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Mo phology and molecula phylogeny e eal new species and
eco ds o Diapo hales om Be ula in Xinjiang, China
Caixia Wang1, Ning Jiang2, Chuli Liu3, ZiYan Xu1, Hailong Lu1,4 , Rong Ma1
1 College o Fo es y and Landscape A chi ec u e, Xinjiang Ag icul u al Uni e si y, U umqi 830052, China
2 Key Labo a o y o Biodi e si y Conse a ion o Na ional Fo es y and G assland Adminis a ion, Ecology and Na u e Conse a ion Ins i u e, Chinese Academy
o Fo es y, Beijing 100091, China
3 The Key Labo a o y o Sil icul u e and Conse a ion o he Minis y o Educa ion, Beijing Fo es y Uni e si y, Beijing 100083, China
4 Fo es y Resea ch Ins i u e o Yili Kazakh Au onomous P e ec u e, Yining 835099, Xinjiang, China
Co esponding au ho : Rong Ma ([email p o ec ed])
Copy igh : © Caixia Wang 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
Xinjiang Uygu Au onomous Region (XUAR), loca ed in he a id hin e land o no hwes
China, se es as impo an subs a es o Be ula species. As he mos widely dis ibu ed
lo al genus o Be ulaceae in Xinjiang, Be ula se es as a i al pionee species due o i s
s ess ole ance and adap a ion o ex eme en i onmen s, playing an i eplaceable ole
in sus aining egional ecosys ems. Du ing disease su eys on Be ula ees in he Tian-
shan and Al ai Moun ains o Xinjiang, we obse ed symp oms o s em and b anch can-
ke s. A o al o 37 ungal s ains we e isola ed and iden i ied, based on mo phological
cha ac e is ics and phylogene ic analyses. As a esul , Cy ospo a al ayensis sp. no . was
p oposed and Co yneum lanci o me, C yp ospo ella be ulae, C . omen ella, Cy. anai ica
and Melanconis g oenlandica a e epo ed o he i s ime in China. In addi ion, Cy o-
spo a sopho iopsis is eco ded o he i s ime on he hos genus Be ula. This s udy
iden i ies po en ial pa hogenic ungi associa ed wi h Be ula, p o iding a ounda ion o
u u e disease managemen and o es heal h esea ch.
Key wo ds: Co yneum, C yp ospo ella, Cy ospo a, Melanconis, molecula phylogeny,
axonomy
In oduc ion
Xinjiang Uygu Au onomous Region (XUAR) is loca ed in he cen al pa o he
Eu asian con inen and ea u es a ypical empe a e con inen al clima e. No a-
bly, i s a id condi ions os e unique biological species. Amongs hese, Be ula
(bi ch ees) a e an impo an plan genus in Xinjiang, p ima ily ep esen ed by
B. pendula and B. ianshanica, which play signi ican ecological oles (Huang e
al. 2020). As key pionee species in he egion, bi ch ees con ibu e g ea ly o
na u al o es conse a ion, ecological es o a ion, sus ainable managemen
and he imp o emen o he egional ecosys em (Dubois e al. 2020).
Howe e , bi ch ees in he egion a e suscep ible o a ious ungal pa ho-
gens. In pa icula , se e al ungal pa hogens ha e been epo ed o cause
b anch canke , dieback and lea spo diseases in bi ch species (I ano á 2014;
Academic edi o : Rajesh Jeewon
Recei ed:
27 Augus 2025
Accep ed:
4 No embe 2025
Published:
18 No embe 2025
Ci a ion: Wang C, Jiang N, Liu C, Xu
ZY, Lu H, Ma R (2025) Mo phology
and molecula phylogeny e eal new
species and eco ds o Diapo hales
om Be ula in Xinjiang, China.
MycoKeys 125: 167–204. h ps://doi.
o g/10.3897/mycokeys.125.169956
MycoKeys 125: 167–204 (2025)
DOI: 10.3897/mycokeys.125.169956
168
MycoKeys 125: 167–204 (2025), DOI: 10.3897/mycokeys.125.169956
Caixia Wang e al.: Diapo hales om Be ula in Xinjiang, China
Fan e al. 2016a, b). Fo ins ance, Melanconis s ilbos oma has been iden i ied
as a causa i e agen o bi ch canke s (Fan e al. 2016a, b), while species o
Diapo he, such as D. be ulae and D. be ulicola, ha e been associa ed wi h bi ch
dieback (Du e al. 2016). Addi ionally, Discula be ulae is known o cause bi ch
lea spo disease (I ano á 2014). Despi e hese indings, sys ema ic s udies on
hese pa hogens ha e no ye been conduc ed in XUAR.
Diapo hales comp ise a di e se g oup o ungi ha colonise a ious ee spe-
cies, whe e hey o en ac as pa hogens in ec ing ba k, lea es and ui s, hough
some species occu as endophy es o sap ophy es (Rossman e al. 2007; Sena-
nayake e al. 2017, 2018; Jiang e al. 2020a; C ous e al. 2023b; Xiao e al. 2023).
In e es ingly, membe s o his o de exhibi a ma ked p e e ence o ha dwood
hos s, pa icula ly wi hin amilies, such as Be ulaceae, Fagaceae and My aceae
(Senanayake e al. 2017; Hyde e al. 2020b). Co espondingly, Be ula, a genus in-
cluding widesp ead ee species, has equen ly been epo ed as a hos associa -
ed wi h Diapo hales (Senanayake e al. 2017; Fan e al. 2018a; C ous e al. 2023a).
Taxa wi hin Diapo hales can be clea ly delinea ed using bo h mo pholog-
ical ea u es and mul i-gene phylogene ic analyses (Senanayake e al. 2017,
2018; Jiang e al. 2025b). On one hand, se e al gene a possess dis inc i e
ai s: o ins ance, Co yneum species p oduce la ge, b own, dis osep a e co-
nidia (Jiang e al. 2018); Cy ospo a species a e cha ac e ised by hyaline, allan-
oid ascospo es and conidia (Fan e al. 2020); and As e ospo ium species a e
ecognised by hei sep a e, b own, s ella e conidia (Senanayake e al. 2017).
On he o he hand, some axa, such as Plagios oma and Pseudoplagios oma,
exhibi o e lapping mo phological ea u es ha complica e hei di e en ia ion
(Senanayake e al. 2017; Jiang e al. 2025b, c).
In his s udy, samples exhibi ing ypical canke symp oms we e collec ed
om Be ula hos s in XUAR. Fungal isola es we e ob ained and iden i ied, based
on mo phological cha ac e is ics and molecula phylogene ic analyses, ollow-
ing con empo a y axonomic amewo ks o he ele an axa. The objec i es
o his s udy we e o cha ac e ise species o Diapo hales associa ed wi h Be -
ula hos s in XUAR and o es ablish links be ween hese ungal species and hei
bi ch hos s o ad ance he unde s anding o canke diseases.
Ma e ials and me hods
Sample collec ion, isola ion and mo phology
Du ing 2023 and 2024, we conduc ed b anch canke disease su eys on hos s
o Be ula in XUAR. Ba ks exhibi ing isible ungal ui ing bodies we e collec ed
o u he analysis. All samples we e s o ed in pape bags and anspo ed o
he labo a o y o ungal isola ion and mo phological cha ac e isa ion.
F ui ing bodies on b anch samples we e examined unde a Zeiss Disco e y V8
s e eomic oscope (Jena, Ge many), sec ioned and ans e ed on o po a o dex-
ose aga (PDA; con aining 200 g po a oes, 20 g dex ose and 20 g aga pe li e)
pla es a 25 °C o ob ain ungal cul u es. Specimens we e deposi ed in he He ba -
ium o Xinjiang Ag icul u al Uni e si y (XJAU), while pu e cul u es we e p ese ed
a he China Fo es y Cul u e Collec ion Cen e (CFCC; h ps://c cc.ca .ac.cn/).
Mo phology was cha ac e ised mainly based on ui ing bodies o ming on
na u al subs a es. S oma a and conidioma a we e hand-sec ioned using a
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MycoKeys 125: 167–204 (2025), DOI: 10.3897/mycokeys.125.169956
Caixia Wang e al.: Diapo hales om Be ula in Xinjiang, China
s e ile double-edged blade and examined unde a Zeiss Disco e y V8 s e eomi-
c oscope. Mic oscopic s uc u es including asci, ascospo es, conidiopho es,
conidiogenous cells and conidia we e obse ed and pho og aphed using an
Olympus BX51 compound mic oscope (Tokyo, Japan).
Phylogene ic analyses
Genomic DNA was ex ac ed om ungal colonies g own on PDA pla es o wo
weeks using he Biospin Fungus Genomic DNA Ex ac ion Ki -BSC14S1 (Bio-
Flux, China), acco ding o he manu ac u e ’s p o ocol. Fo p elimina y ungal
iden i ica ion, he in e nal ansc ibed space (ITS) egion was ampli ied using
p ime pai s ITS1/ITS4 (Whi e e al. 1990). Fu he phylogene ic esolu ion was
achie ed by ampli ying addi ional loci: he la ge subuni n DNA (LSU) o Co y-
neum and Melanconis (p ime s LR0R/LR5); he pa ial ac in (ac ) o Cy ospo a
(p ime s ACT512F/ACT728R); he RNA polyme ase II second la ges subuni
( pb2) gene o Co yneum, Cy ospo a and Melanconis (p ime s RPB2-5F/ RPB2-
7cR); he ansla ion elonga ion ac o 1-alpha ( e 1) gene o Co yneum, C yp-
ospo ella, Cy ospo a and Melanconis (p ime s EF1-728F/EF2; EF688F/1251R)
and he pa ial be a- ubulin ( ub2) gene o C yp ospo ella, Cy ospo a and Mel-
anconis (p ime s B 2a/B 2b) (Vilgalys and Hes e 1990; Glass and Donaldson
1995; Ca bone and Kohn 1999; Liu e al. 1999; Rehne 2001).
