al ma e ial.
Consequen ly, ea u es ha dis inguish he wo axa, such as conidial size
and he ex en o he apical e ile egion, may simply e lec di e ences be-
ween in i o and in i o obse a ions.
Pleu oph agmium pe uamazonicum a . in la um Ma sush., Ma sushima
Mycol. Mem. 7: 61. 1993.
Typus. PERU • Lo e o, o es Rio Neg o; on decaying lea es o a deciduous ee;
No 1990; T. Ma sushima (holo ype MFC-0P570).
No es. The a ie y in la um di e s om he ype a ie y by he conspic-
uous in la ion o he basal po ion o he conidium (Ma sushima 1993).
I is u he cha ac e ised by hick-walled sep a, wi h he wo basal cells
pale b own. Howe e , i emains unce ain whe he hese dis inc ions all
wi hin he na u al in aspeci ic a iabili y o he species o ep esen a
sepa a e species.
40
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Pleu oph agmium sub usi o me Ma sush., Icon. Mic o ung. Ma sush. lec .:
116. 1975.
Typus. JAPAN • Mie P e ec u e, Uni e si y o Mie, Hi aku a Exp. Fo es ; o es ;
Sep 1965; T. Ma sushima (holo ype MFC-1551).
No es. Pleu oph agmium sub usi o me is eadily dis inguished om o he
species o he genus by i s la ge (21–38 × 5.8–9.2 µm), usi o m conidia, which
a e 3–7-sep a e and some imes sligh ly cons ic ed a he sep a. The conidia
a e subhyaline, becoming pale uscous in mass (Ma sushima 1975).
Pleu oph agmium aiwanense Ma sush., Ma sushima Mycol. Mem. 5: 24. 1987.
Synonym. Pleu oph agmium bicolo Ma sush., Icon. mic o ung. Ma sush. lec .:
114. 1975. Nom. illegi . (ICN, A . 53.1) non Pleu oph agmium bicolo Cos an in,
Cos an in, Mucéd. Simpl.: 100. 1888.
Typus. TAIWAN • Ken-Ting Pa k; on o en lea - achis o A enga engle i; 27
May 1980; T. Ma sushima (holo ype MFC-10120).
No es. This species p oduces cylind o-cla a e o somewha u bina e
conidia wi h a sligh ly in la ed apical cell. Obse a ions om cul u e show ha
he e ile egions o he conidiopho e a e discon inuous, occu ing apically as
well as a se e al poin s along he conidiopho e axis, some imes associa ed
wi h nodose swellings (Ma sushima 1987). Mo phologically, i esembles
P. pe uamazonicum bu can be dis inguished by i s sho e conidia and ewe
sep a. Fo addi ional compa ison, see no es o P. asia icum.
Pleu oph agmium icolo Rambelli, Flo a Medi e anea 19: 82. 2009. Nom.
in al. (ICN, A . 40.1).
Typus. ITALY • Pan elle ia, Mon agna G ande; on dead lea es o A bu us unedo
(a7); (holo ype PAL).
Pleu oph agmium a iesep a um Ma sush., Icon. mic o ung. Ma sush. lec .:
117. 1975.
Typus. JAPAN • Kyo o Ci y; on decaying s em o Phyllos achys edulis; Feb 1966;
T. Ma sushima (holo ype MFC-1964).
No es. The species is eadily dis inguished by i s cylind ical, pale oli aceous
conidia, which display conside able a iabili y in bo h size and sep a ion, ang-
ing om one o ou sep a (Ma sushima 1975).
Pleu oph agmium e uculosum D.P. Tiwa i, Indian Phy opa h.: 513. 1970.
Typus. INDIA • Madhya P adesh, Saga ; hizosphe e soil o Pipe be le; Dec
1965; D. P. Tiwa i (holo ype IMI 134426, iso ype ITCC New Delhi 1375).
41
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
No es. Ve ucose conidia a e a e in Pleu oph agmium and a e known only
om a ew species, namely P. cla a um, P. sichuanense, and P. e uculosum. O -
namen ed conidia a e also obse ed in P. p e idophy ophilum as “ ough-walled”
(Zhang e al. 2025), and in P. pa ispo um ( his s udy), whe e he conidial wall
appea s somewha longi udinally c umpled. Among hese axa, molecula da a
a e cu en ly a ailable only o P. sichuanense and P. pa ispo um. Mo phologi-
cally, P. e uculosum is mos simila o P. cla a um, bu can be dis inguished by
i s sep a e conidia and sho e conidiopho es.
Pleu oph agmium yunnanense L.G. Ma & X.G. Zhang, Myco axon 127: 215.
2014.
Typus. CHINA • Yunnan P o ince, he Fo bidden Fo es o Banna; on dead
b anches o Machilus salicina (Lau aceae); 18 Oc 2008; L. G. Ma (holo ype
HSAUP H0086, iso ype HMAS 243412).
No es. The species p oduces pale b own, b oadly usi o m conidia ape -
ing a bo h ends, which a e mos ly 3-sep a e (Ma e al. 2014). In conidial size
and sep a ion, i esembles P. ellipsoideum om he same s udy, bu he la e
di e s in ha ing ellipsoidal conidia ha a e b oadly ounded a he apex. Fo
addi ional in o ma ion, see he no es unde P. cla a um.
Species excluded om Pleu oph agmium and desc ibed in o he gene a
He e, we p o ide a lis o species ha ha e been excluded om Pleu oph agmi-
um and subsequen ly ans e ed o o he gene a by a ious au ho s. Synonymy
ollows he eco ds in MycoBank, unless s a ed o he wise.
Aquap e idospo a bambusinum (D.Q. Dai & K.D. Hyde) D.F. Bao, J. Fungi
7(669): 10. 2021.
Basionym. Pleu oph agmium bambusinum D.Q. Dai & K.D. Hyde, Fungal Di e -
si y 82: 92. 2016.
Dac yla ia a ecae (Ma sush.) R.F. Cas añeda & W.B. Kend ., Uni . Wa e loo,
Biol. Se . 35: 26. 1991.
Basionym. Pleu oph agmium a ecae Ma sush., Ma sushima Mycol. Mem.
5: 23. 1987.
Dac yla ia cylind ospo a (Ma sush.) R.F. Cas añeda & W.B. Kend ., Uni .
Wa e loo, Biol. Se . 35: 27. 1991.
Basionym. Pleu oph agmium cylind ospo um Ma sush., Icon. mic o ung.
Ma sush. lec .: 115. 1975.
