Morphological and phylogenetic analyses reveal novel entomopathogenic fungi infecting scale insects and aphids in China

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Mo phological and phylogene ic analyses e eal no el
en omopa hogenic ungi in ec ing scale insec s and aphids in China
Chunlin Yang1,2 , Xiulan Xu3, Xinyue Li1,2, Feng Liu1,2, Zhen Zeng3, Qiangang Xiao3, Yinggao Liu1,2
1 College o Fo es y, Sichuan Ag icul u al Uni e si y, Chengdu 611130, China
2 Na ional Fo es y and G assland Adminis a ion Key Labo a o y o Fo es Resou ces Conse a ion and Ecological Sa e y on he Uppe Reaches o he Yang ze
Ri e , College o Fo es y, Sichuan Ag icul u al Uni e si y, Chengdu 611130, China
3 Fo es y Resea ch Ins i u e, Chengdu Academy o Ag icul u al and Fo es y Sciences, Chengdu 611130, China
Co esponding au ho s: Chunlin Yang ([email p o ec ed]); Xiulan Xu ([email p o ec ed])
Copy igh : © Chunlin Yang 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
En omopa hogenic ungi exhibi a cosmopoli an dis ibu ion ac oss di e se ecosys ems,
wi h hei ubiqui ous p esence in insically linked o insec habi a s—essen ially occu ing
whe e e insec popula ions exis . These ungi ep esen a i al biological esou ce, pa -
icula ly in ag icul u e and o es y. They se e as a c ucial eposi o y o ungal s ains o
biological pes con ol. This in es iga ion iden i ied se en species om sou hwes Chi-
na based on mul i-gene (ITS, LSU, SSU, ac , pb1, pb2, and e 1-α) phylogene ic analyses
and mo phological cha ac e is ics, including one new species and one newly eco ded
species in Cladospo ium (Cladospo iaceae); ou new species in Moelle iella (Cla icipi a-
ceae); and one new species in Podonec ia (Podonec iaceae). All se en ungi a e in hei
asexual mo phs and we e disco e ed on aphids o scale insec s, which a e common and
signi ican pes s. These include Cladospo ium kuwanaspidis, Cladospo ium guizhouense,
Moelle iella eucalyp i, Moelle iella boehme iae, Moelle iella cinnamomum, Moelle iella ci us,
and Podonec ia mul ia ma a. Desc ip ions and illus a ions o all se en axa a e p o ided.
Six o hese species we e collec ed om scale insec s, speci ically hose ound in bamboo
o es s, b oad-lea ed o es s, o unde s o y ege a ion, and one was collec ed om aphids,
p ima ily dis ibu ed on he unde side o nigh -blooming jasmine lea es. This wo k e eals
he ich di e si y o en omopa hogenic ungi in sou hwes e n China, no only on la ge in-
sec s such as Lepidop e a and Hymenop e a bu also on smalle Hemip e a, many o which
a e signi ican ag icul u al and o es y pes s. This s udy con ibu es ungal esou ces ha
may suppo he de elopmen o inno a i e pes con ol me hods in he u u e.
Key wo ds: 6 new axa, Cladospo iaceae, Cla icipi aceae, ungal en omopa hogens,
Podonec iaceae
In oduc ion
To ensu e en i onmen al sa e y, ungal agen s a e conside ed key biological
egula o s in he applica ion o insec icides o pes con ol (Roy and Co ell
2008; Sha ma and Sha ma 2021). The ini ial ecommenda ion o use mic obi-
al insec pa hogens o con olling insec s was p oposed by LeCon e (1874)
and Pas eu (1874). Subsequen ly, he i s success ul applica ion o Me a -
hizium anisopliae was conduc ed in Russia agains Bo hynode es punc i en is
Academic edi o : João A aújo
Recei ed:
27 Augus 2025
Accep ed:
9 Sep embe 2025
Published:
8 Oc obe 2025
Ci a ion: Yang C, Xu X, Li X, Liu F, Zeng
Z, Xiao Q, Liu Y (2025) Mo phological
and phylogene ic analyses
e eal no el en omopa hogenic
ungi in ec ing scale insec s and
aphids in China. IMA Fungus 16:
e170123. h ps://doi.o g/10.3897/
ima ungus.16.170123
IMA Fungus 16: e170123 (2025)
DOI: 10.3897/ima ungus.16.170123
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IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
(Coleop e a: Cu culionidae) by K assils schik (1888). This achie emen buil
upon he ea lie iden i ica ion o he mic obial agen , ini ially named En omoph-
ho a anisopliae, which a ge ed Anisoplia aus iaca (Coleop e a: Sca abaeidae).
To da e, mo e han 1,000 species o en omopa hogenic ungi ha e been doc-
umen ed ac oss o e 100 gene a (Hywel-Jones 1993; Vega e al. 2012; Hawk-
swo h and Lücking 2017; Co allo e al. 2019; Wu e al. 2024). Among hese
species, such as Akan homyces Lebe , Beau e ia Vuill., Co dyceps F . (=Isa ia
Pe s.), Lecanicillium W. Gams & Za e, and Me a hizium So okin, a e widely ec-
ognized o hei po en ial in biological con ol. No ably, species like A. aculea-
us, B. bassiana, B. b ongnia ii, C. umoso osea, L. lecanii (≡Ve icillium lecanii),
L. musca ium, L. longispo um, and M. anisopliae a e commonly employed in
pes managemen s a egies (Fa ia and W aigh 2007; McKinnon e al. 2017;
Co allo e al. 2019; S one and Bidochka 2020; Nicole i and Becchimanzi 2020;
Nishi e al. 2021; Khonsani e al. 2024). Fo example, 171 p oduc s ha e been
de eloped as biocon ol agen s, p ima ily based on B. bassiana, B. b ongnia ii,
C. umoso osea, and M. anisopliae (Fa ia and W aigh 2007). Beau e ia bassiana,
ma ke ed as “Myco ol”, has been u ilized as a comme cial mycoinsec icide o
a ge a b oad spec um o insec pes s in No h Ame ica (W aigh e al. 2021).
Simila ly, he comme cial p oduc “G een Muscle,” de i ed om M. ac idum
(≡M. anisopliae a . ac idum), has p o en e ec i e agains locus and g asshop-
pe pes s in A ica (Niassy and Dia a 2011). In addi ion, en omopa hogenic un-
gi exhibi di e se nu i ional beha io s— unc ioning as bio ophs, nec o ophs,
o hemibio ophs (Vega e al. 2009)—and demons a e addi ional ecological
oles, including hizosphe e coloniza ion, ungal endophy ism, plan disease an-
agonism, and enhancemen o plan g ow h o s ess esis ance (Kabaluk and
E icsson 2007; Kim e al. 2008; Ownley e al. 2008; Vega 2008; Pa a-Ripoll e al.
2011; Aku se e al. 2013; Jabe and Enke li 2017; Jabe and Ownley 2017; Man-
zoukas and Eliopoulos 2020; Sha ma and Sha ma 2021; Muola e al. 2023).
Insec -associa ed ungi ha e been widely epo ed ac oss he ex ensi e e-
gions sou h o he Yellow Ri e basin, wi h mos species displaying dis ibu ion
pa e ns cha ac e is ic o opical, sub opical, and empe a e zones (Liang e al.
2016; Chen e al. 2021; Wei e al. 2022; Xiao e al. 2023; Chuang e al. 2024; Wang
e al. 2024, 2025), bu ha e been a ely documen ed in he Sichuan egion. The
de elopmen o en omogenous ungi is closely linked o ege a ion and insec
popula ions, as well as clima ic and soil condi ions (Hajek and Lege 1994; Que-
sada-Mo aga e al. 2007, 2024; Vega e al. 2009; Medo and Cagáň 2011), al hough
expe imen al alida ion in his a ea emains limi ed. Sca e ed s udies ha e indi-
ca ed ha en omogenous ungi a e mo e commonly ound in a eas wi h ich eg-
e a ion, ela i ely high humidi y, sui able empe a u es, and shaded en i onmen s.
Chuang e al. (2024) ound ha in Taiwan, China, en omogenous ungi belong-
ing o he Co dycipi aceae a e mo e eadily collec ed in egions wi h an a e age
empe a u e o 19–20.5 °C, ain all anging om 2,300 o 3,150 mm, and ela i e
humidi y be ween 81% and 82.5%. Wang e al. (2025) collec ed and desc ibed
wo new gene a and 13 new species wi hin he Cla icipi aceae, associa ed wi h
whi e lies and scale insec s on a ious hos plan s in Yunnan and Hainan, China.
Sichuan is ecognized as one o he key biodi e si y esea ch ho spo s, ow-
ing o i s di e se ecosys ems, complex habi a condi ions, and ich plan spe-
cies. While he egion suppo s a wide a ie y o insec auna, en omogenous
ungi ha e no been sys ema ically s udied, and hei species di e si y is likely
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IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
signi ican ly unde es ima ed. In ou ongoing sea ch o en omogenous ungi
associa ed wi h minia u e insec s, se e al new axa o en omopa hogenic ungi
(Nec iaceae; Podonec iaceae) we e isola ed om scale insec s collec ed in
Sichuan P o ince, p ima ily om bamboo and walnu hos s (Xu e al. 2021a;
Liu e al. 2023). To u he explo e he di e si y o en omogenous ungi, we
conduc ed ield collec ions a mul iple si es ac oss Sichuan. De ailed mo pho-
logical desc ip ions a e p o ided along wi h ele an ecological da a.
