245
Un eiling species di e si y wi hin Mo ie ellomyco a om China
X: Th ee new species in Linnemannia and one in Mo ie ella
Xin-Yu Ji1, Zi-Ying Ding1, Wen-Xiu Liu1, Fei Li1, Heng Zhao2, Shi Wang1, Xiao-Yong Liu1,3
1 College o Li e Sciences, Shandong No mal Uni e si y, Jinan 250358, China
2 CAS Key Labo a o y o Fo es Ecology and Sil icul u e, Ins i u e o Applied Ecology, Chinese Academy o Sciences, Shenyang 110016, China
3 Ins i u e o Mic obiology, Chinese Academy o Sciences, Beijing 100101, China
Co esponding au ho : Xiao-Yong Liu ([email p o ec ed])
Copy igh : © Xin-Yu Ji 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
The species o he Mo ie ellaceae amily a e di e se and widely dis ibu ed. Fou new
species in his amily a e p oposed om hizosphe e soil h ough a comp ehensi e
axonomic app oach ha combined mul i-locus (SSU-ITS-LSU-RPB1-Ac ) phylogene -
ic analyses wi h de ailed mo phological examina ion. This s udy desc ibes and illus-
a es hese axa, cla i ying hei mo phological ea u es om closely ela ed species
and hei phylogene ic posi ions wi hin he amily. Linnemannia chlamydospo a sp. no .
(phylogene ically p oxima e o L. longigemma a) is cha ac e ized by he abundan p o-
duc ion o hick-walled chlamydospo es. Linnemannia o alispo a sp. no . (a sis e axon
o L. ugosa) is dis inguished by i s o al spo angiospo es. Linnemannia yunnanensis
sp. no . (closely allied o L. bainie ella) is cha ac e ized by i s o al chlamydospo es and
is named a e Yunnan P o ince, i s ype locali y. Mo ie ella i egula ispo a sp. no . (clus-
e ing wi h M. pa ispo a) is dis inguished by i s i egula ly shaped spo angiospo es. As
his is he en h ins almen o ou sys ema ic su ey o Mo ie ellomyco a di e si y in
China, his s udy expands he global species in en o y o Mo ie ellaceae o 158.
Key wo ds: Mo ie ellaceae, Muco omyco a, mul i-gene phylogeny, axonomy, Zygomyco a
In oduc ion
Mo ie ellaceae species ha e an impo an indus ial and ecological alue.
Species wi hin he amily Mo ie ellaceae a e no able o hei abili y o syn-
hesize polyunsa u a ed a y acids (PUFAs), such as a achidonic acid (ARA),
which a e aluable o bio uel p oduc ion and widely u ilized in comme cial
sec o s (Holland 2001; Yada e al. 2014; Telaga ho i e al. 2022). Some spe-
cies can also syn hesize enzymes (lipase, cellulase) wi h po en ial applica ions
in bio uels and ood p ocessing (Holland 2001; Wagne e al. 2013; Yada e
al. 2014; Telaga ho i e al. 2022). Many Mo ie ellaceae species also p oduce
bioac i e an imic obial me aboli es ha se e as e ec i e biocon ol agen s
agains plan pa hogens (Shemshu a e al. 2018). Some Mo ie ellaceae spe-
cies can pa icipa e in o ganic ma e mine aliza ion as decompose s, p omo e
soil nu ien cycling, and imp o e plan s ess esis ance (Holland 2001; Wagne
e al. 2013; Yada e al. 2014; Shemshu a e al. 2018; Telaga ho i e al. 2022).