48
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
unb anched, some imes pe cu en ly elonga ing, lexuous o sinuous, some imes
becoming genicula e exhibi ing a zig-zag pa e n due o local bending abo e he
sep um, each bend is associa ed wi h he o ma ion o a single den icle on he
‘ou side’ gi ing he appea ance o i egula o dicho omous b anching, pale b own
o golden b own, smoo h-walled, sep a e. Conidiogenous cells 10–24 × 3–4 µm,
in eg a ed, e minal, o m ans e se sep a du ing sympodial ex ension and be-
come in e cala y, occasionally la e al g owing di ec ly on hyphae, monoblas ic o
polyblas ic wi h 1–4 peg-like den icles, cylind ical o subula e, subhyaline o pale
oli aceous-b own when in he e minal posi ion, golden-b own when in e cala y,
smoo h-walled; conidiogenesis holoblas ic-den icula e. Conidia 18.5–36(–45) ×
(3–)4–5.5 µm (mean ± SD = 27.1 ± 5.6 × 4.9 ± 0.1 μm), soli a y, d y, ac opleu og-
enous, oblong o cylind ical, elonga e usi o m o na owly ellipsoidal, ape ing a
bo h ends, unca e a he base 1–1.5 µm wide, wi h a conspicuous basal sca ,
usually s aigh , occasionally sligh ly cu ed, o en wi h gu ules o g anules is-
ible inside he cells, smoo h-walled, asep a e and hyaline when young, a ma u-
i y wi h 3–7 ans e se sep a and subhyaline o pale oli aceous-b own, oli a-
ceous-g ey in mass; conidial secession schizoly ic.
Habi a and geog aphical dis ibu ion. Skoliomycella la a is a sap obe oc-
cu ing on plan emnan s. To da e, wo con i med eco ds o igina e om he
Ibe ian Peninsula, speci ically om Po ugal ( his s udy) and Spain (GlobalFun-
gi), wi hin a empe a e, Medi e anean clima e. Acco ding o GlobalFungi, i was
de ec ed in a single ai sample om an an h opogenic biome, MAT ~14.5 °C,
MAP ~575 mm/yea .
No es. Skoliomycella la a is eadily dis inguished om he mo phologically
simila species a ibu ed o Campo esiomyces (Hyde e al. 2020) and Zaaneno-
myces (C ous e al. 2021) by he absence o a achis in he e ile apical po ion
o he conidiogenous cell. Ins ead, i s conidiopho es a e lexuous o sinuous,
some imes exhibi a zig-zag pa e n, wi h a single den icle and/o a sligh ly
p olonged conidiogenous cell o med a each bend on he ou e side o he co-
nidiopho e, gi ing he imp ession o i egula o dicho omous b anching.
The de ec ion o S. la a in u ban ai highligh s i s capaci y o ae ial dispe -
sal, al hough i occu s a e y low ela i e abundance. Howe e , i s global dis-
ibu ion emains unce ain, as no addi ional eco ds ou side he Ibe ian Penin-
sula a e a ailable. Whe he S. la a ep esen s a a e, geog aphically es ic ed
lineage, o whe he i s sca ci y e lec s limi ed sampling, o unde -de ec ion
due o low en i onmen al abundance, emains o be de e mined.
Thysano ea A zanlou, W. Gams & C ous, S ud. Mycol. 58: 80. 2007. emend.
He n.-Res . & C ous, Fungal Sys . E ol. 6: 17. 2020.
Desc ip ion. See A zanlou e al. (2007) and He nández-Res epo e al. (2020).
No es. Thysano ea is cha ac e ised by mic o- o mac onema ous, dema i-
aceous conidiopho es ha a e simple o apically b anched, bea ing e minal
o in e cala y, holoblas ic, polyblas ic conidiogenous cells. Conidia a e soli a y,
ac opleu ogenous, ans e sely sep a e, pale b own, and a iable in shape,
mos commonly oblong, obo oid, o usi o m. In con as , he associa ed syna-
sexual mo ph is phialidic.
49
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Th ee species p e iously classi ied in Uncispo a we e shown o be conge-
ne ic wi h Thysano ea and a e he e o mally ans e ed o he genus, wi h new
combina ions p oposed below.
Thysano ea ac opleu ogena Réblo á & He n.-Res ., sp. no .
MycoBank No: 860719
Fig. 10
E ymology. F om G eek ak os (apex), pleu á (side) and genēs (bo n, p oduced).
Re e ing o he mode o conidium de elopmen , in which conidia a e o med
bo h e minally and la e ally on he conidiogenous cells.
Typus. PAPUA NEW GUINEA • Madang P o ince, oo hills o Finis e e
Range, 40.8 km along oad o Lae; 200 m a.s.l.; on uniden i ied wood; 2 No
1995; A. Ap oo 36665 (holo ype CBS H-25780 d ied cul u e, ex- ype cul u e
CBS 215.96).
Cul u e cha ac e is ics. On CMD colonies 44–45 mm diam., ci cula , la ,
ma gin en i e, lanose, pinkish-b own o b own, da ke a he ma gin, e e se
da k b own. On MLA colonies 36–38 mm diam., ci cula , con ex, ma gin en-
i e, loccose o lanose, oli aceous-g ey o mouse g ey, da k oli aceous a he
ma gin, e e se da k oli aceous. On OA colonies 40–41 mm diam., ci cula , la ,
ma gin en i e, lanose a he cen e and on he inocula ion block, cobwebby o-
wa ds he pe iphe y, wi h a well-de ined g ey-b own cen al zone, da k b own
owa ds he pe iphe y, wi h a di use ligh e halo a he ma gin, e e se da k
b own. On PCA colonies 40–42 mm diam., ci cula , la , ma gin en i e, lanose,
g ey-b own, cobwebby and da k b own owa ds he pe iphe y, e e se da k
b own. Wi h a p ominen subme ged g ow h on all media. Spo ula ion on PCA,
absen on CMD, MLA and OA.
Desc ip ion in cul u e. Colonies on PCA e use. Sexual mo ph. No ob-
se ed. Asexual mo ph. Mycelium composed o subhyaline o pale oli a-
ceous b own, sep a e hyphae, 1.5–2.5 µm wide. Conidiopho es 98–145 ×
3.5–4.5 µm, basal cells some imes sligh ly in la ed 5–6.5 µm wide, mac-
onema ous, mononema ous, loosely sca e ed o densely ascicula e, mos -
ly e ec , s aigh o sligh ly lexuous, cylind ical, sub ly undula e, unb anched
o loosely b anched in he uppe pa , pale o medium b own o pale oli a-
ceous b own, da ke a he base, smoo h-walled, sep a e. Conidiogenous
cells 23.5–28(–40) × 3.5–5(–5.5) µm, in eg a ed, e minal, o ming ans-
e se sep a du ing sympodial p oli e a ion and o en becoming in e cala y,
a anged ei he along mos o he conidiopho e leng h o es ic ed o he
uppe pa , o ming a achis wi h nume ous, closely spaced, minu e den icles,
polyblas ic, cylind ical, pale b own o pale oli aceous b own, end cells end
o be pale , smoo h-walled; conidiogenesis holoblas ic-den icula e. Conidia
(13.5–)15–21 × (3.5–)4–5.5 µm (mean ± SD = 17.2 ± 1.8 × 4.5 ± 0.3 μm),
soli a y, d y, ac opleu ogenous, subcylind ical o oblong o usi o m o u-
si o m-cla a e, ape ing owa ds bo h ends, unca e a he base 1–1.5 µm
wide, wi h a conspicuous basal sca , s aigh o sligh ly cu ed, pale b own
o pale oli aceous b own, da ke a he base, da k oli aceous b own in mass,
smoo h-walled, (1–)3-sep a e; conidial secession schizoly ic.