Polyme ase chain eac ion (PCR) ampli ica ion was pe o med unde he
ollowing condi ions: ini ial dena u a ion a 94 °C o 5 min; 35 cycles o de-
na u a ion (94 °C, 30 sec), annealing (48 °C o ITS/LSU, 54 °C o e 1/ ub2,
55 °C o pb2 o 58 °C o ac , 50 sec) and ex ension (72 °C, 1 min); ollowed
by a inal elonga ion a 72 °C o 7 min. The sequencing se ice was pe o med
by Sangon Bio ech Company Limi ed (Shanghai, China). Raw o wa d and e-
e se eads p oduced in his s udy we e assembled and edi ed in Seqman .
7.1.0 (DNASTAR Inc., USA) and deposi ed in he NCBI da abase (Tables 1–4).
Mul iple sequence alignmen s we e gene a ed using MAFFT . 7 (Ka oh and
S andley 2013) and manually e ined in MEGA . 7.0.21.
Phylogene ic econs uc ions we e pe o med on conca ena ed sequence
da ase s using bo h Maximum Likelihood (ML) and Bayesian In e ence (BI) ap-
p oaches. The da ase s consis ed o : (1) ITS-LSU- pb2- e 1 o Co yneum, (2)
ITS-ac - pb2- e 1- ub2 o Cy ospo a, (3) ITS- e 1- ub2 o C yp ospo ella and (4)
ITS-LSU- pb2- e 1- ub2 o Melanconis. Fo he ML analysis, we employed he GTR-
GAMMA subs i u ion model and conduc ed 1000 boo s ap eplica es using RAx-
ML-HPC .8, implemen ed h ough he CIPRES Science Ga eway po al (h ps://
www.phylo.o g/). The Bayesian analysis was pe o med using pa i ion-speci ic
e olu iona y models selec ed wi h M ModelTes .2.3, based on he Akaike In o -
ma ion C i e ion (AIC). Ma ko Chain Mon e Ca lo (MCMC) simula ions we e un
in M Bayes .3.1.2 (Ronquis and Huelsenbeck 2003) wi h wo independen uns
o 10 million gene a ions each, s a ing om andom ees. We con i med un
con e gence by moni o ing he a e age s anda d de ia ion o spli equencies (<
0.01) and sampled ees e e y 1000 gene a ions. A e disca ding he i s 25% o
ees as bu n-in, pos e io p obabili ies (PP) we e calcula ed om he emaining
ees. Nodal suppo was assessed using boo s ap suppo (BS) alues (ML)
om 1000 eplica es and Bayesian pos e io p obabili ies (PP). The esul ing phy-
logene ic ees we e isualised using FigT ee .1.4.4 (Rambau 2018).
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Caixia Wang e al.: Diapo hales om Be ula in Xinjiang, China
Resul s
Phylogene ic analyses
The conca ena ed alignmen o ITS, LSU, pb2 and e 1 o Co yneum included
32 s ains and 3,380 cha ac e s (ITS: 1–630; LSU: 631–1,471; pb2: 1,472–2,550;
e 1: 2,551–3,380), wi h gaps e ained. ML analysis yielded an op imal ee (likeli-
hood = -13,031.13), wi h he alignmen exhibi ing 633 dis inc pa e ns and 44.19%
unde e mined cha ac e s o gaps. Nucleo ide equencies we e A = 0.240, C =
0.266, G = 0.283, T = 0.211 and subs i u ion a es we e AC = 1.258, AG = 2.217,
AT = 1.350, CG = 1.193, CT = 7.495, GT = 1.0 (α = 0.140). BI employed models
TNe+G4 (ITS), TN+F+I (LSU), TN+F+G4 ( pb2) and TN+F+G4 ( e 1), wi h esul s
ma ching ML opology. Isola es CFCC 71587 and CFCC 71653 clus e ed obus ly
wi h s ain D215, con i ming hei placemen as Co yneum lanci o me (Fig. 1).
Table 1. GenBank accession numbe s used in he phylogene ic analyses o Co yneum.
Species S ains Hos s GenBank accession numbe s Re e ences
ITS LSU pb2 e 1
Co yneum cas aneicola CFCC 52315 Cas anea mollissima MH683559 MH683551 MH685723 MH685731 Jiang e al. (2018)
Co yneum cas aneicola CFCC 52316 Cas anea mollissima MH683560 MH683552 MH685724 MH685732 Jiang e al. (2018)
Co yneum dep essum D202 Que cus pe aea MH674330 MH674330 MH674334 MH674338 Jiang e al. (2018)
Co yneum gigaspo um CFCC 52319* Cas anea mollissima MH683565 MH683557 MH685729 MH685737 Jiang e al. (2018)
Co yneum gigaspo um CFCC 52320 Cas anea mollissima MH683566 MH683558 MH685730 MH685738 Jiang e al. (2018)
Co yneum gigaspo um G14 Cas anea mollissima MK799957 MK799944 MK799820 MK799830 Jiang e al. (2019)
Co yneum gigaspo um G15 Cas anea mollissima MK799958 MK799945 MK799821 MK799831 Jiang e al. (2019)
Co yneum he eanum MFLUCC 17-0369* He ea b asiliensis MH778707 MH778703 NA MH780881 Senwanna e al. (2018)
Co yneum he eanum MFLUCC 17-0376 He ea b asiliensis MH778708 MH778704 NA NA Senwanna e al. (2018)
Co yneum ilicis CFCC 52994* Ilex pe nyi MK799948 MK799935 NA NA Jiang e al. (2019)
Co yneum ilicis CFCC 52995 Ilex pe nyi MK799949 MK799936 NA NA Jiang e al. (2019)
Co yneum ilicis CFCC 52996 Ilex pe nyi MK799950 MK799937 NA NA Jiang e al. (2019)
Co yneum modonium D203 Cas anea sa i a MH674331 MH674331 MH674335 MH674339 Jiang e al. (2018)
Co yneum lanci o me D215 Be ula pubescens MH674332 MH674332 MH674336 MH674340 Jiang e al. (2018)
Co yneum lanci o me CFCC 71587 Be ula pendula PX227525 PX227507 PX233568 NA In his s udy
Co yneum lanci o me CFCC 71653 Be ula pendula PX227526 PX227508 PX233569 NA In his s udy
Co yneum sinense CFCC 52452* Que cus se a a MH683561 MH683553 MH685725 MH685733 Jiang e al. (2018)
Co yneum sinense CFCC 52453 Que cus se a a MH683562 MH683554 MH685726 MH685734 Jiang e al. (2018)
Co yneum sinense X23 Que cus se a a MK799952 MK799939 MK799816 MK799826 Jiang e al. (2019)
Co yneum sinense X60 Que cus se a a MK799953 MK799940 MK799814 MK799824 Jiang e al. (2019)
Co yneum songshanense CFCC 52997* Que cus den a a MK799946 MK799933 MK799812 MK799822 Jiang e al. (2019)
Co yneum songshanense CFCC 52998 Que cus den a a MK799947 MK799934 MK799813 MK799823 Jiang e al. (2019)
Co yneum su onii CFCC 52317* Cas anea mollissima MH683563 MH683555 MH685727 MH685735 Jiang e al. (2018)
Co yneum su onii CFCC 52318 Cas anea mollissima MH683564 MH683556 MH685728 MH685736 Jiang e al. (2018)
Co yneum su onii Z17 Cas anea mollissima MK799955 MK799942 MK799818 MK799828 Jiang e al. (2019)
Co yneum su onii Z86 Cas anea mollissima MK799956 MK799943 MK799819 MK799829 Jiang e al. (2019)
Co yneum umbona um D201 Que cus obu MH674329 MH674329 MH674333 MH674337 Jiang e al. (2018)
Co yneum agi BJFC-S1782* Fagus sp. MW144761 MW144953 NA NA Boonmee e al. (2021)
Co yneum agi BJFC-S1783 Fagus sp. MW144762 MW144954 NA NA Boonmee e al. (2021)
Co yneum sep emsep a um GMB0393 decaying wood OQ540748 OQ540743 NA NA Long e al. (2023)
Co yneum sep emsep a um GMB0392* decaying wood OQ560328 OQ560329 NA NA Long e al. (2023)
Hyalo e minalis alishanensis NCYUCC 19-0400* Ce asus sp. MT447559 MT447557 NA NA Ra hnayaka e al. (2020)
No e. “NA” indica es una ailable sequences, sequences p oduced in he cu en s udy a e in bold and * means ex- ype s ains.