Academic edi o :
Ch is ian Wu zbache
Recei ed:
12 Augus 2025
Accep ed:
31 Oc obe 2025
Published:
20 No embe 2025
Ci a ion: Ji X-Y, Ding Z-Y, Liu W-X, Li
F, Zhao H, Wang S, Liu X-Y (2025)
Un eiling species di e si y wi hin
Mo ie ellomyco a om China X:
Th ee new species in Linnemannia
and one in Mo ie ella. MycoKeys 125:
245–262. h ps://doi.o g/10.3897/
mycokeys.125.168474
MycoKeys 125: 245–262 (2025)
DOI: 10.3897/mycokeys.125.168474
246
MycoKeys 125: 245–262 (2025), DOI: 10.3897/mycokeys.125.168474
Xin-Yu Ji e al.: Th ee new species in Linnemannia and one in Mo ie ella
Addi ionally, ce ain membe s o his amily exhibi unique ecological and bio-
echnological capabili ies, including plan g ow h p omo ion, es uc u ing o
hizosphe e bac e ial communi ies, and decomposi ion o o ganic li e , high-
ligh ing hei mul i ace ed oles in bo h indus ial and en i onmen al con ex s
(Li e al. 2020; Ozimek and Hanaka 2021).
Mo ie ellaceae belongs o Mo ie ellomyco a, Mo ie ellomyco ina,
Mo ie ellomyce es, and Mo ie ellales (h p://www.index ungo um.o g/,
accessed on 14 May 2025) (Smi h e al. 2013; Wijayawa dene e al. 2024).
Mo ie ellaceae species can usually be isola ed om soil, plan hizomes, an-
imal emains, and mosses (Tede soo e al. 2014). The ypical cha ac e is ics
o Mo ie ellaceae species include whi e colony colo , ose e-shaped colony
mo phology, and a dis inc i e odo gene ally desc ibed as esembling ga lic o
we dog hai (Linnemann 1941; Gams 1977; Pe ko i s e al. 2011). This axon
is widely dis ibu ed h oughou he coun y (Linnemann 1941; Holland 2001;
Wagne e al. 2013; Yada e al. 2014; Shemshu a e al. 2018; Li e al. 2020;
Ozimek and Hanaka 2021; Telaga ho i e al. 2022). GBIF da abase documen s
Mo ie ellaceae om A ica (22,727 eco ds, 5.59%), An a c ica (2,889, 0.71%),
Asia (42,876, 10.55%), Oceania (47,301, 11.64%), Eu ope (231,446, 56.97%),
No h Ame ica (26,872, 6.61%) and Sou h Ame ica (32,155, 7.91%; h ps://
www.gbi .o g/, accessed on 22 May 2025). In summa y, he species o he am-
ily Mo ie ellaceae a e mainly concen a ed in Eu ope.
In ecen yea s, Mo ie ellaceae has seen he disco e y o many new species
o Mo ie ella and Linnemannia (Holland 2001; Wagne e al. 2013; Yada e al.
2014; Telaga ho i e al. 2022), bu o he new gene a emain o be u he s ud-
ied. The Ca alogue o Li e da abase con ains 17 gene a wi h a o al o 144 spe-
cies. O hese, Mo ie ella con ains he la ges numbe o species, 80, ollowed
by Linnemannia wi h 24, hen Podia and En omo ie ella in equal hi d place,
bo h wi h nine (h ps://www.ca alogueo li e.o g/, accessed on 22 May 2025).
In his s udy, ex ensi e ield sampling in Yunnan and Xizang, combined wi h
de ailed labo a o y analyses, esul ed in he disco e y o h ee new species in
Linnemannia and one in Mo ie ella. The newly desc ibed species a e Linnemannia
chlamydospo a, L. o alispo a, L. yunnanensis, and Mo ie ella i egula ispo a.
These species we e iden i ied based on molecula phylogene ic e idence, mo -
phological cha ac e is ics, and g ow h empe a u e p o iles. This is he en h e-
po in a se ies o s udies on he di e si y o Mo ie ellomyco a ac oss China
(Tao e al. 2024; Wang e al. 2024; Zhao e al. 2024; Ding e al. 2025a; Ding e al.