50
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Figu e 10. Thysano ea ac opleu ogena (ex- ype CBS 215.96). A Spo ula ing conidiopho es B, C uppe pa o he conid-
iopho e conidiogenous cells o ming a achis and a ached conidia D–G conidiopho es, conidiogenous cells and conidia
H conidia I di e si y o colony mo phology on CMD, MLA, OA, and PCA, espec i ely ( om le o igh ) a e 4 wk. Images:
on PCA (A–H). Scale ba s: 300 µm (A); 10 µm (B–H); 1 cm (I).
51
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Habi a and geog aphical dis ibu ion. The species is a sap obe occu ing
on decaying wood and is cu en ly known only om Papua New Guinea. No
iden ical ITS sequences we e eco e ed in he GlobalFungi da abase.
No es. Phylogene ic analyses (Fig. 2) e ealed ha T. ac opleu ogena o ms
a sis e ela ionship o a subclade comp ising h ee closely ela ed species,
T. hainanensis, T. sinensis, and T. wuzhishanensis. They belong o a subclade
ha also includes T. sei e ii and T. can elliae. Among known species, T. ac op-
leu ogena is closes o T. obscu a and T. sei e ii in ha ing ligh ly pigmen ed,
subcylind ical o oblong, p edominan ly 3-sep a e conidia. Howe e , bo h spe-
cies a e eadily dis inguishable in conidial cha ac e s and ha ing unb anched
conidiopho es. Thysano ea obscu a di e s om he new species in subhya-
line o yellowish b own, la ge conidia, 20–31 × 5–8 µm (Ma sushima 1983).
Thysano ea sei e ii p oduces simila pale b own, subcylind ical o cla a e o
oblong conidia, bu hese a e conside ably sho e , 7–15 × 1.5–3 µm (He nán-
dez-Res epo e al. 2020). In addi ion, T. sei e ii p oduces a phialidic synasex-
ual mo ph in cul u e, a ea u e so a unique among Thysano ea species. In he
phylogene ic ee, all h ee species a e esol ed as dis inc lineages.
The o iginal ma e ial, CBS H-6267, om which he axenic cul u e was de-
i ed, consis s o mul iple wigs. Howe e , we could no loca e he a ge un-
gus. Se e al wigs a e co e ed wi h ae ial mycelium, while o he s ca y e ile
conidiopho es o a dend yphiella-like ungus. Acco dingly, a d ied cul u e was
designa ed as he holo ype o T. ac opleu ogena.
Thysano ea ac opleu ogena appea s o ep esen a a e species, con i med
hus a om a single collec ion on decaying wood. The absence o iden ical
ITS sequences in he GlobalFungi da abase indica es ha i has no ye been de-
ec ed in en i onmen al sequencing da ase s, sugges ing ha he species may
be geog aphically es ic ed o cu en ly o e looked due o unde -sampling.
Thysano ea hainanensis (Jian. Y. Li & Z.F. Yu) Réblo á & He n.-Res ., comb. no .
MycoBank No: 860812
Basionym. Uncispo a hainanensis Jian. Y. Li & Z.F. Yu, Myco axon 129: 474. 2015.
Typus. CHINA • Hainan P o ince, Wuzhishan Na ional Na u e Rese e;
754 m a.s.l., isola ed om decayed lea es; Dec 2011; G. Z. Yang (holo ype YMF
1.04038, ex- ype cul u e YMF1.040381).
Thysano ea melanica (Hong Y. Su, Udayanga & K.D. Hyde) He n.-Res . &
C ous, Fungal Sys . E ol. 6: 18. 2020.
Basionym. Minimelanolocus melanicus Hong Y. Su, Udayanga & K.D. Hyde, Fun-
gal Biol. 119: 1056. 2015.
Cul u e cha ac e is ics. On CMD colonies 51–52 mm diam., ci cula , la o
sligh ly aised in he cen e, ma gin di use, en i e, zona e, loccose o el e y,
whi e-beige in he cen e, b own o da k b own owa ds he ma gin, e e se da k
b own o black. On MLA colonies 49–51 mm diam., ci cula , la , aised a he
cen e, ma gin en i e, zona e, lanose o loccose cen ally, becoming mucoid
and glossy owa ds he pe iphe y, lanose a he ma gin, c eam o bu in he
52
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
cen al egion, sha ply con as ing wi h he su ounding, da k g ey o blackish
mucoid mycelium, oli aceous-g ey a he ma gin, e e se da k oli aceous-g ey
o black. On OA colonies 47–50 mm diam., ci cula , la , ma gin di use, en i e,
loccose o sligh ly lanose a he cen e, becoming cobwebby owa ds he pe-
iphe y, whi ish o pale pinkish-bu , su ounded by a wide zone o subme ged,
da k oli aceous-g ey mycelium ha di uses in o he aga , pale a he ma gin,
e e se uni o mly da k oli aceous-g ey o black. On PCA colonies 41–42 mm
diam., ci cula , la o sligh ly aised in he cen e, ma gin di use, en i e, loccose
o somewha lanose, cen al zone pale pink-b own-bu , su ounded by a da k
b own subme ged zone, pale a he ma gin, e e se da k oli aceous-b own o
nea ly black. Spo ula ion absen on all media.
Desc ip ion in cul u e. Colonies on PCA e use. Sexual mo ph. No obse ed.
Asexual mo ph. Mycelium composed o subhyaline o pale b own, sep a e hy-
phae, 1.5–3 µm wide. Conidiopho es, conidiogenous cells and conidia absen .
Specimen examined. THE NETHERLANDS • No h Holland P o ince, Wi-
e inge mee Polde , Van Bemmelen Hoe e; isola ed om whea ield soil; May
1966; W. Gams (li ing cul u e CBS 862.68).
Habi a and geog aphical dis ibu ion. Thysano ea melanica was o iginally
desc ibed om decaying wood in China (Liu e al. 2015), wi h an addi ional
eco d om whea ield soil in he Ne he lands ( his s udy). Acco ding o he
GlobalFungi da abase, he species has been de ec ed in 266 en i onmen al
samples ac oss h ee con inen s. I s ITS da ase is s ongly Eu ocen ic, wi h
~90% o de ec ions o igina ing om Eu ope, whe e i is eco ded in mul iple
coun ies, showing no able ho spo s in Es onia, Swi ze land, and Ge many.
Asia con ibu es ~8% (d i en la gely by China), and No h Ame ica is spa se-
ly ep esen ed by ~2%. I is mos equen ly de ec ed in c opland (50%),
ollowed by g assland (20%), o es (17%), an h opogenic habi a s (9.4%),
woodland (2.3%) and sh ubland (0.4%) biomes. Mos eco ds a e om soil
samples including opsoil and hizosphe e soil (93.2%), wi h mino ep esen-
a ion in oo s and shoo s. Occu ences a e associa ed wi h MAT ~8.4 °C and
MAP ~749 mm/yea .