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Caixia Wang e al.: Diapo hales om Be ula in Xinjiang, China
Table 2. GenBank accession numbe s used in he phylogene ic analyses o Cy ospo a.
Species S ains GenBank accession numbe s Re e ences
ITS ac pb2 e 1 ub2
Cy ospo a ailan hicola CFCC 54064 PP988711 NA PQ074892 PQ074251 PQ075209 Lin e al. (2024)
Cy ospo a ailan hicola CFCC 89970* MH933618 MH933526 MH933592 MH933494 MH933565 Fan e al. (2020)
Cy ospo a al ayensis sp. no . CFCC 71686 PX227527 PX233570 PX233584 PX233598 PX233612 In his s udy
Cy ospo a al ayensis sp. no . CFCC 71687 PX227528 PX233571 PX233585 PX233599 PX233613 In his s udy
Cy ospo a al ayensis sp. no . CFCC 71688 PX227529 PX233572 PX233586 PX233600 PX233614 In his s udy
Cy ospo a a oci ha a CFCC 59056 PP988739 PQ074599 PQ074912 PQ074274 PQ075229 Lin e al. (2024)
Cy ospo a a oci ha a CFCC 89615 KR045618 KF498673 KU710946 KP310858 KR045659 Fan e al. (2015)
Cy ospo a aue swaldii CBS 153.29 PP988740 PQ074600 PQ074913 PQ074275 PQ075230 Lin e al. (2024)
Cy ospo a be be idis CFCC 89927* KR045620 KU710990 KU710948 KU710913 KR045661 Liu e al. (2015)
Cy ospo a be be idis CFCC 89933 KR045621 KU710991 KU710949 KU710914 KR045662 Liu e al. (2015)
Cy ospo a be ulae CBS 141622* PP988752 PQ074610 PQ074922 PQ074284 PQ075236 Lin e al. (2024)
Cy ospo a cel idicola CFCC 50497* MH933623 NA MH933595 MH933499 MH933566 Fan e al. (2020)
Cy ospo a ch ysospe ma CBS 120.83 PP988773 PQ074626 PQ074942 PQ074304 PQ075255 Lin e al. (2024)
Cy ospo a ch ysospe ma CBS 197.50* PP988777 PQ074630 PQ074946 PQ074308 PQ075259 Lin e al. (2024)
Cy ospo a eas ingensis CFCC 58222* PP988818 NA PQ074980 PQ074346 NA Lin e al. (2024)
Cy ospo a guyuanensis CFCC 55855* PP988853 NA PQ075011 PQ074378 PQ075323 Lin e al. (2024)
Cy ospo a hejingensis CFCC 59571* PP060455 PP059657 PP059663 PP059667 PP059673 Wang e al. (2024)
Cy ospo a hippophaopsis CGMCC 3.18997* PP965505 PP957863 NA PP957877 PP957884 Cai e al. (2024)
Cy ospo a i anica IRAN 4200C* MW295652 MZ014512 MW824359 MW394146 NA Hani eh e al. (2022)
Cy ospo a jiu engensis CFCC 55839* PP988865 PQ074698 NA NA PQ075332 Lin e al. (2024)
Cy ospo a joaquinensis CBS 144235 MG971895 MG972044 NA MG971605 NA Law ence e al. (2018)
Cy ospo a kuanchengensis CFCC 52464* MK432616 MK442940 MK578076 NA NA Jiang e al. (2020b)
Cy ospo a leucospe ma CBS 109491 PP988884 PQ074714 PQ075035 PQ074407 PQ075347 Lin e al. (2024)
Cy ospo a longispo a CBS 144236* MG971905 MG972054 NA MG971615 NA Law ence e al. (2018)
Cy ospo a longis iola a MFLUCC 16-0628 KY417734 KY417700 KY417802 NA NA No phanphoun e al. (2017)
Cy ospo a mac opycnidia CBS 149338* OP038094 OP003977 OP095265 OP106954 OP079909 T a adon e al. (2022)
Cy ospo a melnikii CFCC 89984 MH933644 MH933551 MH933609 MH933515 MH933580 Pan e al. (2018)
Cy ospo a nobilis CFCC 58227 PP988928 PQ074756 PQ075075 PQ074449 PQ075389 Lin e al. (2024)
Cy ospo a nobilis CFCC 58228 PP988929 PQ074757 PQ075076 PQ074450 PQ075390 Lin e al. (2024)
Cy ospo a oleicola CBS 144248* MG971944 MG972098 NA MG971660 NA Law ence e al. (2018)
Cy ospo a pla ycladicola CFCC 50038* KT222840 MH933555 MH933613 MH933519 MH933584 Yang e al. (2015)
Cy ospo a populinopsis CFCC 50032* MH933648 MH933556 MH933614 MH933520 MH933585 Fan e al. (2020)
Cy ospo a p uinopsis CFCC 50034* KP281259 KP310836 KU710970 KP310849 KP310819 Yang e al. (2015)
Cy ospo a p uinopsis CFCC 50035 KP281260 KP310837 KU710971 KP310850 KP310820 Yang e al. (2015)
Cy ospo a p uinosa CBS 200.42 PP988951 PQ074777 PQ075095 PQ074470 PQ075410 Lin e al. (2024)
Cy ospo a p unicola MFLU 17-0995* MG742350 MG742353 MG742352 NA NA Hyde e al. (2018)
Cy ospo a pseudoch ysospe ma CFCC 54081* MZ702631 NA NA OK303613 OK303680 Lin e al. (2024)
Cy ospo a pseudoch ysospe ma CFCC 89629 KF765673 NA KF765705 NA NA Lin e al. (2024)
Cy ospo a pseudoch ysospe ma CFCC 89981* MH933625 MH933533 MH933597 MH933501 MH933568 Lin e al. (2024)
Cy ospo a qinghaiensis CFCC 50026* KP281267 KP310843 KU710972 KP310856 KP310826 Yang e al. (2015)
Cy ospo a ibis CBS 187.36 PP988963 PQ074788 PQ075106 PQ074480 PQ075420 Lin e al. (2024)
Cy ospo a osigena MFLUCC 18-0921* MN879872 NA NA NA NA Hyde e al. (2020a)
Cy ospo a os a a CFCC 89909* KR045643 KU711009 KU710974 NA NA Fan e al. (2020)
Cy ospo a os a a CFCC 89910 KR045644 KU711010 KU710975 KU710933 NA Fan e al. (2020)
Cy ospo a salicacea um MFLUCC 15-0509* KY417746 KY417712 KY417814 NA NA No phanphoun e al. (2017)
Cy ospo a salicina CBS 507.77 PP988981 PQ074804 PQ075122 PQ074497 PQ075435 Lin e al. (2024)
Cy ospo a salicina MFLUCC 15-0862* KY417750 KY417716 KY417818 NA NA No phanphoun e al. (2017)
Cy ospo a shaanxiensis CFCC 56032* PP988987 PQ074810 PQ075128 PQ074502 PQ075441 Lin e al. (2024)
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Caixia Wang e al.: Diapo hales om Be ula in Xinjiang, China
Species S ains GenBank accession numbe s Re e ences
ITS ac pb2 e 1 ub2
Cy ospo a sibi aeae CFCC 50045* KR045651 KU711015 KU710982 KU710938 KR045692 Liu e al. (2015)
Cy ospo a sibi aeae CFCC 50046 KR045652 KU711015 KU710983 KU710939 KR045693 Liu e al. (2015)
Cy ospo a sidaohensis CFCC 56042* PP988992 PQ074815 PQ075133 PQ074507 PQ075446 Lin e al. (2024)
Cy ospo a sinensis CFCC 58231 PP988995 PQ074818 PQ075136 PQ074510 PQ075449 Lin e al. (2024)
Cy ospo a sinensis CFCC 58235* PP988997 PQ074820 PQ075138 PQ074512 PQ075451 Lin e al. (2024)
Cy ospo a sinensis CFCC 58471 PP989002 PQ074824 PQ075143 PQ074517 PQ075455 Lin e al. (2024)
Cy ospo a songshanensis CFCC 56351* PP989006 PQ074828 PQ075147 PQ074521 PQ075459 Lin e al. (2024)
Cy ospo a sopho iopsis CFCC 58464 PP989009 PQ074831 PQ075150 PQ074524 PQ075461 Lin e al. (2024)
Cy ospo a sopho iopsis CFCC 89600* KR045623 KU710992 KU710951 KU710915 KP310817 Fan e al. (2020)
Cy ospo a sopho iopsis CFCC 71679 PX227530 PX233573 PX233587 PX233601 PX233615 In his s udy
Cy ospo a sopho iopsis CFCC 71680 PX227531 PX233574 PX233588 PX233602 PX233616 In his s udy
Cy ospo a sopho iopsis CFCC 71681 PX227532 PX233575 PX233589 PX233603 PX233617 In his s udy
Cy ospo a sopho iopsis CFCC 71682 PX227533 PX233576 PX233590 PX233604 PX233618 In his s udy
Cy ospo a sy ingina CFCC 50036* KP310800 KP310832 NA KP310845 KP310815 Lin e al. (2024)
Cy ospo a anai ica MFLUCC 14-1057* KT459411 KT459413 NA NA NA A iyawansa e al. (2015)
Cy ospo a anai icaCFCC 71675 PX227534 PX233577 PX233591 PX233605 PX233637 In his s udy
Cy ospo a anai ica CFCC 71676 PX227535 PX233578 PX233592 PX233606 PX233638 In his s udy
Cy ospo a anai ica CFCC 71677 PX227536 PX233579 PX233593 PX233607 PX233639 In his s udy
Cy ospo a anai ica CFCC 71678 PX227537 PX233580 PX233594 PX233608 PX233640 In his s udy
Cy ospo a anai icaCFCC 71683 PX227538 PX233581 PX233595 PX233609 PX233641 In his s udy
Cy ospo a anai ica CFCC 71684 PX227539 PX233582 PX233596 PX233610 PX233642 In his s udy
Cy ospo a anai ica CFCC 71685 PX227540 PX233583 PX233597 PX233611 PX233643 In his s udy
Cy ospo a eneb ica CFCC 55841* PP989019 PQ074838 PQ075160 PQ074534 PQ075471 Lin e al. (2024)
Cy ospo a e aspo a CFCC 55847* PP989021 PQ074840 PQ075162 PQ074536 PQ075473 Lin e al. (2024)
Cy ospo a e aspo a CFCC 56279* PP989023 PQ074842 PQ075164 PQ074538 PQ075475 Lin e al. (2024)
Cy ospo a i ici CBS 118561 PP989038 PQ074856 PQ075175 PQ074549 PQ075486 Lin e al. (2024)