2025b; Ji e al. 2025a; Ji e al. 2025b; Wang e al. 2025). These indings no only
en ich he species di e si y o he amily Mo ie ellaceae bu also p o ide new
di ec ions o he axonomic and e olu iona y s udy o he amily.
Ma e ials and me hods
Isola ion
Soil samples we e collec ed om Yunnan and Xizang in 2024 ollowing he
p o ocols es ablished by Zou e al. (2022) and Liu e al. (2019). Each soil
sample (app oxima ely 100 g) was ans e ed o s e ile polye hylene bags
labeled wi h collec ion da e, ege a ion ype, ele a ion, and GPS coo dina es
(la i ude/longi ude) (Ra hnayaka e al. 2025). All samples we e s o ed a
247
MycoKeys 125: 245–262 (2025), DOI: 10.3897/mycokeys.125.168474
Xin-Yu Ji e al.: Th ee new species in Linnemannia and one in Mo ie ella
4 °C pos - anspo a ion un il labo a o y p ocessing. Pu e s ains we e iso-
la ed om he soil samples using a combina ion o soil dilu ion pla ing and
mois -chambe cul i a ion me hods (Zhao e al. 2021). Soil suspensions we e
p epa ed by homogenizing app oxima ely 1 g o soil sample in 10 mL o s e -
ile deionized wa e wi hin a 15 mL conical ube, ollowed by mechanical agi-
a ion on a o ex mixe o 25 minu es a 1,500 pm o achie e ho ough dis-
pe sion. Se ial dilu ions we e pe o med by ans e ing 1 mL o he p ima y
suspension in o 9 mL o s e ile deionized wa e , gene a ing a 10−2 dilu ion.
This p ocess was epea ed sequen ially o ob ain 10−3 and 10−4 dilu ions. Fo
ungal isola ion, 200 μL aliquo s o he 10−3 and 10−4 dilu ions we e asep ically
pipe ed on o Rose Bengal Chlo amphenicol (RBC) aga pla es. The medium
con ained pe li e : pep one (5.00 g), KH2PO4 (1.00 g), MgSO4·7H2O (0.50 g),
Rose Bengal dye (0.05 g), glucose (10.00 g), chlo amphenicol (0.10 g), and
aga (15.00 g), adjus ed o pH 6.8 ± 0.2 (Co y e al. 1995). Samples we e
e enly dis ibu ed using lame-s e ilized glass sp eade s and incuba ed a
26 °C unde ligh - es ic ed condi ions o 2–5 days o p omo e ungal colony
de elopmen . Subsequen ly, ungal colonies exhibi ing ac i e hyphal g ow h
ma gins we e selec i ely sub-cul u ed on o esh Po a o Dex ose Aga (PDA:
20 g/L glucose, 200 g/L po a o in usion, 20 g/L aga , pH 5.6 ± 0.2) o mal
ex ac aga (MEA: 33.6 g/L mal ex ac , 20 g/L aga ) using inocula ion nee-
dles. Mac omo phological ea u es we e documen ed wi h a high- esolu ion
digi al imaging sys em (Canon Powe Sho G7X, Canon, Tokyo, Japan). Fo
he we -chambe p o ocol, homogenized soil aliquo s (1 g) we e asep ically
sp ead on PDA pla es, sealed, and in e ed incuba ed a 15 °C (±0.5 °C) in he
da k o simula e he subsu ace niche. A e 48–72 h o incuba ion, isola e
p ima y ungal colonies by quad an s eak using lame-s e ilized inocula ion
loops. A e wo days, he aga con aining mycelia a he edge o he colony
was ans e ed o esh PDA o MEA, and cul u e as desc ibed abo e. A e
abou i e days, he s ain g ew well.