No es. Al hough s ain CBS 862.68 did no spo ula e on any o he cul u e
media es ed, molecula da a enabled i s placemen in he genus Thysano ea
and as conspeci ic wi h T. melanica (Fig. 2). Thysano ea melanica is nes ed
wi hin a well-suppo ed subclade comp ising six addi ional species. I is cha -
ac e ised by unb anched, da k b own conidiopho es bea ing e minal and in-
e cala y conidiogenous cells, and by pale b own conidia ha a e 1–3-sep a e
when young and 4–6-sep a e a ma u i y, measu ing (9–)13–37(–45) × (2.5–
)3.5–6.5(–8) μm (Liu e al. 2015).
Thysano ea melanica is a cosmopoli an, soil-dwelling sap obe, occasional-
ly associa ed wi h plan issues, ha h i es ac oss a wide ange o en i on-
men s wi h a p e e ence o empe a e o cool- empe a e clima es wi h mod-
e a e ain all. I s equen occu ence in c oplands, g asslands, and o es s
highligh s i s ecological e sa ili y and abili y o pe sis in bo h na u al and
an h opogenic ecosys ems. The p edominance o eco ds om c opland soils
u he implies ha ag icul u al ac i i ies may ha e acili a ed i s sp ead and
pe sis ence, po en ially h ough soil anspo o c op-associa ed dispe sal.
This hypo hesis is consis en wi h he occu ence o CBS 862.68 s ain, which
was isola ed om whea ield soil.
53
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Thysano ea sinensis (G.Z. Yang & Z.F. Yu) Réblo á & He n.-Res ., comb. no .
MycoBank No: 860813
Basionym. Uncispo a sinensis G.Z. Yang & Z.F. Yu, Myco axon 116: 172. 2011.
Typus. CHINA • Yunnan p o ince, Mengla Coun y, Xishuangbanna T opical
Bo anical Ga den; on subme ged lea es o an uniden i ied dico yledonous plan ;
Sep 2010; G. Z. Yang (holo ype YMF 1.03683; ex- ype cul u e YMF 1.03683).
Thysano ea wuzhishanensis (L.P. Chen & Z.F. Yu) Réblo á & He n.-Res .,
comb. no .
MycoBank No: 860814
Basionym. Uncispo a wuzhishanensis L.P. Chen & Z.F. Yu, Sydowia 70: 255. 2018.
Typus. CHINA • Hainan P o ince, Wuzhishan Na ional Na u e Rese e; 754
m a.s.l.; on subme ged decaying lea es in a s eam; 30 Jun 2011; Z. F. Yu (ho-
lo ype YMF 1.04080).
Wongia Khemmuk, Gee ing & R.G. Shi as, IMA Fungus 7: 249. 2016.
Desc ip ion. See Khemmuk e al. (2016).
No es. To da e, 11 species ha e been desc ibed in Wongia. The asexual
mo phs p oduce mac onema ous conidiopho es wi h holoblas ic, polyblas ic,
sympodially elonga ing conidiogenous cells ha gene a e sep a e, subhyaline
o da k b own, ans e sely sep a e o a ely asep a e conidia (e.g. Bao e al.
2021; C ous e al. 2022; Manawasinghe e al. 2024; Wang e al. 2025).
Wongia pallidopola is Réblo á & He n.-Res ., sp. no .
MycoBank No: 860720
Fig. 11
E ymology. F om La in pallidus (pale), and pola is (o o ela ing o he poles).
Re e ing o he conidial pigmen a ion, in which he apical and basal cells a e
dis inc ly pale han he cen al, mo e pigmen ed cells, c ea ing a no iceable
bipola con as .
Typus. THE NETHERLANDS • Gelde land P o ince, Wageningen; isola ed
om sandy soil unde con inuous whea ; Jan 1970; J. H. an Emden No. 4118,
30 (holo ype CBS H-25781 d ied cul u e, ex- ype cul u e CBS 440.70).
Cul u e cha ac e is ics. On CMD colonies 40–41 mm diam., ci cula , la ,
ma gin en i e o sligh ly imb ia e, di use, lanose, wi h a sub le concen ic zon-
ing, beige o g ey-beige a he cen e, b own a he ma gin, e e se da k b own.
On MLA colonies 38–40 mm diam., ci cula , aised, ma gin en i e, lanose, com-
posed o camel b own and pale oli aceous b own concen ic zones, e e se
da k b own. On OA colonies 49–51 mm diam., ci cula , aised, ma gin en i e,
lanose, composed o beige, camel b own, da k b own and cinnamon concen ic
zones, b own a he ma gin, ae ial hyphae a he cen e and ma gin bea ing nu-
me ous colou less exuda es, e e se b own. On PCA colonies 44–45 mm diam.,
54
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
ci cula , con ex, ma gin en i e, lanose, whi ish-g ey a he cen e su ounded
wi h a hin smoke-g ey zone, pale beige o ligh awn owa ds he ma gin, e-
e se da k b own. Spo ula ion abundan on CMD, MLA, and PCA, absen on OA.
Desc ip ion in cul u e. Colonies on MLA e use. Sexual mo ph. No ob-
se ed. Asexual mo ph. Mycelium composed o pale b own, sep a e, spa se-
ly b anched hyphae, 1.5–3 µm wide. Conidiopho es 24–70 × 3.5–5(–5.5) µm,
mac onema ous, mononema ous, soli a y o agg ega ed, e ec , s aigh o
sligh ly lexuous, apically almos sinuous, cylind ical, unb anched, occasionally
p oli e a ing sympodially, b own, da k b own in he lowe pa , smoo h-walled,
sep a e. Conidiogenous cells 12–30 × (4.5–)5–6 µm, in eg a ed, e minal,
mono- o polyblas ic, wi h one o se e al den icles, ex ending sympodially, cy-
lind ical, ape ing, some imes sligh ly swollen a he apex, pale b own, pale
a he apex, smoo h-walled; conidiogenesis holoblas ic-den icula e. Conidia
(20–)22–28(–30) × 5.5–6.5(–5.5) µm (mean ± SD = 25.2 ± 1.7 × 6.1 ± 0.3 μm),
soli a y, d y, ac opleu ogenous, ellipsoid o usi o m o usi o m-cla a e, ape -
ing owa ds bo h ends, unca e a he base 2–2.5 µm wide, wi h a conspicuous
basal sca , mos ly s aigh , occasionally sligh ly cu ed, b own o da k b own,
end cells pale hen he middle ones, apical cell o en wi h a da k b own ip,
smoo h-walled o inely oughened, hick-walled, he ou e wall pa ly de aches
om he conidium, he de ached segmen s appea as apical o side pocke o
wings, some imes he ou e wall is de ached a ound he base imi a ing a min-
u e ill, 3-sep a e, mucoid shea h absen ; conidial secession schizoly ic.