Comp ehensi e phylogene ic analyses we e pe o med using a se en-locus
da ase (ITS, LSU, SSU, e 1-α, pb1, pb2, and ac ) o ensu e accu a e axonomic
posi ion. This s udy en iches he en omogenous ungi da abase in Sichuan and
o e s aluable s ains o sus ainable pes con ol in ag icul u e and o es y.
Ma e ials and me hods
Fungal collec ion and isola ion
Specimens consis ing o whole lea es o b anches bea ing ui ing bodies we e
collec ed om Chengdu, Dazhou, Guangan, Leshan, and Meishan in Sichuan
P o ince, China, placed in s e ilized plas ic bags, and anspo ed o he labo-
a o y o analysis. The p ocedu e o ob aining axenic cul u es, as desc ibed
by Chomnun i e al. (2014), in ol ed isola ing pu e cul u es, ans e ing hem
o PDA slan s o 10 days o cul i a ion, and subsequen ly s o ing hem a 4 °C.
The specimens we e deposi ed in he He ba ium o Sichuan Ag icul u al Uni-
e si y (SICAU), Chengdu, China, while he s ains we e s o ed in he Cul u e
Collec ion o Sichuan Ag icul u al Uni e si y (SICAUCC), China.
Mo phological obse a ions
F ui ing bodies we e obse ed and pho og aphed using a dissec ing mic oscope
NVTGG (Shanghai Ad anced Pho oelec ic Technology Co. L d., Shanghai, Chi-
na) i ed wi h a VS-800C mic o-digi al came a (Shenzhen Weishen Times Tech-
nology Co. L d., Shenzhen, China). The dimensions o conidioma a, pa aphy-
ses, conidiopho es, conidiogenous cells, and conidia we e measu ed om ield
samples and pho og aphed using an Olympus BX43 compound mic oscope
equipped wi h an Olympus DP22 digi al came a, in conjunc ion wi h ACDSee 3.1
so wa e. Measu emen s we e made using Ta oso ® Image F ame Wo k .0.9.7
(Ta oso (R), Non ha Bu i, Thailand). Lac ophenol co on blue eagen was used
o obse e he conidiogenous s uc u es and de e mine he numbe o sep a. To
obse e and documen he colo and ex u e o he colonies, esh pla es we e
p epa ed om pu i ied colonies and incuba ed a 25 °C o 1 o 3 weeks.
DNA ex ac ion, ampli ica ion, and sequencing
To al genomic DNA was ex ac ed om mycelia g own on PDA o single ui -
ing bodies using he Plan Genomic DNA ex ac ion ki (Tiangen, China). P im-
e pai s ITS5 and ITS4 (Whi e e al. 1990), NS1 and NS4 (Whi e e al. 1990),
LR0R and LR5 (Vilgalys and Hes e 1990), EF1-983F and EF1-2218R (Rehne
and Buckley 2005), RPB1-Ac and RPB1-C (Lomba d e al. 2015), RPB2-5F
and RPB2-7cR (Liu e al. 2000), and ACT512F/ACT-783R (Ca bone and Kohn
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IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
1999) we e used o ampli y he in e nal ansc ibed space (ITS), he pa ial
small subuni nuclea DNA (SSU), he nuclea ibosomal la ge subuni (LSU),
he ansla ion elonga ion ac o e 1-α, he la ges subuni o RNA polyme ase
II ( pb1), he second la ges subuni o RNA polyme ase II ( pb2), and he ac in
gene (ac ), espec i ely. Addi ionally, o Cladospo ium species, EF1728F and
EF2 (Lomba d e al. 2015) we e used o ampli y e 1-α.
Polyme ase chain eac ion (PCR) was pe o med in a 25 μL eac ion mix u e
con aining 22 μL Mas e Mix (Beijing TsingKe Bio ech Co. L d., Beijing, China),
1 μL DNA empla e (10–30 ng/μL), and 1 μL o each p ime (120–150 ng/μL).
The ampli ica ion eac ions we e pe o med as desc ibed by Lomba d e al.
(2015), Dai e al. (2016), and Wang e al. (2022a). PCR p oduc s we e se-
quenced a TsingKe Biological Technology Co. L d., Chengdu, China. The newly
gene a ed sequences we e deposi ed in GenBank.
Phylogene ic analyses
Phylogene ic analyses we e conduc ed using sequences om Cladospo ium
(ITS, ac , and e 1-α), Moelle iella (LSU, pb1, and e 1-α), and Podonec ia (ITS,
LSU, SSU, e 1-α, and pb2). Mul igene sequences om a ious species (see
Tables 1–3) we e ob ained om GenBank, along wi h addi ional sequences
gene a ed in his s udy. DNA alignmen s we e ca ied ou using he MAFFT
.7.429 online se ice (Ka oh e al. 2019), and ambiguous egions we e exclud-
ed using BioEdi e sion 7.0.5.3 (Hall 1999). Mul igene sequences we e conca -
ena ed using Mesqui e so wa e (Maddison and Maddison 2019). Mul igene
phylogene ic analyses we e conduc ed using maximum likelihood (ML) and
Bayesian in e ence (BI) me hods. The op imal nucleo ide subs i u ion model
was iden i ied wi h M Model es .2.2 (Nylande 2004).
ML and BI analyses we e pe o med using he CIPRES Science Ga eway web
se e (Mille e al. 2010). Fo he ML analysis, RAxML-HPC2 on XSEDE ( .8.2.10)
(S ama akis 2014) was employed wi h he GTR+GAMMA subs i u ion model
and 1,000 boo s ap i e a ions. Fo BI analyses, he bes - i models we e selec -
ed using M Model es .2.2 o each da ase : Cladospo ium (ITS: SYM+I+G, ac :
GTR+G, e 1-α: GTR+I+G), Moelle iella (LSU: GTR+I+G, pb1: GTR+I+G, e 1-α:
GTR+I+G), and Podonec iaceae (ITS: GTR+I+G, LSU: GTR+I+G, SSU: GTR+I+G,
e 1-α: GTR+I+G, pb2: GTR+I+G). The analyses we e compu ed wi h six si-
mul aneous Ma ko Chain Mon e Ca lo (MCMC) chains, un o 1,000,000 o
10,000,000 gene a ions wi h a sampling equency o e e y 100 gene a ions.
The bu n-in ac ion was se o 0.25, and he un was au oma ically e mina ed
when he a e age s anda d de ia ion o spli equencies ell below 0.01.
Phylogene ic ees we e isualized using FigT ee .1.4.3 (Rambau and
D ummond 2016) and u he edi ed wi h Adobe Illus a o CS6 (Adobe Sys-
ems Inc., Uni ed S a es). Maximum likelihood boo s ap alues (MLBS) o 60%
o g ea e and Bayesian pos e io p obabili ies (BIPP) o 0.95 o highe we e
indica ed on he ees.
Abb e ia ions
ac The ac in gene
BI Bayesian in e ence
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IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
bp Base pai
BIPP Bayesian in e ence pos e io p obabili ies
DNA Deoxy ibonucleic acid
ITS in e nal ansc ibed space
LSU The nuclea ibosomal la ge subuni
MEA Mal ex ac aga medium
ML Maximum likelihood
MLBS Maximum likelihood boo s ap p opo ions
PCR Polyme ase chain eac ion
PDA Po a o dex ose aga medium
pb1 The la ges subuni s o RNA polyme ase II
pb2 The RNA polyme ase II second la ges subuni
SICAU The he ba ium o Sichuan Ag icul u al Uni e si y
SICAUCC The cul u e collec ion in Sichuan Ag icul u al Uni e si y
SNA Syn he ic low nu ien aga medium
SSU The pa ial small subuni nuclea DNA
e 1-α The ansla ion elonga ion ac o 1α
Table 1. Vouche in o ma ion and GenBank accession numbe s o he axa used in he Cladospo ium.