Mo phological obse a ion
Lac ophenol co on blue (LPCB) s aining d ople s we e added o he glass
slide. Then, a small piece o ape was ouched o he mycelial su ace, caus-
ing some hyphae o adhe e o i . The s ained specimen was imme sed in a
lac ol co on blue (LCB) solu ion o easy obse a ion mo phological analysis
u ilized a s e eoscope (Olympus SZX10, Olympus, Tokyo, Japan) and a ligh
mic oscope (Olympus BX53, OLYMPUS, Tokyo, Japan), and a high-de ini ion
colo digi al came a (Olympus DP80 OLYMPU, Tokyo, Japan) o obse e hy-
phal s uc u es and ep oduc i e o gans (Jiang e al. 2024; Tao e al. 2024;
Wang e al. 2024; Ding e al. 2025a; Ding e al. 2025b; Ji e al. 2025a; Ji e
al. 2025b; Wang e al. 2025). Mo phome y analysis was pe o med using
Digimize so wa e ( 5.6.0) wi h a leas 15 indi iduals measu ed pe mo -
phological ai . To de e mine he minimum and maximum g ow h empe a-
u es o he s ains, he empe a u e g adien me hod was used. Fi s , ini i-
a e a he mal acclima iza ion p o ocol by incuba ing he p ima y cul u e a
10 °C (±0.5 °C) o 48 h o s abilize ungal me abolism. Subsequen ly, he
incuba ion empe a u e was educed by 1 °C inc emen s pe day un il adial
g ow h s opped. The inal empe a u e be o e g ow h a es is designa ed as
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MycoKeys 125: 245–262 (2025), DOI: 10.3897/mycokeys.125.168474
Xin-Yu Ji e al.: Th ee new species in Linnemannia and one in Mo ie ella
he minimum g ow h empe a u e. All s ains we e s o ed in 10% s e ile glyc-
e ol a -20 °C. The li ing cul u es we e s o ed in he China Mic obiological
Cul u e Collec ion Cen e , Beijing, China (CGMCC). Equi alen s ains we e
p ese ed in he Shandong No mal Uni e si y Cul u e Collec ion (XG). D y
cul u es o ypes we e submi ed o he He ba ium Mycologicum Academiae
Sinicae, Beijing, China (Funga ium; HMAS). The axonomic in o ma ion was
deposi ed o he Fungal Names eposi o y (h ps://nmdc.cn/ ungalnames/).
DNA ex ac ion, PCR ampli ica ion, and sequencing
Genomic DNAs we e ex ac ed using he DNA Ex ac ion Ki (Ca . No.: 70409-20;
Bea e Biomedical Enginee ing Co., L d.) (Doyle e al. 1990; Wang e al. 2023). Ta -
ge egions (ITS, LSU, SSU, RPB1, and Ac ) we e ampli ied h ough PCR wi h p ime
pai s and p o ocols ou lined in Table 1. Reac ions we e pe o med in a 25 μL i-
nal olume con aining 12.5 μL o 2 × Hie Canace Plus PCR Mas e Mix wi h dye
(Yeasen Bio echnology, Ca No. 10154ES03), 9.5 μL o ddH2O, 1 μL o o wa d p im-
e (10 μM), 1 μL o e e se p ime (10 μM), and 1 µL o empla e genomic DNA
(1 ng/μL). Ampli ied p oduc s we e isualized on a 2% aga ose gel a 254 nm and
pu i ied using he Gel Ex ac ion Ki (Ca # AC0101-C; Shandong Spa kjade Bio ech-
nology Co., L d.), designed o e icien eco e y o DNA agmen s anging om
100 bp o 20 kb (Zhang e al. 2022; Zhang e al. 2025). The same supplie also
p o ided he RNA Rapid Ex ac ion Ki (Ca # AC0305; Shandong Spa kjade Bio-
echnology Co., L d.), which was a ailable o o al RNA ex ac ion had ansc ip-
omic analyses been equi ed. DNA sequencing was pe o med by Beijing Tsingke
Bio ech Co., L d. All sequences gene a ed in his s udy we e deposi ed in GenBank
(accession numbe s p o ided in he Suppl. ma e ial 1), in acco dance wi h s an-
da dized submission p o ocols o public da a accessibili y.