Cy ospo a i ici CBS 118563 PP989039 PQ074857 PQ075176 PQ074550 PQ075487 Lin e al. (2024)
Cy ospo a ulmi MFLUCC 15-0863* KY417759 KY417725 KY417827 NA NA No phanphoun e al. (2017)
Cy ospo a ulmicola MFLUCC 18-1227* MH940220 MH940216 NA NA NA Phookamsak e al. (2019)
Cy ospo a unilocula a CFCC 58460* PP989043 PQ074861 NA PQ074554 NA Lin e al. (2024)
Cy ospo a washing onensis CBS 141619* PP989065 PQ074874 PQ075192 PQ074569 PQ075502 Lin e al. (2024)
Cy ospo a xiaolongmenensis CFCC 58459* PP989068 PQ074877 PQ075194 PQ074572 PQ075505 Lin e al. (2024)
Cy ospo a yakimana CBS 149297* OM976602 ON012555 ON045093 ON012569 ON086750 T a adon e al. (2022)
Cy ospo a yuduensis CFCC 57539* PP989070 PQ074879 PQ075196 PQ074574 PQ075507 Lin e al. (2024)
Diapo he accinii CBS 160.32 KC343228 JQ807297 NA KC343954 KC344196 Gomes e al. (2013)
No e. “NA” indica es una ailable sequences, sequences p oduced in he cu en s udy a e in bold and * means ex- ype s ains.
The conca ena ed da ase o ITS, ac , pb2, e 1 and ub2 o Cy ospo a
ibis species complex, which con ained CFCC 71675, CFCC 71676, CFCC
71677, CFCC 71678, CFCC 71683, CFCC 71684 and CFCC 71685, comp ised
30 s ains, spanning 2,818 aligned cha ac e s (ac : 1–237; ITS: 238–742;
pb2: 743–1,940; e 1: 1,941–2,415; ub2: 2,416–2,818), including gaps. Max-
imum Likelihood (ML) analysis o his da ase p oduced a bes -sco ing ee
wi h a likelihood alue o −10563.20. The alignmen ma ix con ained 711
dis inc pa e ns, wi h 25.23% o si es ep esen ing unde e mined cha ac e s
o gaps. Nucleo ide equencies we e es ima ed as ollows: A = 0.248, C =
0.282, G = 0.236, T = 0.232. Subs i u ion a es we e AC = 1.417, AG = 3.402,
AT = 1.463, CG = 0.735, CT = 6.621 and GT = 1.0, wi h a gamma dis ibu ion
shape pa ame e (α) o 0.245. Fo Bayesian In e ence (BI), he op imal e o-
lu iona y models, selec ed by M ModelTes , we e TIM2e+G4 o ITS, K2P+G4
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Table 3. GenBank accession numbe s used in he phylogene ic analyses o C yp ospo ella.
Species S ains Hos s GenBank accession numbe s Re e ences
ITS e 1 ub2
C yp ospo ella alnicola CBS 121074 Co ylus co nu a EU199204 EU221960 EU219138 Mejía e al. (2008)
C yp ospo ella alni- ub ae CBS 126120 Alnus ub a GU826092 GU826051 GU826010 Mejía e al. (2011)
C yp ospo ella alni- ub ae LCM411 Alnus ub a GU826090 GU826049 GU826008 Mejía e al. (2011)
C yp ospo ella alni- ub ae LCM499.01* Alnus ub a GU826096 GU826055 GU826014 Mejía e al. (2011)
C yp ospo ella alni-sinua ae AR4200 Alnus i idis ssp. sinua a GU826086 GU826045 GU825989 Mejía e al. (2011)
C yp ospo ella alni-sinua ae CBS 125662* Alnus i idis ssp. sinua a GU826087 GU826046 GU826005 Mejía e al. (2011)
C yp ospo ella alni- enui oliae CBS 125663* Alnus incana ssp. enui ol GU826097 GU826056 GU826015 Mejía e al. (2011)
C yp ospo ella alni-co da ae MFLUCC 16-0485 Alnus sp.KY797640 NA NA Tian e al. (2018)
C yp ospo ella alni-co da ae MFLU 16-0812 Alnus sp.KY797639 NA NA Tian e al. (2018)
C yp ospo ella amis adensis LCM618.01 Alnus acumina a GU826109 GU826073 GU826032 Mejía e al. (2011)
C yp ospo ella amis adensis CBS 125664* Alnus acumina a GU826108 GU826072 GU826031 Mejía e al. (2011)
C yp ospo ella be ulae CBS 121078 Be ula pendula EU199213 GU826057 GU826016 Mejía e al. (2008)
C yp ospo ella be ulae LCM477.01 Be ula pendula GU826098 GU826059 GU826018 Mejía e al. (2011)
C yp ospo ella be ulae CBS 121079 Be ula pendula EU199216 GU826058 GU826017 Mejía e al. (2008)
C yp ospo ella be ulae CFCC 71654 Be ula pendula PX227541 PX233644 PX233656 In his s udy
C yp ospo ella be ulae CFCC 71655 Be ula pendula PX227542 PX233645 PX233657 In his s udy
C yp ospo ella be ulae CFCC 71656 Be ula pendula PX227543 PX233646 PX233658 In his s udy
C yp ospo ella be ulae CFCC 71657 Be ula pendula PX227544 PX233647 PX233659 In his s udy
C yp ospo ella be ulae CFCC 71658 Be ula pendula PX227545 PX233648 PX233660 In his s udy
C yp ospo ella con usa CBS 121003 Be ula papy i e a EU199219 NA NA Mejía e al. (2008)
C yp ospo ella co ylina LCM391.04 Co ylus a ellana GU826100 GU826063 GU826022 Mejía e al. (2011)
C yp ospo ella emo alis CBS 121076* Alnus incana ssp. ugosa EU199220 EU221951 EU219139 Mejía e al. (2008)
C yp ospo ella emo alis LCM196.04 Alnus incana ssp. ugosa GU826102 GU826067 GU826025 Mejía e al. (2011)
C yp ospo ella hypode mia CBS 109753 Ulmus mino EU199224 GU826064 GU826023 Mejía e al. (2008)
C yp ospo ella hypode mia CBS 122593* Ulmus mino EU199181 GU826066 GU826024 Mejía e al. (2008)
C yp ospo ella jakli schii LCM112.01 Alnus se ula a GU826089 GU826048 GU826007 Mejía e al. (2011)
C yp ospo ella jakli schii CBS 125665* Alnus se ula a GU826088 GU826047 GU826006 Mejía e al. (2011)
C yp ospo ella ma ylandica LCM386.05 Alnus ma i ima GU826106 GU826070 GU826029 Mejía e al. (2011)
C yp ospo ella ma ylandica LCM581.01 Alnus ma i ima GU826107 GU826071 GU826030 Mejía e al. (2011)
C yp ospo ella ma ylandica CBS 125666* Alnus ma i ima GU826105 GU826069 GU826028 Mejía e al. (2011)
C yp ospo ella mul icon inen alis CBS 126119 Alnus incana ssp. ugosa GU826081 GU826040 GU825999 Mejía e al. (2011)
C yp ospo ella mul icon inen alis LCM93b.02 Alnus incana ssp. ugosa GU826082 GU826041 GU826000 Mejía e al. (2011)
C yp ospo ella mul icon inen alis LCM427.01 Alnus glu inosa GU826085 GU826044 GU826004 Mejía e al. (2011)
C yp ospo ella mul icon inen alis LCM401.01* Alnus glu inosa GU826083 GU826042 GU826001 Mejía e al. (2011)
C yp ospo ella paci ica LCM453.01 Alnus incana ssp. enui olia GU826077 GU826037 GU825995 Mejía e al. (2011)
C yp ospo ella paci ica LCM461.01* Alnus incana ssp. enui olia GU826076 GU826036 GU825994 Mejía e al. (2011)
C yp ospo ella pla yphylla CFCC 50466 Be ula pla yphylla KT732947 KT733015 KT733019 Fan e al. (2016b)
C yp ospo ella pla yphylla CFCC 50465* Be ula pla yphylla KT732946 KT733014 KT733018 Fan e al. (2016b)
C yp ospo ella su usa LCM576.01 Alnus sp.GU826079 GU826039 GU825997 Mejía e al. (2011)
C yp ospo ella su usa LCM576.03 Alnus sp.GU826078 GU826038 GU825996 Mejía e al. (2011)
C yp ospo ella omen ella CBS 126440 Be ula alleghaniensis GU826099 GU826062 GU826021 Mejía e al. (2011)
C yp ospo ella omen ella CBS 121073 Be ula sp.EU199217 GU826060 GU826019 Mejía e al. (2008)
C yp ospo ella omen ella CFCC 71659 Be ula mic ophylla PX227546 PX233649 PX233661 In his s udy
C yp ospo ella omen ella CFCC 71660 Be ula mic ophylla PX227547 PX233650 PX233662 In his s udy
C yp ospo ella omen ella CFCC 71661 Be ula mic ophylla PX227548 PX233651 PX233663 In his s udy
C yp ospo ella omen ella CFCC 71662 Be ula mic ophylla PX227549 PX233652 PX233664 In his s udy
C yp ospo ella omen ella CFCC 71663 Be ula mic ophylla PX227550 PX233653 PX233665 In his s udy
C yp ospo ella omen ella CFCC 71664 Be ula mic ophylla PX227551 PX233654 PX233666 In his s udy
C yp ospo ella omen ella CFCC 71665 Be ula mic ophylla PX227552 PX233655 PX233667 In his s udy
C yp ospo ella wehmeye iana LCM85.02 Tilia ame icana GU826104 GU826068 GU826027 Mejía e al. (2011)
Di opella di opa LCM94.02 Alnus incana ssp. ugosa GU826075 GU826033 GU825990 Mejía e al. (2011)
No e. “NA” indica es una ailable sequences, sequences p oduced in he cu en s udy a e in bold and * means ex- ype s ains.