Habi a and geog aphical dis ibu ion. The examined s ain was isola ed
om sandy ag icul u al soil in he Ne he lands. Acco ding o he GlobalFun-
gi da abase, W. pallidopola is was de ec ed in 812 en i onmen al samples. I
is cosmopoli an, widely dis ibu ed in empe a e o sub opical egions, wi h
mos eco ds om No h Ame ica, Eu ope, and Asia. The main ho spo s a e
in he USA (Michigan, New Yo k and No h Ca olina), China (P o inces Fujian,
Guizhou, Hebei, Jiangxi, Jilin and Yunnan), and cen al Eu ope (pa icula ly
Swi ze land and he Ne he lands), while addi ional, less equen eco ds o igi-
na e om Aus alia and A ica. The species is p ima ily soil-associa ed (86.2%),
wi h occasional de ec ion in oo s, shoo s and a e occu ence in ai o wa-
e . I is p edominan ly associa ed wi h c opland (59.1%) and g assland (28%)
ecosys ems, ollowed by occasional occu ence in he an h opogenic habi a s
(6.4%), o es (4.2%) and woodland, aqua ic, sh ubland and we land biomes.
In bo h USA and China, W. pallidopola is is s ongly associa ed wi h c opland
ecosys ems, pa icula ly ce eals and legumes (Zea mays, Glycine max, O yza
sa i a, Chenopodium quinoa). Howe e , in China he species exhibi s a b oade
ecological ampli ude, occu ing no only in c oplands bu also in o es soils,
hizosphe es, and e en ae osols, whe eas in he USA i appea s o be mo e e-
s ic ed o c opland and an h opogenic soils. Occu ences a e associa ed wi h
MAT ~11.8 °C and MAP ~926 mm/yea .
No es. Wongia pallidopola is closely esembles W. aqua ica (Luo e al. 2019)
in ha ing 3-sep a e, b own conidia wi h pale end cells. Howe e , W. aqua ica
di e s in possessing sho e conidia, measu ing 17–21 × 5–7 µm. Phyloge-
ne ically, bo h species o m a s ongly suppo ed sis e ela ionship wi hin a
monophyle ic clade ha is basal o all emaining Wongia species (Fig. 3).
Mos conidia lacked any shea h-like s uc u e; howe e , in a ew cases,
a simila ea u e was obse ed (Fig. 11O). I appea ed on one o bo h sides
55
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Figu e 11. Wongia pallidopola is (ex- ype CBS 440.70). A Spo ula ing conidiopho es B–K conidiopho es, conidiogenous
cells and conidia L–O conidia P di e si y o colony mo phology on CMD, MLA, OA, and PCA, espec i ely ( om le o
igh ) a e 4 wk. Images: on MLA (A–O). Scale ba s: 300 µm (A); 10 µm (B–O); 1 cm (P).
56
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
nea he base o apex, only a he apex, o occasionally a se e al poin s on
he same conidium. This s uc u e is sligh ly pigmen ed and esembles
an ou e wall laye ha de aches in one o se e al places a he han a mu-
coid shea h. In o he Wongia species, a mucoid shea h has no been epo -
ed. I is possible ha in cul u e, whe e osmo ic condi ions di e om hose
in na u e, he ou e conidial wall de e io a es and pa ially de aches. In con-
as , a simila s uc u e obse ed in P. pa ispo um ep esen s a ue mu-
coid bu epheme al shea h ha is p ac ically in isible on conidia om na -
u al ma e ial ye clea ly isible in cul u e, posi ioned la e ally a ound he
middle o he conidium (Fig. 7N–P) o co e ing he uppe wo- hi ds (Fig. 7D, E).
Based on eDNA da a, W. pallidopola is is ega ded as a cosmopoli an soil
sap obe wi h s ong ecological associa ions o c opland and g assland ecosys-
ems. I s p e alence in ag icul u al soils sugges s ha human ac i i y, pa icu-
la ly h ough ag icul u al p ac ices, may ha e acili a ed i s dissemina ion. I s
occasional de ec ion in o es soils, hizosphe es, and e en ae osols indica es
ha i can exploi a b oade ange o habi a s. Despi e i s widesp ead occu -
ence in en i onmen al samples, W. pallidopola is ep esen s a mo phologically
c yp ic ungal lineage ha has likely been o e looked in adi ional su eys.
Wongia hachidopho a Réblo á & He n.-Res ., sp. no .
MycoBank No: 860721
Fig. 12
E ymology. F om G eek hachis (spine, axis) and pho os de i ed om pho a
(bea ing). Re e ing o he e ile, achis-like uppe pa o he conidiopho e,
which bea s conidia along i s leng h.
Typus. INDIA • Bangalo e, A bo e um o Fo es y Depa men ; on dead lea
o Bambusa sp.; Jun 1973; W. Gams (holo ype CBS H-11671 d ied cul u e, ex-
ype cul u e CBS 531.73, pa a ype CBS H-5414).
Cul u e cha ac e is ics. On CMD colonies 45–51 mm diam., ci cula , la ,
powde y, sienna, ma gin pale , cinnamon o awn, di use, imb ia e o sligh -
ly loba e, e e se b own wi h di e en ones o da k b ick, b ick o sienna in
concen ic zones. On MLA colonies 65–67 mm diam., ci cula , la , ma gin
en i e o imb ia e, el e y o powde y, cen e osy bu , cinnamon o sa on
owa ds he pe iphe y, e e se b own. On OA colonies 58–60 mm diam., ci cu-
la , la , ma gin en i e o imb ia e o loccose, el e y, wi h concen ic b own
zones o di e en ones such as ul ous, sienna o cinnamon, och eous o-
wa ds he ma gin, e e se da k b own (umbe ) in he cen e wi h och eous
ma gin. On PCA colonies 53–54 mm diam., ci cula , la , ma gin en i e im-
b ia e o di use, el e y o sandy, cen e ul ous o och eous, isabelline o-
wa ds he ma gin, wi h nume ous exuda es a he cen e, e e se o he same
colou . Spo ula ion abundan on all media.
Desc ip ion in cul u e. Colonies on OA e use. Sexual mo ph. No obse ed.
Asexual mo ph. Mycelium composed o hyaline o pale b own, smoo h hyphae,
e ucose close o he conidiopho e base, 1–2 μm wide. Conidiopho es up o
63 μm long, 2.5–3.5 μm wide a he base, mac onema ous, mononema ous,
soli a y, e ec , s aigh o sligh ly lexuous, cylind ical, unb anched, pale b own
o b own-o ange. Conidiogenous cells 13–52 × 3–4 μm, in eg a ed, e minal,
57
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Figu e 12. Wongia hachidopho a (ex- ype CBS 531.73). A–C Spo ula ing conidiopho es D–G conidiopho es, conidioge-
nous cells and conidia H–K conidia L di e si y o colony mo phology on CMD, MLA, OA, and PCA, espec i ely ( om le
o igh ) a e 4 wk. Images: on OA (A–K). Scale ba s: 50 µm (A, C); 100 µm (B); 10 µm (D–K); 1 cm (L).