Species Vouche in o ma ion ITS ac e 1-α Re e ences
Cladospo ium acalyphae CBS 125982 T HM147994 HM148481 HM148235 Bensch e al. (2010)
Cl. albo la escens CBS 140690 T LN834420 LN834604 LN834516 Sando al-Denis e al. (2016)
Cl. angulosum CBS 140692 T LN834425 LN834609 LN834521 Sando al-Denis e al. (2016)
Cl. angus ispo um CBS 125983 T HM147995 HM148482 HM148236 Bensch e al. (2010)
Cl. angus i e minale CBS 140480 T KT600379 KT600575 KT600476 Bensch e al. (2015)
Cl. an h opophilum CBS 140685 T LN834437 LN834621 LN834533 Sando al-Denis e al. (2016)
Cl. a enosum CHFC-EA 566 T MN879328 MN890008 MN890011 C ous e al. (2020a)
Cl. a mandiae CBS 153756 T PQ066521 PQ067354 PQ067355 Tan e al. (2024)
Cl. aspe ula um CBS 126340 T HM147998 HM148485 HM148239 Bensch e al. (2010)
Cl. aulonemiae COAD 2269 T MZ318427 MT373119 MT680198 Cos a e al. (2022)
Cl. aulonemiae COAD 2270 MZ318428 MT373120 MT680199 Cos a e al. (2022)
Cl. aus aliense CBS 125984 T HM147999 HM148486 HM148240 Bensch e al. (2010)
Cl. aus oa icanum CBS 140481 T KT600381 KT600577 KT600478 Bensch e al. (2015)
Cl. aus oli o ale CBS 148321 T MN879327 MN890007 MN890010 C ous e al. (2021a)
Cl. bambusicola COAD 2256 T MZ318433 MT373125 MT680204 Cos a e al. (2022)
Cl. bambusicola COAD 2565 OP535372 OP598124 OP676083 Cos a e al. (2022)
Cl. benschiae COAD 2263 T MZ318436 MT373128 MT680207 Cos a e al. (2022)
Cl. benschiae COAD 2265 MZ318437 MT373129 MT680208 Cos a e al. (2022)
Cl. ben i oglioae BRIP 74745a T OR947065 OR964960 OR964961 Tan and Shi as (2023a)
Cl. b igadei ense COAD 2257 T MZ318435 MT373127 MT680206 Cos a e al. (2022)
Cl. cap i imosum FMR 16532 T LR813198 LR813205 LR813210 I u ie a-González e al. (2021)
Cl. ca e nicola URM 8389 T MZ518829 MZ555746 MZ555733 Pe ei a e al. (2022)
Cl. chalas ospo oides CBS 125985 HM148001 HM148488 HM148242 Bensch e al. (2010)
Cl. chasman hicola CPC 21300 T KY646221 KY646224 KY646227 Ma in-Felix e al. (2017)
Cl. chlamydospo igenum AUMC 11340 T MN826919 OL514009 Moha am e al. (2022)
Cl. chlamydospo i o mans COAD 2571 T OP535374 OP598126 OP676085 Pe ei a e al. (2024)
Cl. chlamydospo i o mans COAD 2568 OP535378 OP598130 OP676089 Pe ei a e al. (2024)

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IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Species Vouche in o ma ion ITS ac e 1-α Re e ences
Cl. chusqueae COAD 2258 T MZ318430 MT373122 MT680201 Cos a e al. (2022)
Cl. chusqueae COAD 2261 MZ318431 MT373124 MT680203 Cos a e al. (2022)
Cl. chubu ense CBS 124457 T FJ936158 FJ936165 FJ936161 Schube e al. (2009)
Cl. cladospo ioides CBS 112388 T HM148003 HM148490 HM148244 Bensch e al. (2010)
Cl. compac ispo um AUMC 11366 T MN826822 OL514010 Moha am e al. (2022)
Cl. colocasiae CBS 386.64 T HM148067 HM148555 HM148310 Bensch e al. (2010)
Cl. colombiae CBS 274.80B T FJ936159 FJ936166 FJ936163 Schube e al. (2009)
Cl. congjiangense GUCC 21208.3 T OP852675 OP863094 OP859042 Yang e al. (2023)
Cl. congjiangense GUCC 21208.5 OP852676 OP863095 OP859043 Yang e al. (2023)
Cl. cop ophilum FMR 16164 T LR813201 LR813207 LR813213 I u ie a-González e al. (2021)
Cl. co icola BRIP 74385a T OP256851 OP288997 OP288998 C ous e al. (2023)
Cl. c ousii CBS 140686 T LN834431 LN834615 LN834527 Sando al-Denis e al. (2016)
Cl. cucume inum CBS 171.52 T HM148072 HM148561 HM148316 Bensch e al. (2010)
Cl. delica ulum CBS 126344 HM148081 HM148570 HM148325 Bensch e al. (2010)
Cl. de ikae BRIP 72278a T MZ303808 MZ344212 MZ344193 P asanna h e al. (2021a, 2021b)
Cl. diaman inense COAD 3108 T ON062328 ON141933 ON982817 Du a e al. (2023)
Cl. endo i icola JZBH 390018 T MN654960 MN984220 MN984228 Manawasinghe e al. (2020)
Cl. eucommiae GUCC 401.1 T OL587465 OL519775 OL504966 Wang e al. (2022b)
Cl. eucommiae GUCC 401.9 ON334729 ON383337 Wang e al. (2022b)
Cl. eu opaeum CBS 134914 T HM148056 HM148543 HM148298 Bensch e al. (2018)
Cl. eu opaeum CBS 116744 HM148053 HM148540 HM148294 Bensch e al. (2018)
Cl. exaspe a um CBS 125986 HM148090 HM148579 HM148334 Bensch e al. (2010)
Cl. exile CBS 125987 HM148091 HM148580 HM148335 Bensch e al. (2010)
Cl. labelli o me CBS 126345 HM148092 HM148581 HM148336
Cl. la o i ens CBS 140462 T LN834440 LN834624 LN834536 Sando al-Denis e al. (2016)
Cl. uniculosum CBS 122129 T HM148094 HM148583 HM148338 Bensch e al. (2010)
Cl. usco i ide FMR 16385 T LR813200 LR813206 LR813212 I u ie a-González e al. (2021)
Cl. gamsianum CBS 125989 T HM148095 HM148584 HM148339 Bensch e al. (2010)
Cl. globispo um CBS 812.96 T HM148096 HM148585 HM148340 C ous e al. (2011)
Cl. g e illeae CBS 114271 T JF770450 JF770473 JF770472 C ous e al. (2011)
Cl. guizhouense GUCC 401.7 T OL579741 OL519780 OL504965 Wang e al. (2022b)
Cl. guizhouense GUCC 401.8 ON334728 ON383338 ON383470 Wang e al. (2022b)
Cl. guizhouense SICAUCC 25-0057 PV156487 PV153466 PV153474 In his s udy
Cl. guizhouense SICAUCC 25-0058 PV156488 PV153467 PV153475 In his s udy
Cl. hemileiicola COAD 2567 T OP535376 OP598128 OP676087 Pe ei a e al. (2024)
Cl. hemileiicola COAD 3350 OP535381 OP598133 OP676092 Pe ei a e al. (2024)
Cl. he e opogonicola BRIP 72465a T OL307932 OL332743 OL332742 Tan e al. (2022)
Cl. hillianum CBS 125988 T HM148097 HM148586 HM148341 Bensch e al. (2010)
Cl. in e sicolo CBS 401.80 T HM148101 HM148590 HM148345 Bensch e al. (2010)
Cl. ipe eniae CBS 140483 T KT600394 KT600589 KT600491 Bensch e al. (2015)
Cl. i anicum CBS 126346 T HM148110 HM148599 HM148354 Bensch e al. (2010)
Cl. kaiyangense GUCC 21265.2 T OP852665 OP863097 OP859045 Yang e al. (2023)
Cl. kenpeggii CPC 19248 T KY646222 KY646225 KY646228 Ma in-Felix e al. (2017)
Cl. kuwanaspidis SICAUCC 25-0063 T PV156489 PV153468 PV153480 In his s udy
Cl. kuwanaspidis SICAUCC 25-0064 PV156490 PV153469 PV153481 In his s udy
Cl. lagena ii o me SFC20230103-M23 T OQ186119 OQ185167 OQ185128 Lee e al. (2023)
7
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Species Vouche in o ma ion ITS ac e 1-α Re e ences