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Table 4. GenBank accession numbe s used in he phylogene ic analyses o Melanconis.
Taxa S ains Hos s GenBank accession numbe s Re e ences
ITS LSU pb2 e 1 ub2
Juglanconis juglandina CBS 133343 Juglans egia KY427149 KY427149 KY427199 KY427218 KY427234 Voglmay e al. (2017)
Juglanconis p e oca yae CBS 144326* P e oca ya axini olia MK229175 MK229175 MK238324 MK238332 MK238338 Voglmay e al. (2017)
Melanconis alni CBS 131693 Alnus glu inosa MN784962 MN784962 MN780745 MN780774 MN780803 Jakli sch and
Voglmay (2020)
Melanconis alni CBS 131695* Alnus glu inosa MN784963 MN784963 MN780746 MN780775 MN780804 Jakli sch and
Voglmay (2020)
Melanconis alni MEW* Alnus glu inosa MN784964 MN784964 MN780747 MN780776 MN780805 Jakli sch and
Voglmay (2020)
Melanconis alni MAIV Alnus incana MN784965 MN784965 MN780748 MN780777 MN780806 Jakli sch and
Voglmay (2020)
Melanconis alni D156 Alnus glu inosa MN784966 MN784966 MN780749 MN780778 MN780807 Jakli sch and
Voglmay (2020)
Melanconis be ulae CFCC 50471* Be ula albosinensis KT732952 KT732971 KT732984 KT733001 KT733022 Fan e al. (2016a)
Melanconis be ulae CFCC 50472 Be ula albosinensis KT732953 KT732972 KT732985 KT733002 KT733023 Fan e al. (2016a)
Melanconis be ulae CFCC 50473 Be ula albosinensis KT732954 KT732973 KT732986 KT733003 KT733024 Fan e al. (2016a)
Melanconis g oenlandica CBS 116450* Be ula nana KU878552 KU878553 NA KU878554 KU878555 Lomba d e al. (2016)
Melanconis g oenlandica MAFF
410219
Be ula maximowic-
ziana MN784967 MN784967 MN780750 MN780779 MN780808 Jakli sch and
Voglmay (2020)
Melanconis g oenlandica CBS 133341 Be ula papy i e a MN784968 MN784968 MN780751 MN780780 MN780809 Jakli sch and
Voglmay (2020)
Melanconis g oenlandica CBS 133339 Be ula sp. MN784969 MN784969 MN780752 MN780781 MN780810 Jakli sch and
Voglmay (2020)
Melanconis g oenlandica CBS 133340 Be ula papy i e a MN784970 MN784970 MN780753 MN780782 MN780811 Jakli sch and
Voglmay (2020)
Melanconis g oenlandica CFCC 71566 Be ula ianchanica PX227553 PX227509 NA PX233619 PX233628 In his s udy
Melanconis g oenlandica CFCC 71567 Be ula ianchanica PX227554 PX227510 NA PX233620 PX233629 In his s udy
Melanconis g oenlandica CFCC 71568 Be ula ianchanica PX227555 PX227511 NA PX233621 PX233630 In his s udy
Melanconis g oenlandica CFCC 71569 Be ula ianchanica PX227556 PX227512 NA PX233622 PX233631 In his s udy
Melanconis i oana MAFF
410080 Be ula e manii MN784971 MN784971 MN780754 MN780783 MN780812 Jakli sch and
Voglmay (2020)
Melanconis i oana CFCC 50474 Be ula albosinensis KT732955 KT732974 KT732987 KT733004 KT733025 Fan e al. (2016a)
Melanconis i oana CFCC 52876 Be ula albosinensis MK096324 MK096364 MK096409 MK096284 NA Fan e al. (2016a)
Melanconis i oana CFCC 52877 Be ula albosinensis MK096326 MK096366 MK096411 MK096286 NA Fan e al. (2016a)
Melanconis i oana CFCC 52878 Be ula albosinensis MK096327 MK096367 MK096412 MK096287 NA Fan e al. (2016a)
Melanconis la issae CBS 123196* Be ula sp. MN784972 MN784972 MN780755 MN780784 MN780813 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. eu opaea D157 Alnus alnobe ula MN784973 MN784973 MN780756 MN780785 NA Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. eu opaea D158 Alnus alnobe ula MN784974 MN784974 MN780757 MN780786 MN780814 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. eu opaea D257 Alnus incana MN784975 MN784975 MN780758 MN780787 MN780815 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. eu opaea CBS 131692* Alnus incana MN784976 MN784976 MN780759 MN780788 MN780816 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. eu opaea CBS 131694 Alnus alnobe ula MN784977 MN784977 MN780760 MN780789 MN780817 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. eu opaea MAV1 Alnus alnobe ula MN784978 MN784978 MN780761 MN780790 MN780818 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. i alica
MFLUCC 16-
1199* Alnus co da a MF190151 MF190096 NA NA NA Senanayake e al.
(2017)
Melanconis ma ginalis
subsp. i alica
MFLUCC 17-
1659* Alnus co da a MF190152 MF190097 MF377602 NA NA Senanayake e al.