64
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
his o ically placed in Pleu oph agmium con inues o hinde a obus axonomic
eassessmen o he genus. To cla i y he axonomy o Pleu oph agmium, we
conduc ed a mo phological compa ison o he accep ed species, which p o-
ides addi ional insigh s in o he mo phological a ini ies and dis inc ions o
i s species (Table 3). Based on his e alua ion, we did no iden i y any Neomy -
mec idium species ha could be ega ded as conspeci ic wi h al eady known
Pleu oph agmium species, al hough some axa me i close examina ion.
He po ichiellaceae: no el lineages in Thysano ea
The holoblas ic-den icula e mode o conidiogenesis, wi h conidia o en bo ne
on a sympodially elonga ing achis, occu s in se e al gene a o he He po-
ichiellaceae, including Aciculomyces (To es-Ga cia e al. 2023), Fonsecaea
(Naja zadeh e al. 2009), Neo e onaea (Qiu e al. 2022), Rhinocladiella (Melin
and Nann eld 1934; de Hoog and He manides-Nijho 1977; Mülle e al. 1987),
Ve onaea (Ci e i and Mon ema ini 1957), and Thysano ea. In addi ion, He nán-
dez-Res epo e al. (2020) epo ed a phialidic synasexual mo ph o Thysano-
ea sei e ii, which is p oduced only in cul u e. Fonsecaea and Rhinocladiella
also display ma ked pleomo phism, exhibi ing conside able a ia ion in asex-
ual mo phology, including se e al ypes o synasexual mo phs, and conidial
on ogeny unde a ying condi ions (Schol-Schwa z 1968; McGinnis 1983; de
Hoog and He manides-Nijho 1977; Tsuneda e al. 1986). The pleomo phic
na u e o se e al gene a in his amily u he complica es iden i ica ions
based solely on mo phology and has his o ically con ibu ed o hei polyphyly
wi hin he He po ichiellaceae.
The genus Thysano ea, based on T. papuana, was es ablished o ungi wi h
ma u e conidiopho es bea ing se e al ie s o b anchle s a anged apically in
a compac clus e , impa ing a dis inc ly a bo eous appea ance (A zanlou e
al. 2007). The sympodially elonga ing conidiogenous cells a e e minal o in-
e cala y on b anchle s, occasionally disc e e, wi h a conspicuous den icula e
achis. Recen s udies ha e shown ha he adi ionally diagnos ic b anching
pa e n o Thysano ea conidiopho es, used o dis inguish he genus om Pe i-
coniella (Sacca do and Be lese 1885), can a y wi h cul u e condi ions and may
be less p onounced in na u e o in young cul u es (Ki schne 2016; Wang e al.
2019; He nández-Res epo e al. 2020).
He nández-Res epo e al. (2020) poin ed ou ha Thysano ea is closely
ela ed o Minimelanolocus (Cas añeda-Ruiz 2001). Howe e , he placemen
o he M. na icula is, he ype species o he genus, emains unce ain due
o he absence o DNA sequence da a, wi h i s p esumed phylogene ic posi-
ion in he He po ichiellaceae in e ed om o he species cu en ly accep ed
in he genus (e.g. Liu e al. 2015; Wang e al. 2019). He nández-Res epo e
al. (2020) highligh ed mo phological di e ences be ween M. na icula is, and
species cu en ly assigned o Minimelanolocus based on molecula da a. Con-
sequen ly, hese au ho s ans e ed sequenced Minimelanolocus species o
Thysano ea and emended he genus.
In addi ion, molecula da a suppo he ans e o h ee Uncispo a species o
Thysano ea, o which we p opose new combina ions. Ou phylogene ic analy-
ses esol ed ex- ype s ains o U. hainanensis (Li e al. 2015), U. sinensis (Yang e
al. 2011), and U. wuzhishanensis (Liu e al. 2018a), wi hin he Thysano ea clade.
65
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
This e idence expands he mo phological concep o Thysano ea by inco po-
a ing new conidial cha ac e is ics. These axa sha e dis inc i e apically ape -
ing, cylind ical conidia, cu ed in he uppe po ion o e mina ing in a hooked
apical cell, and a e mo phologically incompa ible wi h U. ha oldiae, he ype
species o Uncispo a (Sinclai and Mo gan-Jones 1979). The la e is cha ac-
e ised by holoblas ic, de e mina e conidiogenous cells wi h a single la locus,
whe eas he analysed species possess polyblas ic, holoblas ic cells ha elon-
ga e sympodially, and conidia a e bo ne on den icles. The phylogene ic posi ion
o Uncispo a s. s . emains un esol ed. These indings emphasise he need o
ca e ul iden i ica ion o Uncispo a species, pa icula ly conside ing he diag-
nos ic alue o conidiogenous cell mo phology and shape o conidia, bu also
sugges he polyphyly o Uncispo a, wa an ing a na owe ci cumsc ip ion o
he genus based on eshly ecollec ed ma e ial.
Papulosaceae: expanding he di e si y o Wongia
Al hough Wongia was o iginally es ablished o wo sexually ep oducing
species (Khemmuk e al. 2016), se e al subsequen ly desc ibed axa ha e
been de ined solely by asexual ea u es (Luo e al. 2019; Bao e al. 2021;
Manawasinghe e al. 2024; Yu e al. 2024; Wang e al. 2024). Bo h new spe-
cies, W. pallidopola is and W. hachidopho a, con o m o he gene ic concep
o Wongia in p oducing mac onema ous, unb anched, da k b own conidio-
pho es wi h polyblas ic, sympodially p oli e a ing conidiogenous cells and
sep a e, b own conidia. Thei phylogene ic placemen wi hin he genus ex-
pands he known di e si y o Wongia.
Cu en ly, he genus Wongia comp ises 13 species. Wongia pallidopola is
and W. aqua ica o med a s ongly suppo ed subclade and include species
wi h ela i ely sho conidiopho es (≤ 90 µm long) and da k b own conidia
wi h dis inc ly pale end cells. By con as , all o he known species, including
W. hachidopho a, ep esen mo phologically dis inc and phylogene ically
s ongly suppo ed g oup cha ac e ised by subhyaline o pale b own conid-
ia ha a e mos ly e enly pigmen ed, and only a ely ha e end cells sligh ly
pale han he median cells. Wi hin he second subclades a e g ouped he
only species known ha ep oduce also sexually, namely W. iche ai, W. g i -
inii, and W. gu ula a (Khemmuk e al. 2016; Wang e al. 2024). They a e
cha ac e ised by pe i hecial, non-s oma ic ascoma a, ili o m pa aphyses,
uni unica e asci wi h a non-amyloid apical ing, and 3-sep a e, da k b own
ascospo es wi h pale b own o subhyaline end cells.
Wongia hachidopho a was nes ed wi hin a complex comp ising ou o h-
e closely ela ed species. While W. suae, W. bandungensis, and W. bambu-
sae a e mo phologically indis inguishable, di e ing only by sub le a ia ion
in conidial size, W. hachidopho a and W. usi o mis a e clea ly sepa a ed
om hem by conidial colou , sep a ion, and shape. Compa isons o ITS,
e 1, and pb2 sequences u he suppo W. hachidopho a as a dis inc
species. In con as , W. bandungensis and W. suae show almos comple e
sequence iden i y in e 1 and pb2 and di e by 1.4% in ITS. Gi en hei mo -
phological indis inguishabili y and minimal gene ic di e gence, hese wo
axa may ep esen e y ecen ly di e ged popula ions o a e likely conspe-
ci ic, pending u he popula ion-le el sampling.