Cl. len ulum FMR 16288 T LR813203 LR813209 LR813215 I u ie a-González e al. (2021)
Cl. licheniphilum CBS 125990 HM148111 HM148600 HM148355 Bensch e al. (2010)
Cl. longica ena um CBS 140485 KT600403 KT600598 KT600500 Bensch e al. (2015)
Cl. longissimum CBS 300.96 DQ780352 EF101385 EU570259
Cl. lycope dinum CBS 574.78C HM148115 HM148604 HM148359 P asanna h e al. (2021a, 2021b)
Cl. macadamiae BRIP 72269a T MZ303810 MZ344214 MZ344195 Jayasi i e al. (2019)
Cl. magnoliigena MFLUCC 18-1559 T MK347813 MK340864 Lee e al. (2023)
Cl. mal i imosum SFC20230103-M51 T OQ186147 OQ185195 OQ185155 Lee e al. (2023)
Cl. ma inum SFC20230103-M33 T OQ186129 OQ185177 OQ185137 Bensch e al. (2015)
Cl. mon ecillanum CBS 140486 T KT600406 KT600602 KT600504 Bensch e al. (2010)
Cl. my acea um CBS 126350 T HM148117 HM148606 HM148361 Yang e al. (2023)
Cl. nayongense GUCC 21260.3 T OP852669 OP863106 OP859054 Zimowska e al. (2021)
Cl. neapoli anum MgPo1 T MK387890 MK416051 MK416094 Bensch e al. (2018)
Cl. needhamense CBS 143359 T MF473142 MF473991 MF473570 Bensch e al. (2018)
Cl. nee landicum CBS 143360 T KP701887 KP702010 KP701764 Bensch e al. (2018)
Cl. neopsych o ole ans CGMCC 3.18031 T KX938383 KX938366 KX938400 Bensch e al. (2010)
Cl. oxyspo um CBS 125991 HM148118 HM148607 HM148362 Bensch e al. (2010)
Cl. pa acladospo ioides CBS 171.54 T HM148120 HM148609 HM148364 Bensch e al. (2015)
Cl. pa apenidielloides CBS 140487 T KT600410 KT600606 KT600508 Rosado e al. (2019)
Cl. passi lo ae COAD 2135 T MH682175 MH729795 MH724943 Rosado e al. (2019)
Cl. passi lo icola COAD 2140 T MH729800 MH724948 Bensch e al. (2010)
Cl. pe angus um CBS 125996 T HM148121 HM148610 HM148365 Bensch e al. (2018)
Cl. pe angus um CBS 126365 HM148123 HM148612 HM148367 Bensch e al. (2018)
Cl. pe angus um CPC 11663 HM148128 HM148617 HM148372 Sando al-Denis e al. (2016)
Cl. pe angus um CPC 13870 HM148142 HM148631 HM148386 Bensch e al. (2010)
Cl. pe angus um FMR 13321 LN834380 LN834564 LN834476 Sando al-Denis e al. (2016)
Cl. pe nambucoense URM 8390 T MZ518830 MZ555747 MZ555734 Pe ei a e al. (2022)
Cl. pe nambucoense URM 8391 MZ518828 MZ555745 MZ555732 Pe ei a e al. (2022)
Cl. phaenocomae CBS 128769 JF499837 JF499881 JF499875 C ous and G oenewald (2011)
Cl. phyllac iniicola CBS 126352 HM148150 HM148639 HM148394
Cl. phyllophilum CBS 125992 T HM148154 HM148643 HM148398 Bensch e al. (2010)
Cl. pini-ponde osae CBS 124456 T FJ936160 FJ936167 FJ936164 Schube e al. (2009)
Cl. polonicum MgPo1 T MK387894 MK416055 MK416098 Zimowska e al. (2021)
Cl. p o eacea um BRIP 72301a T MZ303809 MZ344213 MZ344194 P asanna h e al. (2021a, 2021b)
Cl. p uni-salicinae GUCC 21206.1 T OP852683 OP863092 OP859041 Yang e al. (2023)
Cl. pseudocladospo ioides CBS 125993 T HM148158 HM148647 HM148402 Bensch e al. (2010)
Cl. pseudochalas opo oides CBS 140490 T KT600415 KT600611 KT600513 Bensch e al. (2015)
Cl. pseudo enuissimum COAD 2266 T MZ318439 MT373132 MT680211 Cos a e al. (2022)
Cl. punicae GUCC 21271.5 T OP852672 OP863108 OP859056 Yang e al. (2023)
Cl. pu is COAD 2487 T MK253337 MK249980 MK293777 F ei as e al. (2021)
Cl. queenslandicum BRIP 72447a T OL307928 OL332736 OL332735 Tan e al. (2022)
Cl. queenslandicum BRIP 72452a OL307929 OL332738 OL332737 Tan e al. (2022)
Cl. queenslandicum BRIP 72455a OL307931 OL332741 OL332740 Tan e al. (2022)
Cl. ec oides CBS 125994 T HM148193 HM148683 HM148438 Bensch e al. (2010)
Cl. ibis GUCC 21244.1 T OP852666 OP863098 OP859046 Yang e al. (2023)
Cl. ub um CMG 28 T MN053018 MN066639 MN066644 Vicen e e al. (2021)
8
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Species Vouche in o ma ion ITS ac e 1-α Re e ences
Cl. ugulo a ians CBS 140495 T KT600459 KT600656 KT600558 Bensch e al. (2015)
Cl. u hsange ae BRIP 75808a T OR290122 OR335736 OR335741 Tan and Shi as (2023b)
Cl. scab ellum CBS 126358 T HM148195 HM148685 HM148440 Bensch e al. (2010)
Cl. se oides COAD 3470 T OP535379 OP598131 OP676090 Pe ei a e al. (2024)
Cl. se oides COAD 2576 OP535380 OP598132 OP676091 Pe ei a e al. (2024)
Cl. silenes CBS 109082 T EF679354 EF679506 EF679429 Schube e al. (2007)
Cl. sinua um CGMCC 3.18096 T KX938385 KX938368 KX938402 Bensch e al. (2018)
Cl. speluncae COAD 3116 T ON062329 ON141934 ON982818 Du a e al. (2023)
Cl. spla iae BRIP 75807a OR290120 OR335734 OR335739 Tan and Shi as (2023b)
Cl. sphae ospe mum CBS 193.54 DQ780343 EF101380 EU570261 C ous e al. (2021b)
Cl. s ipag os idicola CBS 146978 T MZ064420 MZ078146 MZ078223 Bensch e al. (2010)
Cl. subuli o me CBS 126500 T HM148196 HM148686 HM148441 Bensch e al. (2010)
Cl. enuissimum CBS 125995 T HM148197 HM148687 HM148442 Bensch e al. (2018)
Cl. ianshanense CGMCC 3.18033 T KX938381 KX938364 KX938398 B aun e al. (2003); Bensch e al. (2010)
Cl. u edinicola ACC 46649 AY251071 HM148712 HM148467 Bensch e al. (2018)
Cl. uweb aunianum CBS 143365 T MF473306 MF474156 MF473729 Bensch e al. (2010)
Cl. a ians CBS 126362 T HM148224 HM148715 HM148470 Bensch e al. (2010)
Cl. e ucocladospo ioides CBS 126363 T HM148226 HM148717 HM148472 Bensch e al. (2018)
Cl. icinum CBS 143366 T MF473311 MF474161 MF473734 Bensch e al. (2018)
Cl. ignae CBS 121.25 HM148227 HM148718 HM148473 Ma in-Felix e al. (2017)
Cl. welwi schiicola CPC 18648 T KY646223 KY646226 KY646229 Yang e al. (2023)
Cl. wenganense GUCC 21220.1 T OP852682 OP863101 OP859049 Bensch e al. (2018)
Cl. wes e dijkiae CBS 113746 T HM148061 HM148548 HM148303 Sando al-Denis e al. (2016)
Cl. xan hoch oma icum CBS 140691 T LN834415 LN834599 LN834511 Bensch e al. (2010)
Cl. xylophilum CBS 125997 T HM148230 HM148721 HM148476 Xu e al. (2021b)
Cl. yunnanensis KUN-HKAS 121704 T OK338502 OL466937 OL825680 Zala e al. (2007); Dugan e al. (2008)
Cl. longissimum CBS 300.96 T DQ780352 EF101385 EU570259 Zala e al. (2007); Dugan e al. (2008)
T Type ma e ial. New species we e shown in bold.
Table 2. Vouche in o ma ion and GenBank accession numbe s o he axa used in he Moelle iella.