(2017)
Melanconis ma ginalis
subsp. ma ginalis D321* Alnus alnobe ula
subsp. c ispa MN784979 MN784979 MN780762 MN780791 MN780819 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. ma ginalis D321a* Alnus alnobe ula
subsp. c ispa MN784980 MN784980 MN780763 MN780792 MN780820 Jakli sch and
Voglmay (2020)
175
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Taxa S ains Hos s GenBank accession numbe s Re e ences
ITS LSU pb2 e 1 ub2
Melanconis ma ginalis
subsp. ma ginalis D321b* Alnus alnobe ula
subsp. c ispa MN784981 MN784981 MN780764 MN780793 MN780821 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. ma ginalis CBS 109496 Alnus alnobe ula sub-
sp. maximowiczii MN784982 MN784982 MN780765 MN780794 MN780822 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. ma ginalis AR 4864 Alnus alnobe ula MN784983 MN784983 MN780766 MN780795 MN780823 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. ma ginalis CBS 133346 Alnus alnobe ula MN784984 MN784984 MN780767 MN780796 MN780824 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. ma ginalis
MAFF
410218
Alnus alnobe ula sub-
sp. maximowiczii MN784985 MN784985 MN780768 MN780797 MN780825 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. i olensis CBS 122310* Alnus alnobe ula MN784986 MN784986 MN780769 MN780798 MN780826 Jakli sch and
Voglmay (2020)
Melanconis ma ginalis
subsp. i olensis D322a Alnus alnobe ula MN959458 MN959458 MN989415 MN989416 MN989417 Jakli sch and
Voglmay (2020)
Melanconis paci ica CBS 109744 Alnus ub a EU199197 AF408373 DQ862022 DQ862038 EU219103 Mejía e al. (2008)
Melanconis s ilbos oma D143 Be ula pendula KY427156 KY427156 KY427206 KY427225 KY427241 Voglmay e al. (2017)
Melanconis s ilbos oma D258 Be ula ae nensis MN784987 MN784987 MN780770 MN780799 MN780827 Jakli sch and
Voglmay (2020)
Melanconis s ilbos oma CBS 109778* Be ula pendula MN784988 MN784988 MN780771 MN780800 MN780828 Jakli sch and
Voglmay (2020)
Melanconis s ilbos oma CBS 121894 Be ula pendula KY427156 KY427156 JQ926302 JQ926368 MN780830 Voglmay e al. (2017)
Melanconis s ilbos oma CBS 133338 Be ula papy i e a MN784990 MN784990 MN780773 MN780802 MN780831 Jakli sch and
Voglmay (2020)
Melanconis s ilbos oma CFCC 50475 Be ula pla yphylla KT732956 KT732975 KT732988 KT733005 KT733026 Fan e al. (2016a)
Melanconis s ilbos oma CFCC 50476 Be ula pla yphylla KT732957 KT732976 KT732989 KT733006 KT733027 Fan e al. (2016a)
Melanconis s ilbos oma CFCC 50477 Be ula pla yphylla KT732958 KT732977 KT732990 KT733007 KT733028 Fan e al. (2016a)
Melanconis s ilbos oma CFCC 50478 Be ula pla yphylla KT732959 KT732978 KT732991 KT733008 KT733029 Fan e al. (2016a)
Melanconis s ilbos oma CFCC 50479 Be ula pla yphylla KT732960 KT732979 KT732992 KT733009 KT733030 Fan e al. (2016a)
Melanconis s ilbos oma CFCC 50480 Be ula pla yphylla KT732961 KT732980 KT732993 KT733010 KT733031 Fan e al. (2016a)
Melanconis s ilbos oma CFCC 50481 Be ula pla yphylla KT732962 KT732981 KT732994 KT733011 KT733032 Fan e al. (2016a)
Melanconis s ilbos oma CFCC 50482 Be ula pla yphylla KT732963 KT732982 KT732995 KT733012 KT733033 Fan e al. (2016a)
Melanconis s ilbos oma CFCC 50483 Be ula pla yphylla KT732964 KT732983 KT732996 KT733013 KT733034 Fan e al. (2016a)
Melanconis s ilbos oma CFCC 71570 Be ula pendula PX227557 PX227513 NA PX233623 PX233632 In his s udy
Melanconis s ilbos oma CFCC 71571 Be ula pendula PX227558 PX227514 NA PX233624 PX233633 In his s udy
Melanconis s ilbos oma CFCC 71572 Be ula pendula PX227559 PX227515 NA PX233625 PX233634 In his s udy
Melanconis s ilbos oma CFCC 71573 Be ula pendula PX227560 PX227516 NA PX233626 PX233635 In his s udy
Melanconis s ilbos oma CFCC 71574 Be ula pendula PX227561 PX227517 NA PX233627 PX233636 In his s udy
No e. “NA” indica es una ailable sequences, sequences p oduced in he cu en s udy a e in bold and * means ex- ype s ains.
o ac , TNe+R2 o pb2, TIM2e+G4 o e 1 and HKY+F+G4 o ub2. BI e-
sul s we e cong uen wi h ML opology. Phylogene ic econs uc ion placed
he se en isola es om his s udy (CFCC 71675, CFCC 71676, CFCC 71677,
CFCC 71678, CFCC 71683, CFCC 71684 and CFCC 71685) in o a well-sup-
po ed clade wi h MFLUCC 14-1057, iden i ied as Cy ospo a anai ica (Fig. 2).
The conca ena ed da ase o ITS, ac , pb2, e 1 and ub2 o Cy ospo a
ch ysospe ma species complex, which con ained CFCC 71679, CFCC 71680,
CFCC 71681 and CFCC 71682, comp ised 33 s ains, spanning 3,182 aligned
cha ac e s (ac : 1–254; ITS: 255–754; pb2: 755–1,952; e 1: 1,953–2,441; ub2:
2,442–3,182), including gaps. Maximum Likelihood (ML) analysis o his da a-
se p oduced a bes -sco ing ee wi h a likelihood alue o -9050.97. The align-
men ma ix con ained 608 dis inc pa e ns, wi h 32.72% o si es ep esen ing
unde e mined cha ac e s o gaps. Nucleo ide equencies we e es ima ed as
ollows: A = 0.248, C = 0.291, G = 0.236, T = 0.223. Subs i u ion a es we e AC =
182
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Caixia Wang e al.: Diapo hales om Be ula in Xinjiang, China
Figu e 6. Maximum Likelihood ee o Melanconis gene a ed om combined ITS, LSU, pb2, e 1 and ub2 sequence
da a. Boo s ap suppo alues ≥ 50% and Bayesian pos e io p obabili ies ≥ 0.90 a e demons a ed a he b anches.
Ex- ype s ains a e ma ked in bold.
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Taxonomy
Co yneaceae Co da, Icon. ung. (P ague) 3: 36 (1839)
No es. Co yneaceae, ypi ied by Co yneum, ep esen s a bo h mo pholog-
ically and phylogene ically dis inc lineage wi hin Diapo hales, p ima ily
cha ac e ised by he p oduc ion o ans e sely dis osep a e, b own conidia
(Senanayake e al. 2017, 2018; Jiang e al. 2018, 2019; Ra hnayaka e al.
2020). To da e, h ee gene a ha e been accep ed in Co yneaceae, based
on in eg a ed mo phological and phylogene ic e idence, iz. Co yneum,
Hyalo e minalis and Subellipsoidispo a (Tang e al. 2023). While phyloge-
ne ic analyses ha e placed Talekpea in close ela ionship wi h Co yneum
(Ra hnayaka e al. 2020; Tang e al. 2023), his genus exhibi s signi ican
mo phological di e gence om he h ee accep ed gene a. Resolu ion o i s
axonomic s a us will equi e addi ional sampling and mo e comp ehensi e
s udies in he u u e. No ably, Hyalo e minalis is dis inguished om Co y-
neum, based on i s pycnidial conidioma a (Ra hnayaka e al. 2020). Subel-
lipsoidispo a has been desc ibed only om i s sexual mo ph, dis inguished
om Co yneum by ascoma a ypically bea ing a single pe idium (Tang e al.
2023). This dispa i y in known mo phologies highligh s he need o u he
esea ch o ully unde s and he li e his o ies and e olu iona y ela ionships
wi hin Co yneaceae.
Co yneum Nees, Sys . Pilze (Wü zbu g): 34 (1816)
= Pseudo alsa Ces. & De No ., Comm. Soc. c i og. I al. 1( asc. 4): 206 (1863).
No es. Co yneum, ypi ied by C. umbona um, is he la ges genus wi hin he
amily Co yneaceae. The asexual mo phs o Co yneum a e commonly ob-
se ed and a e p edominan ly ound on ee gene a such as Be ula, Cas a-
nea and Que cus (Senanayake e al. 2017; Jiang e al. 2018, 2019). Species
in his genus a e cha ac e ised by ans e sely dis osep a e, b own conidia
(Senanayake e al. 2017). Cu en ly, he delimi a ion o species wi hin Co y-
neum elies on a combina ion o mo phological cha ac e is ics and molecu-
la phylogene ic analyses, based on ITS, LSU, pb2 and e 1 loci (Jiang e al.
2018, 2019). Ne e heless, he lack o molecula da a o many desc ibed
species emains a cons ain o accu a e species iden i ica ion (Jiang e al.
2018, 2019). Species o Co yneum a e commonly epo ed as plan pa ho-
gen causing canke diseases and some species a e sap obic on decaying
wood con ibu ing o nu ien cycling (Senanayake e al. 2017, 2018).
Co yneum lanci o me (F .) Voglmay & Jakli sch, IMA Fungus 6(1): 146 (2015)
Fig. 7
= Co yneum b achyu um Link, Sp. pl., Edn 4 6(2): 124 (1825).
≡ Pseudo alsa lanci o mis (F .) Ces. & De No ., Comm. Soc. c i og. I al. 1( asc.
4): 206 (1863).
184
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Desc ip ion. Pseudos oma a semi-imme sed in he ba k, sca e ed, conical,
950–1700 μm diam., 600–900 μm high, wi h 3–8 pe i hecia a anged i egu-
la ly. Ec os oma ic disc dis inc , ci cula , b own, 430–650 μm diam. Os ioles
black, 75–140 μm diam. Pe i hecia globula , somewha la ened a base wi h
black neck, 350–650 μm diam. Asci hyaline, wi h chi inoid, e ac i e ing,
cla a e o elonga e-obo oid, (190–)207–241(–246) × (30–)33–40(–42) μm,
Figu e 7. Co yneum lanci o me om Be ula pendula. A, B. Habi o ascos oma a on b anch; C. Os ioles embedded in
ascos oma in sec ion; D. T ans e se sec ion h ough ascos oma; E. Longi udinal sec ion h ough ascos oma; F. Asci;
G. Ascospo es. Scale ba s: 500 μm (B–E); 20 μm (F, G).
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8-spo ed, bise ia e. Ascospo es usi o m, ends poin ed, da k b own, 5–8-dis-
osep a e, (37.5–)39.5–48.5(–53) × (13.5–)14–17(–18.5) (a . = 44.1 ± 4.4 ×
15.4 ± 1.6, n = 50) (n = 50) μm, L/W a io = 2.4–3.6.