66
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Tubeu iaceae: new gene a and species wi h holoblas ic-den icula e
conidiogenesis
Skoliomycella la a and Z. hili e a e new addi ions o he amily Tubeu iaceae.
The holoblas ic conidiogenesis on den icula e conidiogenous cells is a ea u e
ha is widesp ead and con e gen ac oss he amily. Skoliomycella and Zaanen-
omyces a e membe s o a well-suppo ed subclade comp ising ungi cha ac e -
ised by holoblas ic (mos ly holoblas ic-den icula e) conidiogenesis and h ee
conidial mo pho ypes, including Acan hos igma (De No a is 1863; Réblo á and
Ba 2000; Boonmee e al. 2014) and Helicospo ium (Nees on Esenbeck 1816;
Linde 1929; Lu e al. 2018a) wi h helicospo ous conidia, Neodic yospo a (Zhang
e al. 2023b) wi h dic yospo ous conidia, and Campo esiomyces (Hyde e al.
2020) and Zaanenomyces (C ous e al. 2021) wi h s aigh , a ely helicospo ous
conidia. Among hese gene a, Skoliomycella is mo phologically mos simila o
Campo esiomyces and Zaanenomyces, bu di e s in ha ing sinuous o genicu-
la e conidiopho es bea ing sca e ed den icles along hei en i e leng h.
Zaanenomyces, ypi ied by Z. quad ipa is, was es ablished by C ous e al.
(2021) o dema iaceous hyphomyce es wi h simple, e ec conidiopho es, e -
minal conidiogenous cells ha ex end sympodially o ming a achis and d y,
soli a y, hyaline, na owly obcla a e, sep a e conidia. All h ee Zaanenomyces
species ha e been eco ded om dead culms o Juncus spp. (C ous e al.
2021). Among hese, Z. hili e is closely ela ed o Z. quad ipa is. Bo h species
exhibi compa able conidiopho e and conidiogenous cell a chi ec u e; howe e ,
hey can be dis inguished by di e ences in conidial mo phology. In ou p elim-
ina y phylogene ic analyses o ITS–LSU sequences (da a no shown), Z. mod-
e a icis-academiae (ex- ype CBS 148312, and CBS149453) and Z. e sa ilis
(ex- ype CBS 148315), o med ei he a lineage dis inc om Z. quad ipa is–Z.
hili e subclade, o a monophyle ic bu s a is ically unsuppo ed clade in he ML
analyses. The e o e, addi ional loci ( pb2 and e 1) we e sequenced o hese
h ee species o comple e hei da ase . In he inal ou -gene analyses (Fig. 4),
all ou species we e eco e ed as a monophyle ic clade wi h mode a e s a is-
ical suppo in he ML analysis.
Gi en he a iable opology o he Zaanenomyces clade obse ed in se e -
al phylogene ic analyses, ex- ype s ains o h ee Zaanenomyces species we e
cul i a ed and examined o cul u al cha ac e is ics and mic omo phology.
Compa a i e analysis showed ha Z. mode a icis-academiae and Z. e sa ilis
di e om Z. hili e and Z. quad ipa is in se e al dis inc mo phological and
cul u al ea u es. Zaanenomyces mode a icis-academiae and Z. e sa ilis ha e
much pale , o en hyaline and mic onema ous conidiopho es, pa icula ly p o-
nounced in he o me species, and hei conidia ge mina e apidly. In con as ,
Z. hili e and Z. quad ipa is p oduce b own, mac onema ous conidiopho es
wi h a conspicuous, pigmen ed basal cell. Colony mo phology also a ies; col-
onies o Z. mode a icis-academiae and Z. e sa ilis a e uni o mly da k om
he cen e o pe iphe y, whe eas Z. quad ipa is and Z. hili e display a mo e
dis inc , pale ma gin and ce ain zona ion.
The ins abili y in phylogene ic esolu ion o he Zaanenomyces clade ac oss
analyses sugges s ha i s cu en ci cumsc ip ion may no adequa ely cap-
u e he ull ex en o mo phological a iabili y among i s species. This wa -
an s u he in es iga ion h ough expanded axon sampling, he inclusion o
67
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
addi ional molecula loci, and conside a ion o a possible e ision o he ge-
ne ic concep . Co espondingly, ou mo phological obse a ions indica e ha
Z. mode a icis-academiae and Z. e sa ilis o m a mo phologically cohesi e
subg oup wi hin Zaanenomyces, dis inguishable om he lineage comp ising
Z. hili e and Z. quad ipa is. This aises he possibili y ha hey may ep esen
a sepa a e, ye closely ela ed axon.
Mo phologically, Zaanenomyces is ema kably simila o Campo esiomyces
(Hyde e al. 2020), making bo h gene a di icul o dis inguish. Campo esiomy-
ces was o iginally desc ibed o wo sexually ep oducing species and he asex-
ual Ca. accinii (syn. Helicoma accinii, Ca is 1989), which has helicospo ous
conidia bo ne on den icula e conidiogenous cells. Han e al. (2025) expanded
he genus by adding h ee species de ined solely by asexual cha ac e is ics,
namely dema iaceous, mac onema ous conidiopho es o en in small clus-
e s, holoblas ic-den icula e conidiogenous cells, and hyaline o pale b own,
sep a e, s aigh conidia, he eby b oadening he gene ic concep o include
a non-helicospo ous conidial shape. The ex- ype s ain o H. accinii (ATCC
66068) is also p ese ed as CBS 216.90, and he la e is he sou ce o he
DNA sequences a ailable in GenBank (Tsui e al. 2006). In his s udy, we ex-
amined CBS 216.90 in i o, and we con i m ha i ma ches he p o ologue and
p oduces he cha ac e is ic helicospo ous conidia. Howe e , helicoid conidial
mo phology was omi ed om he p o ologue o Campo esiomyces (Hyde e
al. 2020) and was no adequa ely add essed by Han e al. (2025), who sug-
ges ed ha he occu ence o wo conidial ypes wi hin he genus migh be he
esul o geog aphical isola ion in Ca. accinii.
Campo esiomyces accinii is eadily dis inguished om Helicoma by i s co-
nidiopho es and conidial mo phology. In Ca. accinii, conidiopho es e mina e
in an in eg a ed, den icula e conidiogenous cell, and he conidia a e coiled bu
asymme ical wi h a dis inc basal cell ha is elonga ed, ape ing, and unca e
a he base. In con as , Helicoma de elops se i o m conidiopho es wi h nume -
ous in e cala y conidiogenous cells, usually bea ing one o wo den icles, and
p oduces symme ical, coiled conidia wi h cells ha o en become p og essi ely
smalle om he cen e owa d bo h ends. The conidiopho e a chi ec u e o Ca.
accinii is consis en wi h o he Campo esiomyces species ha o m s aigh
conidia. No ably, he s aigh , dis inc ly unca e basal cell o he o he wise
coiled conidium o Ca. accinii may e lec an in e media e s age in he mo -
phogene ic di e si ica ion o conidial o ms wi hin Campo esiomyces, poin ing
o a b oade mo phological plas ici y in he genus han p e iously ecognised.