Species Vouche in o ma ion LSU pb1 e 1-α Re e ences
Hypoc ella ci ina P.C. 606 EU392556 EU392640 EU392694 Cha e i e al. (2008)
H. ci ina P.C. 597 AY986905 AY986930 DQ000331 Cha e i e al. (2005a)
H. discoidea BCC2097 AY986945 DQ000346 Cha e i e al. (2005a)
H. discoidea I93-901d EU392567 EU392646 EU392700 Cha e i e al. (2008)
H. disci o mis P.C. 655 EU392560 EU392643 EU392697 Cha e i e al. (2008)
H. disci o mis P.C. 676 EU392566 EU392645 EU392699 Cha e i e al. (2008)
H. hi su a P.C. 436.2 AY986922 AY986949 DQ000350 Cha e i e al. (2005a)
H. hi su a P.C. 543 EU392569 EU392648 EU392702 Cha e i e al. (2008)
H. i idans P.C. 635 EU392572 EU392651 EU392705 Cha e i e al. (2008)
H. i idans I89-490 EU392570 EU392649 EU392703 Cha e i e al. (2008)
Moelle iella a icana P.C. 736 AY986917 AY986943 DQ000344 Cha e i e al. (2005a)
M. alba BCC49492 JQ269645 KX254424 JQ256905 Mongkolsam i e al. (2015)
M. alba BCC49409 T JQ269646 KX254423 JQ256906 Mongkolsam i e al. (2015)
M. basicys is P.C. 374 AY986903 AY986928 DQ000329 Cha e i e al. (2005a)
9
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Species Vouche in o ma ion LSU pb1 e 1-α Re e ences
M. basicys is F183147 EU392577 EU392653 Cha e i e al. (2008)
M. boehme iae SICAUCC 25-0061 T PV124784 PV153478 PV153492 In his s udy
M. boehme iae SICAUCC 25-0062 PV124785 PV153479 PV153493 In his s udy
M. boli iensis P.C. 603 AY986923 AY986950 DQ000351 Cha e i e al. (2005a)
M. chiangmaiensis BCC18029 T MT659360 MW091560 Khonsani e al. (2021)
M. chiangmaiensis BBH33051 MT659362 MT672277 MT672269 Khonsani e al. (2021)
M. chiangmaiensis BCC60941 MT659361 MT672278 MT672270 Khonsani e al. (2021)
M. chumphoensis BCC47575 JQ269648 KX254422 JQ256908 Mongkolsam i e al. (2015)
M. chumphoensis BCC47574 T JQ269647 KX254421 JQ256907 Mongkolsam i e al. (2015)
M. cinnamomum SICAUCC 25-0067 T PV124788 PV153484 PV153494 In his s udy
M. cinnamomum SICAUCC 25-0068 PV124789 PV153485 PV153495 In his s udy
M. ci us SICAUCC 25-0059 T PV124782 PV153476 PV153490 In his s udy
M. ci us SICAUCC 25-0060 PV124783 PV153477 PV153491 In his s udy
M. disjunc a J.B. 205 EU392578 EU392654 Cha e i e al. (2008)
M. epiphylla P.C. 545 EU392585 EU392660 EU392711 Cha e i e al. (2008)
M. epiphylla I93-813 EU392583 EU392656 EU392707 Cha e i e al. (2008)
M. eucalyp i SICAU 25-0072 T PV124778 PV153470 PV153486 In his s udy
M. eucalyp i SICAU 25-0073 PV124779 PV153471 PV153487 In his s udy
M. eucalyp i SICAU 25-0074 PV124780 PV153472 PV153488 In his s udy
M. eucalyp i SICAU 25-0075 PV124781 PV153473 PV153489 In his s udy
M. e ansii P.C. 627 T AY986916 AY986942 DQ000343 Cha e i e al. (2005a)
M. la a BCC60924 T KF951146 KX254430 MT672271 Khonsani e al. (2021)
M. la a BCC60925 KF951147 KX254431 MT672272 Khonsani e al. (2021)
M. la a BCC60929 KX298238 KX254432 MT672273 Khonsani e al. (2021)
M. la a BCC60930 KX298237 KX254429 Khonsani e al. (2021)
M. globos oma a YFCC 22109275 T OR828408 OR831942 OR831952 Wang e al. (2025)
M. globos oma a YHH 221010 OR828403 OR831940 OR831950 Wang e al. (2025)
M. g acilispo a CGMCC 3.18989 T KC964202 KC964191 KC964179 Yuan e al. (2020)
M. g acilispo a CGMCC 3.18990 KC964203 KC964192 KC964180 Yuan e al. (2020)
M. hainanensis YHH 2303020 OR828400 OR831938 OR831948 Wang e al. (2025)
M. hainanensis YFCC 23039277 T OR831939 OR831949 Wang e al. (2025)
M. inspe a a ARSEF 2396 T AY518374 DQ070029 EU392713 Cha e i e al. (2008)
M. jinghongensis YFCC 23089312 T OR854253 OR837093 Wang e al. (2025)
M. jinghongensis YHH 2308025 OR828411 OR854254 OR837094 Wang e al. (2025)
M. jinuoana YHH MJBP2309031 T PP178643 PP776170 PP776160 Wang e al. (2024)
M. jinuoana YHH MJBP2309032 PP178644 PP776171 PP776161 Wang e al. (2024)
M. jinuoana YFCC MJBP23099451 PP178645 PP776172 PP776162 Wang e al. (2024)
M. kanchanabu iensis BCC75979 MT659363 MT672279 MT843900 Khonsani e al. (2021)
M. kanchanabu iensis BCC75980 MT659364 MT672280 MT843901 Khonsani e al. (2021)
M. kanchanabu iensis BCC75981 T MT659365 MT672281 Khonsani e al. (2021)
M. libe a P.C. 444 EU392591 EU392662 EU392714 Cha e i e al. (2008)
M. libe a P.C. 445 AY986900 AY986925 DQ000326 Cha e i e al. (2005a)
M. longzhuensis YHH MLFSL2310012 T PP178646 PP776173 PP776163 Wang e al. (2024)
M. longzhuensis YHH MLFSL2310013 PP178647 PP776174 PP776164 Wang e al. (2024)
M. longzhuensis YFCC-MLFSL23109453 PP776175 PP776165 Wang e al. (2024)
M. mac os oma P.C. 605 T AY986919 AY986946 DQ000347 Cha e i e al. (2005a)
16
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Figu e 2. Phylogene ic ee based on maximum likelihood analysis o he combined LSU, pb1, and e 1-α sequences
wi hin he Cla icipi aceae. Numbe s abo e he nodes indica e MLBS alues (≥ 60%, le ) and BIPP alues (≥ 0.95, igh ),
wi h lowe alues deno ed as “–”. The ee is oo ed o Pu pu eocillium lilacinum (CBS 431.87, CBS 284.36). Ex- ype
s ains a e highligh ed in bold, and isola es om he p esen s udy a e shown in ed.

17
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Figu e 3. Phylogene ic ee based on maximum likelihood analysis o he combined ITS, LSU, SSU, e 1-α, and pb2 se-
quences wi hin he Pleospo ales. Numbe s abo e he nodes ep esen MLBS alues (≥ 60%, le ) and BIPP alues (≥ 0.95,
igh ), wi h lowe alues deno ed as “–”. The ee is oo ed o Tubeu ia ja anica (MFLUCC 12-0545) and T. chiangmaiensis
(MFLUCC 11-0514). Ex- ype s ains a e highligh ed in bold, and isola es om he p esen s udy a e shown in ed.
18
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Taxonomy
Cladospo ium kuwanaspidis X.L. Xu & C.L. Yang, sp. no .
MycoBank No: MB 858369
Fig. 4
E ymology. In e e ence o he gene ic name o he associa ed scale insec
(Kuwanaspis howa di).
Diagnosis. Simila o Cladospo ium pe angus um in ha ing conidiopho es o
compa able size, bu C. kuwanaspidis di e s by i s unb anched conidiopho es
and la ge amoconidia.
Type. CHINA • Sichuan P o ince, Meishan Ci y, Hongya Coun y. In ec -
ed scale insec s (Kuwanaspis howa di) we e ound on he culms o bam-
boo (Pleioblas us ama us (Keng) P. C. Keng), 29°41.88'N, 103°14.04'E, al .
540 m, 13 Ma . 2021, C.L. Yang, YCL202103004 (SICAU 25-0082 – holo ype
p ese ed in he He ba ium o Sichuan Ag icul u al Uni e si y; li ing cul u e
SICAUCC 25-0063 – ex-holo ype s o ed in he Cul u e Collec ion in Sichuan
Ag icul u al Uni e si y).
Desc ip ion. Pa asi ic on scale insec om Pleioblas us ama us (Poaceae).
Sexual mo ph: No obse ed. Asexual mo ph: Hyphomyce ous. Mycelium
supe icial and imme sed, wi h abundan spo ula ion on he su ace o scale
insec . Conidiopho es e ec , ascicula e, usually mac onema ous, cylind ical,
subnodulose o nodulose, genicula e, sep a e, unb anched, pale b own o
b own, sligh ly oughened o e uculose, hick-walled, and p onounced loci,
45–120 × 3.5–6 μm. Conidiogenous cells e minal o in e cala y, cylind ical,
some imes sinuous, p oli e a ion sympodial, 14–25(–80) × 2.5–6.5 μm, conid-
iogenous loci a he apex (2–5) o in la e al shoulde s (0–2). Ramoconidia oli e
o b own, sep a e o asep a e, ellipsoidal o subcylind ical, smoo h- and hick-
walled, 6–13.5 × 3–5.5 μm. Seconda y amoconidia oblong, pale b own, 0–1
sep a e, 2–4 dis al hila, 5.5–9 × 2.5–5 μm. Conidia nume ous, ca ena e, o m-
ing sho b anched chains, asep a e, oli e o b own, smoo h- and hin- walled,
ellipsoid-o oid, obo oid, 2–7.5 × 2–4 μm. In e cala y conidia limoni o m, o al
o ellipsoid, wi h hila p o ube an , 3.5–7.5 × 2.5–4 μm. Te minal conidia glo-
bose o ellipsoid, some imes hila e iden , 2–4 × 2–3.5 μm. Mic ocyclic conidio-
genesis absen . In i o on SNA: Mycelium abundan , subme ged, o e g owing
whole cul u e dishes, hyphae s aigh o sligh ly sinuous, sep a e and b anched,
oli e o b own, and hick-walled, 2–3.5 µm wide. Conidiopho es e ec , occa-
sionally b anched, b own, hick-walled, 33–118 × 2.5–4 µm. Ramoconidia ol-
i e o b own, na owly ellipsoid o cylind ical-oblong, subcylind ical, sep a e o
asep a e, smoo h- and hick-walled, 6–17.5 (–20) × 2–5 µm. Conidia in simple
and b anched ac ope al chains, ligh oli e, asep a e, smoo h- and hin-walled,
ellipsoido oid, obo oid, usi o m, 3–6.5 × 2–4 µm. In e cala y conidia o al o
ellipsoid, wi h hila p o ube an , 3.5–6.5 × 2.5–4 μm. Te minal conidia globose
o ellipsoid, 2.5–4 × 2–4 μm.