Phylogene ic analyses
The newly acqui ed sequence da a we e p ocessed using MEGA 7 o en-
su e consis ency (Kuma e al. 2016; La sson 2014). Re e ence sequences
o Mo ie ellaceae we e e ie ed om GenBank using he me hodology
Table 1. PCR in o ma ion used in his s udy.
Locus PCR
p ime s P ime sequences (5’–3’) PCR cycle Re e ence
ITS ITS5
ITS4
GGA AGT AAA AGT CGT AAC AAG G TCC
TCC GCT TAT TGA TAT GC
95 °C 5 min; (95 °C 30 s, 55 °C 30 s,
72 °C 1 min) × 35 cycles; 72 °C 10 min
(Whi e e al. 1990)
LSU LR0R
LR5
GTA CCC GCT GAA CTT AAG C TCC TGA
GGG AAA CTT CG
95 °C 5 min; (94 °C 30 s, 52 °C 45 s,
72 °C 90 s) × 30 cycles; 72 °C 10 min
(Hu deal e al. 2023)
SSU NS1
NS4
GTA GTC ATA TGC TTG TCT CC CTT CCG
TCA ATT CCT TTA AG
95 °C 5 min; (94 °C 60 s, 54 °C 50 s,
72 °C 60 s) × 37 cycles; 72 °C 10 min
(Hu deal e al. 2023)
RPB1 RPB1-A
RPB1-C
GAR TGY CCD GGD CAY TTY GG CCN GCD
ATN TCR TTR TCC ATR TA
95 °C 3 min; (94 °C: 40 s, 60 °C: 40 s, 72 °C:
2 min) × 9 (94 °C: 45 s, 55 °C: 1.5 min,
72 °C: 2 min) × 37 cycles; 72 °C 10 min
(S ille and Hall 1997)
Ac ACT-1
ACT-4R
TGG GAC GAT ATG GAI AAI ATC TGG CA TC
ITC GTA TIC TIG CTI IGA IAT CCA CA T
95 °C 3 min; (95 °C: 60 s, 55 °C: 60 s, 72 °C:
1 min) × 30 cycles; 72 °C 10 min
(Voig and
Wös emeye 2000)
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MycoKeys 125: 245–262 (2025), DOI: 10.3897/mycokeys.125.168474
Xin-Yu Ji e al.: Th ee new species in Linnemannia and one in Mo ie ella
o Telaga ho i e al., and phylogene ic analyses we e conduc ed o each
gene ic ma ke (Telaga ho i e al. 2022). The e olu iona y ela ionships wi h-
in Mo ie ellaceae we e econs uc ed h ough bo h Maximum Likelihood
(ML) and Bayesian In e ence (BI) app oaches, implemen ed ia he CIPRES
Science Ga eway (h ps://www.phylo.o g/, accessed 16 May 2025) (Nie
e al. 2020a; Nie e al. 2020b). The maximum likelihood (ML) analysis was
pe o med using RAxML e sion 8.2.4 on he CIPRES Science Ga eway Ve -
sion 3.3 pla o m, wi h 1,000 boo s ap eplica es conduc ed o assess he
obus ness o he phylogene ic ee (Mille e al. 2010; Nguyen e al. 2015).
The Bayesian in e ence (BI) analysis was conduc ed using he GTR + I + G
model, wi h samples collec ed e e y 1,000 gene a ions. A o al o eigh cold
Ma ko chains we e un concu en ly o wo million gene a ions (Ronquis
e al. 2012; S ama akis 2014). The esul ing phylogene ic ees we e isually
op imized and anno a ed using iTOL (h ps://i ol.embl.de, accessed May 16,
2025) and Adobe Illus a o CC 2019 (Wang e al. 2023).