Conidial mo phology wi hin he Tubeu iaceae is highly di e se. The helicoid,
sep a e, hyaline o pale b own conidial ype is he mos common mo pho ype
ha occu s in he majo i y o gene a in he amily. In addi ion o he s aigh ,
hyaline, sep a e conidia o Campo esiomyces, Skoliomycella, and Zaanenomy-
ces, o he o ms include da kly pigmen ed dic yoconidia o monodic ys-like
asexual mo phs, desc ibed in he li e cycles o Chlamydo ubeu ia (Boonmee
e al. 2011), Dic yospo a (B ahmanage e al. 2017), Manoha acha iella (Bag-
yana ayana e al. 2009), Mu ipulch a (Luo e al. 2017), Neochlamydo ubeu ia
(Lu e al. 2018a), Neodic yospo a (Zhang e al. 2023b), and Tubeu ia amazon-
ensis (Samuels e al. 1987). Whe eas he helicospo ous and s aigh conidia
a e p oduced on holoblas ic-den icula e conidiogenous cells, he dic yoconidia
a ise om holoblas ic conidiogenous cells wi h a la , non-den icula e locus.
68
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Tubeu ia amazonensis also p oduces a pycnidial, as e omella-like synasexual
mo ph (synanamo ph). The sys ema ic placemen o Tu. amazonensis emains
unce ain. Based on ascoma al wall cha ac e s, C ane e al. (1998) ans e ed
his species o Thax e iella, a genus cha ac e ised by helicospo ous conidia.
Howe e , no molecula da a a e cu en ly a ailable o Thax e iella spp., lea ing
i s phylogene ic posi ion un esol ed.
Conclusions
This s udy highligh s he impo ance o ou ine molecula and mo phological
e i ica ion o s ains ob ained e en om well-cu a ed cul u e collec ions. The
case o Pleu oph agmium pa ispo um demons a es how sub le mo phologi-
cal cha ac e s, i misin e p e ed, can obscu e ue phylogene ic ela ionships.
Ou indings indica e ha his o ical misiden i ica ions pe sis and may pe pe -
ua e axonomic inaccu acies i le un e i ied.
Phylogene ic analyses e ealed ha se en s ains deposi ed unde he name
P. pa ispo um ep esen a polyphyle ic assemblage dis ibu ed ac oss ou am-
ilies and/o o de s in h ee classes, namely in he Do hideomyce es (Tubeu iales),
Eu o iomyce es (Chae o hy iales), and So da iomyce es (My mec idiales and Pap-
ulosaceae). Only one s ain, CBS 770.83, con o ms o he species concep o P.
pa ispo um, con i ming i s placemen wi hin he My mec idiales. These esul s
led o he synonymisa ion o Neomy mec idium wi h Pleu oph agmium and he
p oposal o one new genus, se e al new species, new names and combina ions.
Acknowledgemen s
We since ely hank Ga y J. Samuels o eading he manusc ip and o his
aluable commen s and sugges ions. We hank he cul u e collec ion manage
Ge a d Ve kleij (CBS) o a ailabili y o he s udied s ains, and cul u e and col-
lec ion cu a o s T ix Me kx (CBS) and Dana Lančo á (PRA) o hei in aluable
assis ance in ob aining li ing cul u es and acili a ing he deposi ion o new
he ba ium specimens and d ied s ains. We a e g a e ul o Kazuaki Tanaka o
checking axa and hei GenBank Accessions used in his s udy.
Addi ional in o ma ion
Con lic o in e es
The au ho s ha e decla ed ha no compe ing in e es s exis .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
Adhe ence o na ional and in e na ional egula ions
All he ungal s ains used in his s udy ha e been legally ob ained, espec ing he Con-
en ion on Biological Di e si y (Rio Con en ion).
69
IMA Fungus 16: e173033 (2025), DOI: 10.3897/ima ungus.16.173033
Ma ina Réblo á e al.: Pleu oph agmium pa ispo um
Funding
This s udy was suppo ed by he p ojec o he Czech Academy o Sciences ‘S a e-
gie AV21 MycoLi e – s ě hub’ (M.R.), and long- e m esea ch de elopmen p ojec s,
no. RVO 67985939 o he Czech Academy o Sciences (M.R., L.B.) and MH CZ – DRO
(UHHK, 00179906) o he Uni e si y Hospi al H adec K álo é (J.N.).
Au ho con ibu ions
Concep ualiza ion: MR, MHR. Fo mal analysis: JN, MR, MHR, LB. Funding acquisi ion:
MR. In es iga ion: JN, MR, LB, MHR. Me hodology: JN, MR, MHR. P ojec adminis a ion:
MR. Resou ces: MR. Supe ision: MR. Valida ion: MR. W i ing - o iginal d a : MR, MHR.
W i ing - e iew and edi ing: JN, MHR, MR.
Au ho ORCIDs
Ma ina Réblo á h ps://o cid.o g/0000-0001-5229-1709
Jana Nek indo á h ps://o cid.o g/0000-0002-2861-5483
Lucie Baucho á h ps://o cid.o g/0009-0001-2658-9103
Ma ga i a He nández-Res epo h ps://o cid.o g/0000-0001-5157-3392
Da a a ailabili y
All o he da a ha suppo he indings o his s udy a e a ailable in he main ex o
Supplemen a y In o ma ion.
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Supplemen a y ma e ial 1
Alignmen s o DNA sequences
Au ho s: Ma ina Réblo á
Da a ype: x
Explana ion no e: Alignmen s o sequences o membe s o he My mec idiales, He po-
ichiellaceae, Papulosaceae and Tubeu iaceae.
Copy igh no ice: This da ase is made a ailable unde he Open Da abase License
(h p://openda acommons.o g/licenses/odbl/1.0/). The Open Da abase License
(ODbL) is a license ag eemen in ended o allow use s o eely sha e, modi y, and
use his Da ase while main aining his same eedom o o he s, p o ided ha he
o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/ima ungus.16.173033.suppl1
Supplemen a y ma e ial 2
Biogeog aphical dis ibu ion, sample ype, habi a and o he de ailed me ada a
o newly desc ibed species in e ed om he GlobalFungi da abase
Au ho s: Ma ina Réblo á
Da a ype: xlsx
Copy igh no ice: This da ase is made a ailable unde he Open Da abase License
(h p://openda acommons.o g/licenses/odbl/1.0/). The Open Da abase License
(ODbL) is a license ag eemen in ended o allow use s o eely sha e, modi y, and
use his Da ase while main aining his same eedom o o he s, p o ided ha he
o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/ima ungus.16.173033.suppl2