Cul u e cha ac e is ics. Conidia o all ypes can ge mina e wi hin 12 h in
s e ile dis illed wa e , wi h ge mina ion ubes o ming om any pa o he co-
nidial body. Colonies on MEA a aining 40–50 mm diame e , a e 1 week a
25 °C, g eyish-g een o g eyish owa d egula ma gins, el e y, spo ula ion p o-
use, adially u owed, ae ial mycelium abundan , e e se da k g een. Colonies
19
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
on PDA a aining 35–40 mm diame e , a e 1 week a 25 °C, g ey-oli aceous o
i on g ey, pale g eenish g ey owa d egula ma gins, e e se da k g een. Colo-
nies on SNA a aining 20–30 mm diame e , a e 1 week a 25 °C, smoke-g ey o
oli aceous-g ey, ae ial mycelium loose, e e se oli aceous o b own.
Hos . Kuwanaspis howa di (Diaspididae).
Habi a . Sub opical e e g een b oad-lea ed o es s, pa icula ly bamboo
o es s, a e common habi a s. Kuwanaspis howa di o en causes damage in
he lowe canopy o bamboo o es s. Occasionally, Cladospo ium kuwanaspidis
can be obse ed on bamboo culms. Scale insec s a e mo e easily ound in
shaded and shel e ed a eas.
Dis ibu ion. China, Sichuan P o ince, Meishan Ci y.
Figu e 4. Cladospo ium kuwanaspidis. A–C. Symp oms and appea ance o colonies obse ed on Kuwanaspis howa di.
D–G. Conidiopho es. H. Ramoconidia (whi e a ows), seconda y amoconidia ( ed a ows), and e minal conidia (yellow
a ows). I–K. Colonies on MEA, PDA, and SNA o 7 days. Scale ba s: 200 µm (B, C); 10 µm (D–H).
20
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Ma e ial examined. CHINA • Sichuan P o ince, Meishan Ci y, Hongya Coun-
y. In ec ed scale insec s (Kuwanaspis howa di) we e ound on he culms o
bamboo (Pleioblas us ama us), 29°41.88'N, 103°14.04'E, al . 540 m, 9 Ma .
2021, C.L. Yang, YCL202103004 (li ing cul u e SICAUCC 25-0063); • ibid.
YCL202103004-1 (SICAU 25-0083, li ing cul u e SICAUCC 25-0064).
No es. The ITS base-pai compa ison be ween Cladospo ium kuwanaspidis
(SICAUCC 25-0063) and he phylogene ically a ilia ed ex- ype cul u e o
C. pe angus um (CBS 125996) shows no nucleo ide di e ences. Howe e , nucle-
o ide di e ences in he ac and e 1-α egions a e 1.86% (4 bp, 0 gap) and 4.01%
(15 bp, 2 gaps), espec i ely. Mo phologically, C. pe angus um has la ge amoco-
nidia compa ed o hose obse ed in C. kuwanaspidis on bo h he hos and SNA
medium (25–45 µm s. 6–13.5 µm and 6–20 µm, espec i ely). In addi ion, com-
pa ed o he leng h o in e cala y conidia (4–19 µm) obse ed in C. pe angus um
(CBS 125996), ou isola es exhibi sho e in e cala y conidia on bo h he hos and
PDA medium (3.5–7.5 µm and 3.5–6.5 µm, espec i ely) (Bensch e al. 2010).
Cladospo ium guizhouense S.Y. Wang, Yong Wang bis & Y. Li, MycoKeys 91:
151–168 (2022)
MycoBank No: MB 842407
Fig. 5
Desc ip ion. Pa asi ic on aphids om Telosma co da a (Bu m. .) Me .
(Apocynaceae). Sexual mo ph: No obse ed. Asexual mo ph: Hyphomyce ous.
Mycelium supe icial and imme sed, wi h abundan spo ula ion on he su ace
o aphids. Conidiopho es e ec , ascicula e, mac onema ous, cylind ical, sub-
nodulose o nodulose, genicula e, sep a e, b anched, pale b own, sligh ly ough-
ened o e uculose, hick-walled, and p onounced loci, 111–367 × 3.5–5 μm.
Conidiogenous cells in eg a ed, e minal o in e cala y, cylind ical, cylind i-
cal-oblong, some imes genicula e, 20–65 × 3–6 μm, conidiogenous loci a he
apex (2–5) o in la e al shoulde s (0–4). Ramoconidia oli e-g een, 0–4-sep a e,
ellipsoidal o subcylind ical, smoo h- and hick-walled, 10–26 × 3.5–7.5 μm.
Seconda y amoconidia pale b own, oblong, oblong-ellipsoid, 0–1sep a e, 1–4
dis al hila, 8.5–13.5 × 3.5–7.5 μm. Conidia nume ous, ca ena e, o ming sho
b anched chains, asep a e, oli e o ligh oli e, smoo h- and hin-walled, a iable
in size and shape, ellipsoid-o oid, obo oid, and usi o m, 4–10 × 3–6.5 μm.
In e cala y conidia asep a e, oli e o ligh oli e, ellipsoid-o oid, usi o m, 6–10
× 4–6.5 μm. Te minal conidia asep a e, oli e o ligh oli e, obo oid, 4–6.5 ×
3–5.5 μm. Mic ocyclic conidiogenesis absen . In i o on SNA: Mycelium abun-
dan , subme ged, o e g owing whole cul u e dishes, hyphae s aigh o sligh ly
sinuous, sep a e and b anched, ligh oli e-g een o oli e-g een, and hick-walled,
1.5–4.5 µm wide. Conidiopho es e ec , b anched, ligh oli e-g een, hick-walled,
43–163 × 2.5–4 µm. Ramoconidia pale oli eg een, na owly ellipsoid o cylin-
d ical-oblong, subcylind ical, sep a e o asep a e, smoo h- and hick-walled,
6–27 × 3–5 µm. Conidia in simple and b anched ac ope al chains, ligh oli e,
asep a e, smoo h- and hin-walled, a iable in size and shape, ellipsoid-o oid,
obo oid, usi o m, 3–8 × 2.5–4 µm. In e cala y conidia ligh oli e, asep a e,
smoo h- and hin-walled, mos ly ellipsoid, 3.5–8 × 2.5–4 µm. Te minal conidia
ligh oli e, asep a e, smoo h- and hin-walled, mos ly o oid, 3–4.5 × 2.5–3.5 µm.
21
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Cul u e cha ac e is ics. All conidia can ge mina e wi hin 12 h in s e ile
dis illed wa e , wi h ubes o ming om any pa . Colonies on MEA a aining
35–45 mm diame e , a e 1 week a 25 °C, da k g een, whi e o g eyish owa d
egula ma gins, el e y, spo ula ion p o use, adially u owed, ae ial mycelium
abundan , e e se da k g een. Colonies on PDA a aining 30–40 mm diame e ,
a e 1 week a 25 °C, g eyishg een, g eyish owa d i egula ma gins, el e y,
spo ula ion p o use, adially u owed, ae ial mycelium abundan , e e se da k
g een. Colonies on OA a aining 30–40 mm diame e , a e 1 week a 25 °C,
g eyish oli aceous, whi e owa d i egula ma gins, spo ula ion p o use, e e se
oli aceous. Colonies on SNA a aining 25–30 mm diame e , a e 1 week a
25 °C, oli aceous, la , whi e owa d egula and wide ma gins, ae ial mycelium
loose, e e se oli aceous.
Hos . Aphids (Aphididae), lea es o plan s, and u edinia o Hemileia as a ix
(Pucciniaceae).
Figu e 5. Cladospo ium guizhouense. A. Symp oms obse ed on hos . B, C. Appea ance o colonies on aphids.
D–H. Conidiopho es. I. Ramoconidia (whi e a ows), seconda y amoconidia ( ed a ows), and e minal conidia (yellow
a ows). J–M. Colonies on MEA, PDA, OA, and SNA o 7 days. Scale ba s: 1000 µm (B); 200 µm (C); 20 µm (D–I).

22
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Habi a . In opical and sub opical egions, i can in ec insec s, ac sap o-
phy ically on plan issues, and unc ion as a ungicolous ungus on he u edin-
ia o us . Du ing he middle and la e s ages o aphid damage, nume ous aphids
die on he unde side o plan lea es, becoming en eloped in mycelial issue.
Dis ibu ion. China, Sichuan P o ince, Guangan Ci y. China, Guizhou P o ince,
Guiyang Ci y. E hiopia, O omia Region, Illubabo Go e. B azil, Minas Ge ais, Viçosa.
Ma e ial examined. CHINA • Sichuan P o ince, Guangan Ci y, Yuechi Coun y.
In ec ed aphids we e ound on he unde side o he lea es o Telosma co da a,
30°44.18"N, 106°30.80"E, al . 480 m, 31 Jan. 2020, X.L. Xu, XXL202001003
(SICAU 25-0076, li ing cul u e SICAUCC 25-0057); • ibid. XXL202001003-1
(SICAU 25-0077, li ing cul u e SICAUCC 25-0058).
No es. Phylogene ically, ou collec ions g ouped wi h isola es o Cladospo-
ium guizhouense wi hin he C. cladospo ioides complex, showing s ong s a-
is ical suppo (92% MLBS, 0.97 BIPP) (Fig. 1). A o al o 6 bp nucleo ide di -
e ences we e obse ed be ween ou isola e SICAUCC 25-0057 and he ex- ype
cul u e o C. guizhouense (GUCC 401.7), wi h 0 bp in ITS, 2 bp in ac , and 4 bp in
e 1-α (Wang e al. 2022b). Mo phologically, ou isola e is ully consis en wi h
C. guizhouense, showing no no iceable di e ences (Wang e al. 2022b; Pe ei a
e al. 2024).