Cul u e cha ac e is ics. Colonies on PDA la , sp eading, wi h mode a e loc-
culen ae ial mycelium and undula ing ma gin, ini ially whi e, becoming da k
b own and eaching 90 mm diam. a e 3 weeks a 25 °C.
Ma e ials examined. China • Xinjiang Uygu Au onomous Region, Al ay
P e ec u e, Al ay Ci y, Alahake Town, Haxionggou, om dead b anches o
Be ula pendula, 7 Oc obe 2024, Rong Ma, Caixia Wang & Hailong Lu (XJAU
4053, li ing cul u es CFCC 71587 and CFCC 71653).
No es. Co yneum lanci o me is he ype species o Pseudo alsa (Su on
1975). Howe e , since Co yneum umbona um ( he ype species o Co yneum)
was desc ibed ea lie (Nees on Esenbeck 1816), Pseudo alsa was synony-
mised unde Co yneum and P. lanci o mis was ea ed as a synonym o Co. lan-
ci o me (Rossman e al. 2015). In his s udy, we epo he i s disco e y o Co.
lanci o me along wi h i s eleomo ph in Xinjiang, China, signi ican ly expanding
he known geog aphical dis ibu ion o his ungus.
Cy ospo aceae F . [as ‘Cy ispo ei’], Sys . o b. eg. (Lundae): 118 (1825)
No es. Cy ospo aceae is a mo phological and phylogene ical amily o Diapo -
hales, con aining a single genus Cy ospo a (Lin e al. 2024).
Cy ospo a Eh enb., Syl . mycol. be ol. (Be lin): 28 (1818)
No es. Cy ospo a is cha ac e ised by he single o laby in hine, locula e s o-
ma a, ilamen ous conidiopho es and allan oid hyaline conidia and ascospo es
(Jiang e al. 2020b; Zhu e al. 2020; Cai e al. 2024; Jia e al. 2024; Li e al.
2024; Lin e al. 2024; Jiang e al. 2025a; Ilyukhin e al. 2025). Membe s o his
genus a e commonly known as causal agen s o ee canke diseases, bu also
include endophy ic and sap ophy ic species (Fan e al. 2020).
Cy ospo a al ayensis C.X. Wang, Ning Jiang & R. Ma, sp. no .
MycoBank MB 860537
Fig. 8
E ymology. Named a e he collec ion si e o he holo ype, Al ay P e ec u e.
Desc ip ion. Conidioma a pycnidial, sca e ed, imme sed o semi-imme sed
in he ba k, discoid o conical, 400–600 μm diam., 250–400 μm high, wi h an
undi ided locule. Concep acle absen . Ec os oma ic disc isabelline, ci cula o
o oid, 150–250 μm diam., wi h a single os iole pe disc in he cen e. Os iole
g ey o black, 50–100 μm diam. Conidiopho es bo ne along he locules, hyaline,
unb anched o b anched a he base, (29–)35.5–43.5(–47) × 1.5–2 μm. Conid-
iogenous cells en e oblas ic, phialidic, subcylind ical o cylind ical. Conidia hy-
aline, allan oid, hin-walled, egu ula e, asep a e, smoo h, (8.5–)9.5–11.5(–13)
× 2.5–3(–3.5) (a . = 10.6 ± 1 × 2.8 ± 0.2, n = 50) μm, L/W a io = 3.4–4.2.
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Cul u e cha ac e is ics. Colonies on PDA la , sp eading, wi h mode a e ae i-
al mycelium and undula ing ma gin, sa on o och eous, eaching 90 mm diam.
a e 2 weeks a 25 °C, s e ile.
Ma e ials examined. China • Xinjiang Uygu Au onomous Region, Al ay P e-
ec u e, Habahe Coun y, Bi ch Fo es Scenic A ea, om b anches o Be ula pen-
dula, 4 Oc obe 2024, Rong Ma, Caixia Wang & Hailong Lu (holo ype XJAU 3988,
ex-holo ype cul u es CFCC 71686, CFCC 71687 and CFCC 71688).
No es. Th ee isola es o Cy ospo a om Be ula pendula in his s udy o med
a dis inc subclade phylogene ically close o Cy. be be idis om Be be is
dasys achya, ep esen ing a new species (Fig. 4). Howe e , Cy. al ayensis can
be dis inguished om Cy. be be idis by conidial size (9.5–11.5 × 2.5–3 μm in
Cy. be be idis s. 6–6.9 × 1.9–2.1 μm in Cy. be be idis) (Liu e al. 2015). A
he nucleo ide le el, Cy. al ayensis di e s om Cy. be be idis (ITS, 15/581; ac ,
22/254; pb2, 58/721; e 1, 161/597; ub2, 75/415) (Jeewon and Hyde 2016).
Figu e 8. Mo phology o Cy ospo a al ayensis om Be ula pendula. A, B. Habi o conidioma a on b anch; C, D. T ans e se
sec ion h ough conidioma a; E. Longi udinal sec ion h ough conidioma a; F. Conidiopho es and conidiogenous cells;
G. Conidia. Scale ba s: 300 μm (B–E); 10 μm (F, G).
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Cy ospo a sopho iopsis X.L. Fan & C.M. Tian, Pe soonia 45: 39 (2019).
Desc ip ion. See Fan e al. (2020).
Ma e ials examined. China • Xinjiang Uygu Au onomous Region, Al ay
P e ec u e, Al ay Ci y, Alahake Town, Haxionggou, om b anches o Be ula
pendula, 7 Oc obe 2024, Rong Ma, Caixia Wang & Hailong Lu (XJAU 4048, cul-
u es CFCC 71679, CFCC 71680, CFCC 71681 and CFCC 71682).
No es. Cy ospo a sopho iopsis was in oduced on canke ed b anches o
S yphnolobium japonicum (Fan e al. 2020). This ungus was la e con i med
as popla and willow canke pa hogens in China (Lin e al. 2022, 2023). In
his s udy, we i s ly disco e ed his species om Be ula pendula in XUAR
(Fig. 3).
Cy ospo a anai ica No ph., Bulgako & K.D. Hyde, Fungal Di e si y 75: 172
(2015)
Fig. 9
Desc ip ion. Conidioma a pycnidial, se ied, semi-imme sed in he ba k, coni-
cal, 1200–2000 μm diam., 400–650 μm high, wi h mul iple subdi ided locules
wi h common walls. Concep acle absen . Ec os oma ic disc honey, ci cula o
o oid, 350–650 μm diam., wi h a single os iole pe disc in he cen e. Os iole
g ey o black, 75–140 μm diam. Conidiopho es bo ne along he locules, hyaline,
b anched, (13–)15–28.5(–32) × 1.5–2 μm. Conidiogenous cells en e oblas ic,
phialidic, subcylind ical o cylind ical. Conidia hyaline, allan oid, hin-walled,
egu ula e, asep a e, smoo h, (5–)5.5–6(–7) × 1.5–2 (a . = 5.8 ± 0.4 × 1.7 ± 0.1,
n = 50) μm, L/W a io = 3.2–3.8.
Cul u e cha ac e is ics. Colonies on PDA la , sp eading, wi h abundan ae i-
al mycelium and ough ma gin, g ey o da k g een, eaching 90 mm diam. a e
2 weeks a 25 °C, s e ile.
Ma e ials examined. China • Xinjiang Uygu Au onomous Region, Al ay P e-
ec u e, Al ay Ci y, Alahake Town, Haxionggou, om b anches o Be ula pen-
dula, 7 Oc obe 2024, Rong Ma, Caixia Wang & Hailong Lu (XJAU 4047, cul-
u es CFCC 71675, CFCC 71676); • Xinjiang Uygu Au onomous Region, Al ay
P e ec u e, Al ay Ci y, om b anches o Be ula pendula, 7 Oc obe 2024, Rong
Ma, Caixia Wang & Hailong Lu (XJAU 4064, cul u es CFCC 71677 and CFCC
71678); • Xinjiang Uygu Au onomous Region, Al ay P e ec u e, Qinghe Coun-
y, Xileasheke Village, om b anches o Be ula pendula, 8 Oc obe 2024, Rong
Ma, Caixia Wang & Hailong Lu (XJAU 4084, cul u es CFCC 71683, CFCC 71684,
CFCC 71685).
No es. Cy ospo a anai ica was i s desc ibed om Be ula pubescens a .
glab a a in Russia (A iyawansa e al. 2015). In he p esen s udy, se en isola es
ob ained om Be ula pendula in Xinjiang, China, clus e ed wi hin a single clade
oge he wi h he ex- ype s ain o Cy. anai ica in he phylogene ic ee (Fig. 2).
Howe e , he conidia o ou specimens we e la ge han hose o he holo ype
(5.5–6 × 1.5–2 μm s. 3.5–4 × 0.6–0.7 μm) (A iyawansa e al. 2015). Based on
he molecula phylogene ic e idence and he sha ed hos genus, he isola es
ob ained in his s udy a e iden i ied as Cy. anai ica, ep esen ing a new hos
and geog aphic eco d o his species.
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Gnomoniaceae G. Win e [as ‘Gnomonieae’], Rabenh. K yp .-Fl. 1(2): 570 (1886)
No es. Gnomoniaceae is cha ac e ised by imme sed, a ely e umpen o supe i-
cial ascoma a, wi hou a s oma o agg ega ed wi h a udimen a y s oma (Sena-
nayake e al. 2017). Membe s o Gnomoniaceae a e endophy es, pa hogens and
sap obes inhabi ing a ious hos s and subs a es (Senanayake e al. 2018).