Resul s
Phylogeny
Fo Linnemannia, phylogene ic analyses we e pe o med on a da ase comp is-
ing 43 s ains ep esen ing 34 species, wi h Mo ie ella longicollis (CBS 879.97)
as an ou g oup (Genbank numbe s see Suppl. ma e ial 1: able S1). The se-
quence ma ix comp ises 4,863 conca ena ed cha ac e s: 1–690 (ITS), 691–
1,672 (LSU), 1,673–2,728 (SSU), 2,729–4,099 (RPB1), and 4,100–4,863 (Ac ).
Among hese cha ac e s, 1,130 a e pa simony-in o ma i e, along wi h 3,417
cons an and 316 pa simony-unin o ma i e. Bayesian ee opology is cong u-
en wi h ha o he ML ee (Fig. 1).
Fo Mo ie ella, phylogene ic analyses we e pe o med on a da ase com-
p ising 89 s ains ep esen ing 75 species, wi h Umbelopsis au o ophica (CBS
310.93) as an ou g oup (Genbank numbe s see Suppl. ma e ial 1: able S2).
The sequence ma ix comp ises 5,292 conca ena ed cha ac e s: 1–962 (ITS),
963–1,968 (LSU), 1,969–3,049 (SSU), 3,050–4,422 (RPB1), and 4,423–5,292
(Ac ). Among hese, 1,806 a e pa simony-in o ma i e, along wi h 2,581 con-
s an and 905 pa simony-unin o ma i e. Bayesian ee opology is consis en
wi h he ML ee (Fig. 2).
Taxonomy
Linnemannia chlamydospo a X.Y. Ji, H. Zhao & X.Y. Liu, sp. no .
Fig. 3
Fungal Names: FN 572057
Type. China • Xizang, Nyingchi Ci y, Bayi Dis ic (29°33'40"N, 94°33'13"E,
al i ude 3713 m), om soil, 29 Augus 2024, X.Y. Ji, holo ype HMAS 354073,
ex-holo ype li ing cul u e CGMCC 3.28892 (=XG10460-10-1).
E ymology. The epi he chlamydospo a (La .) e e s o a la ge numbe o
chlamydospo es.
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MycoKeys 125: 245–262 (2025), DOI: 10.3897/mycokeys.125.168474
Xin-Yu Ji e al.: Th ee new species in Linnemannia and one in Mo ie ella
Figu e 1. The Maximum Likelihood phylogene ic ee o he genus Linnemannia based on combined ITS, LSU, SSU, RPB1,
and Ac sequences wi h Mo ie ella longicollis as ou g oup. Node suppo s a e indica ed by wo me ics: Maximum
Likelihood Boo s ap Values (le , MLBV ≥ 70%) and Bayesian In e ence Pos e io P obabili ies ( igh , BIPP ≥ 0.90),
sepa a ed by a slash (/). No el species a e emphasized in ed. Bold en ies wi h as e isks (*) deno e ex- ype o ex-holo ype
s ains. The scale ba in he lowe le ep esen s 0.1 subs i u ions pe si e.