Moelle iella eucalyp i X.L. Xu, Feng Liu & C.L. Yang, sp. no .
MycoBank No: MB 858370
Fig. 6
E ymology. In e e ence o he gene ic name o hos plan s.
Diagnosis. Simila o Moelle iella sinensis in ha ing somewha simila s o-
ma a, M. eucalyp i di e s by ha ing longe and wide pa aphyses, sho e and
wide conidiogenous cells, and wide conidia.
Type. CHINA • Sichuan P o ince, Dazhou Ci y, Heibaoshan Na ional Fo es
Pa k. In ec ed scale insec s we e ound on he unde side o lea es o Euca-
lyp us sp. (My aceae), 31°55.30'N, 107°47.75'E, al . 690 m, 21 Jul. 2022, Feng
Liu, LF202207001A (SICAU 25-0072 – holo ype p ese ed in he He ba ium o
Sichuan Ag icul u al Uni e si y).
Desc ip ion. Pa asi ic on scale insec s ound on eucalyp us lea es. Sexual
mo ph: No obse ed. Asexual mo ph: Coelomyce ous. S oma a yellow o da k
o ange in esh specimens, pale yellow o whi e in old, en i ely co e ing he insec
hos s, hickened pul ina e, umbona e o hemisphe ic, co ony, sessile, globose
o subglobose, ube cules on he su ace, 1.5–2.5 mm in diame e , 1–2 mm in
high. Conidioma a o i ice sca e ed o ci cula ly a anged, o al o elonga e lask
shaped, na ow o i ices, 120–460 × 80–155 μm. Phialides hyaline, s ick-shaped
o cylind ical, 10–18 × 1–2.5 μm. Conidia hyaline, usi o m, yellow conidial mass-
es, 8–12 × 1.5–3 μm. Pa aphyses p esen , hyaline, ili o m, 90–180 × 1.0–1.6 μm.
Cul u e cha ac e is ics. No ge mina ion was obse ed due o he specimens
being d ied and he cul u e no being ob ained.
Hos . Scale insec s (Coccidae).
Habi a . Sub opical monsoon e e g een b oad-lea o es . Scale insec s
we e in ec ed on he unde side o eucalyp us lea es.
Dis ibu ion. China, Sichuan P o ince, Dazhou Ci y.
23
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Ma e ial examined. CHINA • Sichuan P o ince, Dazhou Ci y, Heibaoshan Na-
ional Fo es Pa k. In ec ed scale insec s we e ound on he unde side o lea es
o Eucalyp us sp., 31°55.30'N, 107°47.75'E, al . 690 m, 21 Jul. 2022, Feng Liu,
LF202207001A (SICAU 25-0072); • ibid. LF202207001B (SICAU 25-0073); • ibid.
LF202207001C (SICAU 25-0074); • ibid. LF202207001D (SICAU 25-0075).
No es. Phylogene ically, ou collec ions a e closely ela ed o Moelle iella sin-
ensis, showing ew sequence di e ences wi h he ype s ain CGMCC3.18911,
speci ically 1 bp in LSU (0.12%, 0 gap) and 8 bp in pb1 (1.10%, 0 gap), while
he e 1-α sequence is in alid. Moelle iella sinensis was i s desc ibed on
whi e ly nymphs (Hemip e a) and la e ound on scale insec s (Coccidae) in
Thailand (Chen e al. 2020; Khonsani e al. 2021). The desc ibed specimens
o M. sinensis a e p ima ily cha ac e ized by la o umbona e, pale yellow
pul ina e s oma a and usi o m conidia. Howe e , ou collec ions di e om
M. sinensis in ha ing pul ina e s oma a ha a e yellow o da k o ange when
esh, pale yellow o whi e when old, and hicke in appea ance. Addi ionally,
hey ha e longe and wide pa aphyses (90–180 × 1.0–1.6 μm s. 43.2–68.9 ×
0.6–0.8 μm), sho e and wide conidiogenous cells (10–18 × 1–2.5 μm s. up
Figu e 6. Moelle iella eucalyp i. A. Fungus-in ec ed scale insec on he unde side o lea es. B–D. S oma a con aining
conidioma a wi h conidial masses. E–I. Sec ion o s oma showing conidioma. J–M. Phialides and conidia a he ips
wi h pa aphyses. N. Conidia. Scale ba s: 500 mm (B–D); 200 μm (E, F); 50 μm (H, I); 10 μm (J, K).
24
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
o 30 μm × 0.8–1.3 μm), and wide conidia (1.5–3 μm s. 1.3–1.8 μm). Hence,
we in oduce M. eucalyp i as a new species, based on he dis inc mo phologi-
cal di e ences obse ed.
Moelle iella boehme iae X.L. Xu & C.L. Yang, sp. no .
MycoBank No: MB 858372
Fig. 7
E ymology. In e e ence o he gene ic name o hos plan s.
Diagnosis. Simila o Moelle iella jinuoana in ha ing yellow o o ange, glo-
bose s oma a wi h a na ow hypo hallus, M. boehme iae di e s by i s slowe
g ow h on PDA, unique colony cha ac e is ics, and wide conidia.
Type. CHINA • Sichuan P o ince, Leshan Ci y, Muchuan Coun y. In ec ed
scale insec s we e ound on he s ems o Boehme ia spica a (Thunb.) Thunb.,
28°47.91'N, 103°55.63'E, al . 900 m, 12 Ma . 2021, C.L. Yang, YCL202103003
(SICAU 25-0080 – holo ype p ese ed in he He ba ium o Sichuan Ag icul u al
Uni e si y; li ing cul u e SICAUCC 25-0061 – ex-holo ype s o ed in he Cul u e
Collec ion in Sichuan Ag icul u al Uni e si y).
Desc ip ion. Pa asi ic on scale insec om Boehme ia spica a (U icaceae).
Sexual mo ph: No obse ed. Asexual mo ph: Coelomyce ous. S oma a yel-
low o o ange in esh specimens, en i ely co e ing he insec hos s, hickened
pul ina e, globose, ube cules on he su ace, closely agg ega ed, some wi h
na ow hypo hallus, 0.8–2.5 mm diame e . Hyphae o s oma a o m compac
ex u a epide moidea. Conidioma a simple dep essions o su ace, ound o
i egula shape, no ma u e conidioma a obse ed.
Cul u e cha ac e is ics. The conidia ge mina e wi hin 12 h in s e ile dis illed
wa e , wi h ge m ubes ex ending om bo h ends o he conidia. Colonies on
PDA slowg owing, a aining a diame e o 5–7 mm in 28 days a 25 °C. Col-
onies compac pul ina e, su ace elu inous, pale yellow o yellow. Conidial
masses usually abundan , yellow. Re e se o colony pale o ange. Conidioma a
U-shaped, subci cula o ci cula . Conidia unicellula , hyaline, smoo h, usoid
wi h ounded ends, 9.5–12.5 × 3–4 µm. No pa aphyses we e obse ed.
Hos . Scale insec s (Coccidae).
Habi a . Sub opical monsoon e e g een b oad-lea o es s se e as he hab-
i a o Moelle iella boehme iae, which can in ec scale insec s ound on he
s ems o Boehme ia spica a. In ec ions a e ypically obse ed a lowe heigh s,
close o he g ound.
Dis ibu ion. China, Sichuan P o ince, Leshan Ci y.
Ma e ial examined. CHINA • Sichuan P o ince, Leshan Ci y, Muchuan Coun-
y. In ec ed scale insec s we e ound on he s ems o Boehme ia spica a,
28°47.91'N, 103°55.63'E, al . 900 m, 12 Ma . 2021, C.L. Yang, YCL202103003
(SICAU 25-0080, li ing cul u e SICAUCC 25-0061); • ibid. YCL202103003-1 (SI-
CAU 25-0081, li ing cul u e SICAUCC 25-0062).
No es. Phylogene ic analysis e ealed ha Moelle iella boehme iae clus e s
wi hin he Globose clade and is closely ela ed o M. jinuoana (Fig. 2), sha -
ing cha ac e is ics such as yellow o o ange, globose s oma a wi h a na ow
hypo hallus (Wang e al. 2024). Al hough de ailed mo phological compa i-
sons we e limi ed due o he absence o ma u e conidioma a in ou specimen,
25
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
M. boehme iae di e s by i s slowe g ow h on PDA and dis inc colony cha ac-
e is ics. The colonies o M. boehme iae a e yellow on he on and ligh o ange
on he e e se, whe eas hose o M. jinuoana a e pale o ange o o ange on he
on and b ownish on he e e se. Addi ionally, M. boehme iae has wide co-
nidia (3–4 µm) obse ed in cul u e compa ed o M. jinuoana, which has conid-
ia measu ing 2.1–2.9 µm on he subs a e. In addi ion, M. boehme iae di e s
om he ype s ain M. jinuoana (YHH MJBP2309031) by 16 bp (1.76%, 0 gap)
in he e 1α egion, 2 bp (0.28%, 0 gap) in he pb1 egion, and 4 bp (0.47%, 2
gaps) in he LSU egion, espec i ely.