C yp ospo ella Sacc., Michelia 1(no. 1): 30 (1877)
No es. C yp ospo ella is cha ac e ised by agg ega ed ascoma a below he ba k
su ace, wi h con e ging necks and ellipsoid o elonga ed, asep a e o a e-
ly 1-sep a e ascospo es (Mejía e al. 2008, 2011). Species o his genus a e
usually dis ibu ed in empe a e egions as endophy es and occasionally as
Figu e 9. Mo phology o Cy ospo a anai ica om Be ula pendula. A, B. Habi o conidioma a on b anch; C, D. T ans e se
sec ion h ough conidioma a; E. Longi udinal sec ion h ough conidioma a; F. Conidiopho es and conidiogenous cells; G.
Conidia. Scale ba s: 500 μm (B–E); 10 μm (F–G).
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sap obes and pa hogens on ha dwood ees, such as Be ulaceae, Tiliaceae and
Ulmaceae (Ba 1978; Mejía e al. 2011; Fan e al. 2016 b).
C yp ospo ella be ulae (Tul. & C. Tul.) L.C. Mejía & Cas l., Mycol. Res. 112(1):
32 (2008)
Fig. 10
Desc ip ion. Conidioma a ace ula , imme sed o semi-imme sed in he ba k,
sca e ed, conical, 850–1200 μm diam., 650–800 μm high. Cen al column be-
nea h he disc mo e o less conical, g ey o black. Ec os oma ic disc b own,
ci cula o o oid, 350–420 μm diam. Conidiopho es educed o conidiogenous
cells. Conidiogenous cells na owly cylind ical, smoo h, hyaline, p oducing a co-
Figu e 10. Mo phology o C yp ospo ella be ulae om Be ula pendula. A–C. Habi o conidioma a on b anch; D. T ans-
e se sec ion h ough conidioma a; E. Longi udinal sec ion h ough conidioma a; F. Conidiopho es and conidiogenous
cells; G. Conidia. Scale ba s: 500 μm (B–E); 10 μm (F, G).
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nidium a apex, (6–)8–10.5(–13) × (3–)4.5–6.5(–8) μm. Conidia hyaline, asep-
a e, cylind ical o cla a e, cu ed, (52–)55.5–66.5(–70) × (4.5–)5.5–6.5 μm
(a . = 61 ± 5.2 × 5.8 ± 0.6 μm, n = 50) μm, L/W a io = 9.1–11.9.
Cul u e cha ac e is ics. Colonies on PDA la , sp eading, wi h mode a e ae-
ial mycelium and e en ma gin, whi e o awn, eaching 70 mm diam. a e 2
weeks a 25 °C, s e ile.
Ma e ials examined. China • Xinjiang Uygu Au onomous Region, Al ay P e-
ec u e, Al ay Ci y, Alahake Town, Haxionggou Valley, om b anches o Be ula
pendula, 7 Oc obe 2024, Rong Ma, Caixia Wang & Hailong Lu (XJAU 4059 cul-
u es CFCC 71654 and CFCC 71655); ibid. (XJAU 4060, cul u es CFCC 71656,
CFCC 71657 and CFCC 71658).
No es. The species concep o C yp ospo ella be ulae was concei ed mo e
na owly han p e ious s udies, wi h only he sexual mo ph desc ibed (Mejía e
al. 2008, 2011). This ungus has been eco ded in Aus ia and Russia inhabi ing
Be ula len a and B. pendula (Mejía e al. 2008, 2011). In his s udy, wo new
specimens collec ed om Xinjiang, China and i e isola es we e ob ained. They
we e iden i ied as C . Be ulae, based on he molecula phylogeny (Fig. 5), ep e-
sen ing a new hos eco d in China. In addi ion, asexual mo ph o his ungus is
i s ly disco e ed and desc ibed he ein.
C yp ospo ella omen ella (Peck) L.C. Mejía, Mycologia 103(2): 397 (2011)
Fig. 11
Desc ip ion. Pseudos oma a imme sed o semi-imme sed in he ba k,
sca e ed, conical, 1450–1800 μm diam., 450–600 μm high, wi h 8–15
pe i hecia a anged ci cula ly o i egula ly. Ec os oma ic disc b own, ci cula o
o oid, 350–450 μm diam. Os ioles b own o black, 75–130 μm diam. Pe i hecia
lask-shaped o sphe ical, 250–400 μm diam. Asci hyaline, wi hou e ac i e
ing, cla a e, (85.5–)108–115(–126.5) × (9–)12.5–20(–21.5) μm, 8-spo ed.
Ascospo es 2–4-se ia e, cylind ical, sligh ly cu ed, ape ing owa ds ounded
ends, hin-walled, hyaline, asep a e, (39–)47–61.5(–67) × (4.5–)5–5.5(–6.5)
(a . = 54.2 ± 7 × 5.2 ± 0.6, n = 50) μm, L/W a io = 9–11.8.
Cul u e cha ac e is ics. Colonies on PDA la , sp eading, wi h mode a e ae-
ial mycelium and undula e ma gin, yellowish, eaching 70 mm diam. a e 2
weeks a 25 °C, s e ile.
Ma e ials examined. China • Xinjiang Uygu Au onomous Region, Al ay P e ec-
u e, Jeminay Coun y, Kizilkayin Red Bi ch Fo es , om b anches o Be ula mic o-
phylla, 5 Oc obe 2024, Rong Ma, Caixia Wang & Hailong Lu (XJAU 4005, cul u e
CFCC 71659); ibid. (XJAU 4006, cul u es CFCC 71660, CFCC 71661 and CFCC
71662); ibid. (XJAU 4010, cul u es CFCC 71663, CFCC 71664, CFCC 71665).
No es. C yp ospo ella omen ella was p e iously ega ded as a synonym o
C . be ulae (Reid and Boo h 1987; Mejía e al. 2008). This species has been cu -
en ly only documen ed in he Uni ed S a es, whe e i in ec s Be ula populi olia
and o he Be ula species (Mejía e al. 2008, 2011). The ascospo e mo pholo-
gy o ou specimens was consis en wi h ha desc ibed o he C . omen ella
lec o ype (Mejía e al. 2011). In he p esen s udy, we epo o he i s ime
he occu ence o C . omen ella on Be ula mic ophylla in China, suppo ed by
molecula phylogene ic e idence (Fig. 5).
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Caixia Wang e al.: Diapo hales om Be ula in Xinjiang, China
Melanconidaceae G. Win e [as ‘Melanconideae’], Rabenh. K yp .-Fl., Edn 2
(Leipzig) 1.2: 764 (1886)
No es. Melanconidaceae was es ablished by Win e (1886). This amily ini ial-
ly encompassed nume ous gene a cha ac e ised by pe i hecia imme sed in
well-de eloped s oma a, wi h os ioles eme ging h ough an ec os oma ic disc
(Ba 1978). Howe e , based on analyses o LSU sequence da a, Cas lebu y e al.
(2002) and Rossman e al. (2007) la e educed his amily o include only he ype
genus, Melanconis. Fan e al. (2018b) u he con i med ha Melanconidaceae
should be ea ed as a mono ypic amily, con aining only he genus Melanconis.
Melanconis Tul. & C. Tul., Selec . ung. ca pol. (Pa is) 2: 115 (1863)
No es. Melanconis is cha ac e ised by ci cula ly a anged pe i hecia imme sed
in well-de eloped o educed en os oma a wi h a concolou ous cen al col-
Figu e 11. Mo phology o C yp ospo ella omen ella om Be ula mic ophylla. A, B. Habi o ascos oma a on b anch;
C. Os ioles embedded in ascos oma in sec ion; D. T ans e se sec ion h ough ascos oma; E. Longi udinal sec ion
h ough ascos oma; F–H. Asci; I. Ascospo es. Scale ba s: 500 μm (B–E); 20 μm (F–I).
198
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Caixia Wang e al.: Diapo hales om Be ula in Xinjiang, China
Cai G, Zhao Y, Zhai Y, Yan M, Ma R, Zhang D (2024) Two new species o Cy ospo a
(Diapo hales, Cy ospo aceae) causing canke disease o Malus domes ica and
M. sie e sii in Xinjiang, China. MycoKeys 109: 305–318. h ps://doi.o g/10.3897/my-
cokeys.109.131456
Ca bone I, Kohn LM (1999) A me hod o designing p ime se s o specia ion s udies in
ilamen ous ascomyce es. Mycologia 3(3): 553–556. h ps://doi.o g/10.1080/00275
514.1999.12061051
Cas lebu y LA, Rossman AY, Jakli sch WJ, Vasilye a LN (2002) A p elimina y o e iew
o he Diapo hales based on la ge subuni nuclea ibosomal DNA sequences. Myco-
logia 94: 1017–1031. h ps://doi.o g/10.2307/3761867
C ous PW, Cos a MM, Kandemi H, Ve maas M, Vu D, Zhao L, Abell SE, A umugam E,
Flakus A, Ju je ić Ž, Kaliyape umal M, Mahade akuma S, Ma ney TS, Mu ugadoss
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