Moelle iella cinnamomum X.L. Xu & C.L. Yang, sp. no .
MycoBank No: MB 858373
Fig. 8
E ymology. In e e ence o he gene ic name o hos plan s.
Figu e 7. Moelle iella boehme iae. A, B. Imma u e s oma a o e scale insec s. C. Conidioma o med in PDA. D. Phialides wi h
conidia a he ips. E–G. Phialides wi h de eloping conidia. H. Conidia. I. Ge mina ing conidia. J, K. Colonies ob e se and
e e se on PDA a 25 °C a e 3 weeks. L. Conidial mass on PDA. Scale ba s: 1000 µm (B); 50 µm (C); 20 µm (D); 10 µm (E–I).
32
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
and u edinia o Hemileia as a ix on lea es o Co ea a abica om A ica (Pe ei-
a e al. 2024). This is he i s epo o C. guizhouense om aphids on Telosma
co da a (Apocynaceae). The scale insec Kuwanaspis howa di is commonly dis-
ibu ed and ha m ul o bamboo. In ou p e ious s udy, wo Podonec ia species
and a Mic oce a species associa ed wi h na i e bamboo plan s we e disco e ed
(Xu e al. 2021a). Cladospo ium kuwanaspidis is he second en omopa hogenic
species ound on bamboo (Pleioblas us ama us) in ou in es iga ions. As en o-
mopa hogenic C. cladospo ioides is a po en ial candida e o biocon ol agains
whi e lies and aphids (Islam e al. 2019; Nicole i e al. 2024), and as endophy ic
Cladospo ium sp. has demons a ed insec icidal ac i i y o α-glucosidase inhib-
i o s agains Spodop e a li u a in i o (Singh e al. 2015), mo e Cladospo ium
species need o be in es iga ed as po en ial agen s o biocon ol.
The genus Moelle iella (Ascomyco a, Hypoc eales, Cla icipi aceae) was es ab-
lished o accommoda e M. sulphu ea (B esadola 1896) and is cha ac e ized by
globose o py i o m pe i hecia imme sed in a b igh ly colo ed s oma wi h open-
ings p o uding om he s oma on scale insec s (Coccidae, Homop e a) o whi e-
lies (Aley odidae, Homop e a), as well as mul i-sep a e ascospo es ha disa -
icula e a he sep a inside he ascus. In con as , species o Hypoc ella and he
subsequen ly p oposed genus Samuelsia, ha ing Asche sonia-like anamo phs,
ha e ascospo es ha emain whole (Pe ch 1921; Mains 1959; Ki k e al. 2001;
Cha e i e al. 2008). The genus cu en ly has abou 52 eco ds (h ps://www.spe-
cies ungo um.o g/Names/Names.asp, accessed on 4 No embe 2024), and spe-
cies ha e been epo ed in Belize, Boli ia, Cos a Rica, Cô e d’I oi e, China, Ecuado ,
Ghana, Gua emala, Guiana, Hondu as, Jamaica, Mexico, Panama, he Philippines,
Thailand, T inidad, Venezuela, and Vie nam, equen ly om Thailand, indica ing
i is mo e common in opical egions (Liu e al. 2006; Cha e i e al. 2008; Qiu e
al. 2009; Mongkolsam i e al. 2011b, 2015; Li e al. 2016; Tibp omma e al. 2017;
Chen e al. 2020; Yuan e al. 2020; Khonsani e al. 2021). Moelle iella libe a has
been widely used due o i s pa asi ism o la ge popula ions o whi e lies and scale
insec s in he wild (Zhang e al. 2018; Ingle e al. 2019; Qasim e al. 2020). In Chi-
na, M. och acea has p e iously been eco ded om Fujian P o ince on homop e -
an cada e s (Qiu e al. 2009). In ecen yea s, se e al species ha e been epo ed
om he sub opical egions o Yunnan and Fujian p o inces, iz. M. g acilispo a
on whi e ly nymphs (Hemip e a) (Yuan e al. 2020), M. sinensis in ec ing whi e-
ly nymphs (Hemip e a) (Chen e al. 2020), M. pue ensis on whi e lies (Wang e
al. 2022c), M. simaoensis on whi e lies (Yang e al. 2023a), and M. jinuoana and
M. longzhuensis on scale insec s and whi e lies (Wang e al. 2024). Based on
ou obse a ion, ou Moelle iella species we e newly eco ded on scale insec s
inhabi ing Boehme ia spica a, Ci us sp., and Eucalyp us sp. in Sichuan P o ince.
Moelle iella sinensis, M. cinnamomum, and M. ci us we e accommoda ed in he
E use clade (Cha e i e al. 2008; Wang e al. 2024), which is cha ac e ized by e -
use o hin, pul ina e s oma a wi h loose hyphal issue, mos ly ha ing hypo halli.
The new species M. boehme iae was nes ed in he Globose clade (Cha e i e al.
2008), cha ac e ized by globose, da ke s oma a wi h compac issue.
Podonec iaceae was p oposed as a amily in he Pleospo ales o accommo-
da e he genus Podonec ia, which was con i med by phylogene ic analyses (Dao
e al. 2016; Yang e al. 2019b). Podonec ia was adi ionally eco ded wi h a
Te ac ium-like conidial s age (Pe ch 1921; Rossman 1978). P e ious s udies con-
i med he link be ween he sexual mo phs in Podonec ia and he asexual mo phs

33
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
in Te ac ium wi h iden ical molecula sequences (Dao e al. 2015, 2016; Xu e
al. 2021a). In addi ion, an asexual genus, Te anac ium, ypi ied by T. g amineum
(Hudson and Su on 1964), was eco ded as an associa ed anamo ph o Podonec-
ia gahnia on scale insec s. Howe e , al hough he shape and de elopmen o
conidia in he associa ed Te anac ium a e co ela ed wi h Te ac ium, he conidi-
oma a in Te anac ium a e pycnidia, whe eas in Te ac ium hey a e spo odochia.
The ela ionship be ween Te anac ium and Podonec ia equi es u he phyloge-
ne ic and axonomic s udies wi h mo e samples. In his s udy, ou collec ions had
hyphomyce ous anamo phs, which we e consis en wi h Podonec ia, iden i ied
based on he phylogene ic esul s o combined ITS, LSU, SSU, e 1-α, and pb2
da a. Mos species o Podonec ia a e associa ed wi h a mo ed scale insec s on
Ci us spp. Podonec ia mul ia ma a is he hi d en omopa hogenic species asso-
cia ed wi h Kuwanaspis howa di on bamboo (Pleioblas us ama us) in China.
Conclusion
Se en en omopa hogenic ungi om Sichuan P o ince, China, a e desc ibed,
including six new species and one newly eco ded species. These species a e
mo phologically simple hyphomyce es o coelomyce es, wi h de ailed iden-
i ica ion based on mo phological cha ac e is ics and phylogene ic analyses.
These ungi wa an a en ion, pa icula ly hose associa ed wi h majo ag icul-
u al and o es y pes s. We p edic ha sou hwes e n China ha bo s a ich di-
e si y o en omopa hogenic ungi, wi h many species ye o be disco e ed and
e alua ed. These ungi ep esen a c ucial esou ce o u u e d ug de elopmen
and hold signi ican po en ial o pes managemen in ag icul u e and o es y.
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
No applicable.
Funding
This esea ch was inanced by he Sichuan Science and Technology P og am o China
(No. 2024NSFSC1192).
Au ho con ibu ions
C-LY, X-LX, and FL collec ed samples. C-LY, X-LX, and FL isola ed cul u es and pe o med
DNA isola ion and PCR ampli ica ion. C-LY, X-LX, X-YL, and FL analyzed da a. C-LY and
X-LX w o e he o iginal d a . Z-Z, QGX, and Y-GL e iewed and edi ed he d a . All au-
ho s ead and app o ed he inal manusc ip .
34
IMA Fungus 16: e170123 (2025), DOI: 10.3897/ima ungus.16.170123
Chunlin Yang e al.: No el en omopa hogenic ungi in ec ing scale insec s and aphids in China
Au ho ORCIDs
Chunlin Yang h ps://o cid.o g/0000-0002-3854-8835
Xiulan Xu h ps://o cid.o g/0000-0002-6832-5421
Da a a ailabili y
All sequence da a gene a ed o his s udy a e a ailable in GenBank (h ps://www.ncbi.
nlm.nih.go /genbank/).
Re e ences
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bio-con ol agen agains whi e lies and aphids in Egyp . Jou nal o Applied En omol-
ogy 127: 228–235. h ps://doi.o g/10.1046/j.1439-0418.2003.00662.x
Abdollahzadeh J, G oenewald JZ, Coe zee MPA e al. (2020) E olu ion o li es yles in
Capnodiales. S udies in Mycology 95: 381–414. h ps://doi.o g/10.1016/j.simy-
co.2020.02.004
Aku se KS, Maniania NK, Fiaboe KKM e al. (2013) Endophy ic coloniza ion o Vicia
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