M.longicollis CBS 209.32*
L. iede ikiana P 3s8
L.hyalina CBS 223.35
L.sco diella HFSF81*
L.scle o iella CBS 529.68*
L.ac o ona CBS 386.71*
L.olea CGMCC 3.28579*
L.zychae CBS 316.52*
L.s ella is ks2-4*
L.nimbosa HFSF57*
L.exigua NNIBRFG5521
L.mannui P 2s5
L. a shede ae CBS 388.71*
L.elizabe hkennyiae BRIP 74948a*
L. lu iae EML-YR25716-1*
L.gamsii CBS 749.68*
L.diaoluoshan XG08196-6*
L.schmucke i CBS 295.59*
L.cama gensis CBS 221.58*
L.cama gensis CBS 221.58*
L.amoeboidea CBS 889.72*
L. ama indoides CGMCC 3.28576*
L. hizomo pha XG07310-1*
L.bi amosa RS5
L.bi amosa SYFGD6-2
L.bi amosa SYFGP2-1
L.nan ahalensis CBS 610.70*
L.b e ispho a CGMCC 3.28577*
L.soli a ia OAS3*
L.elonga a 7
L. ugosa XG10541-6*
L.o alispo a XG09524-11-2
L.o alispo a CGMCC 3.28891*
L.yunnanensis CGMCC 3.28890*
L.yunnanensis XG08668-7-2
L.longigemma a CBS 653.93*
L. o unda CGMCC 3.28764*
L. o unda XG08755-7-2
L.chlamydospo a CGMCC 3.28892*
L.chlamydospo a XG10460-10-2
L. bainie ella P 1s13
L. bainie ella P 1s20
L. bainie ella P 1s21
91/0.99
99/0.99
99/0.98
100/0.99
92/0.99
93/1
73/0.95
78/0.92
81/0.94
93/0.97
100/1
73/0.96
75/0.99
95/0.99
83/0.96
91/0.99
0.1
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MycoKeys 125: 245–262 (2025), DOI: 10.3897/mycokeys.125.168474
Xin-Yu Ji e al.: Th ee new species in Linnemannia and one in Mo ie ella
Umbelopsis au o ophica CBS 310.93*
M.longicollis CBS 209.32*
M.capi a a CBS 110.640
M.wol ii CBS 651.93
M.wol ii CBS 209.69
M.wol ii CBS 612.70
M.mic ozygospo a CBS 880.97*
M.epigama CBS 489.70*
M. e icilla a CBS 346.66
M.humilis FSU828
M.humilis CBS 745.68
M.humilis CBS 222.35*
M.ho icola CBS 305.52*
M.clonocys is CBS 357.76*
M.minu issima CBS 307.52
M.epicladia CBS 355.76*
M.lapis OBS3*
M.an a c ica CBS 609.70*
M. iangula is OAS8
M.alpina CBS 210.32
M.alpina CBS 210.32*
M.mul ispo a KUMCC 20-0005*
M.calciphila WA 18944*
M. o micicola CBS 109.589
M.pa aensis CBS 547.89
M.beljako ae CBS 123.72*
M.lignicola CBS 207.37*
M.gemmi e a CBS 134.45*
M.kuhlmanii CBS 157.71*
M.spinospo a XG06904-41
M.echinosphae a CBS 575.75*
M.chlamydospo a CBS 120.34
M.mu abilis CBS 308.52*
M.s ylospo a CBS 211.32*
M.simplex CBS 243.82
M.angus a CBS 293.61*
M.pa azychae CBS 868 71*
M.s angula a CBS 455.67*
M. os a inskii CBS 522.70*
M.selenospo a CBS 811.68*
M.hypsicladia CBS 116.202*
M.indohii FSU830
M.indohii FSU831
M.indohii CBS 720.71*
M.polycephala FSU696
M.polycephala FSU866
M.polygonia CBS 685.71*
M.acu a XG08182-4-2
M.acu a CGMCC 3.28761*
M.amoeboidea CBS 889.72*
M.yunnanensis KUMCC 20-0009*
M.globalpina CBS 360.70*
M.basipa ispo a CBS 517.72*
M. o micae WA 49853*
M.dicho oma CBS 221.35*
M. ibe ensis XG00421-2-2
M. ibe ensis CGMCC 3.28763*
M.pa ispo a CBS 311.52
M.i egula ispo a CGMCC 3.28893*
M.i egula ispo a XG00435-2-2
M.mac ocys is CBS 314.85
M.oedema CGMCC 3.28762*
M.oedema XG00420-1-2
M.elonga ula CBS 488.70*
M.cys ojenkinii CBS 456.71*
M.pulchella CBS 312.52
M. u icola CBS 432.76*
M. imb icys is CBS 943.70*
M.a milla iicola CBS 914.73*
M.bainie i CBS 220.35
M.zona a CBS 228.35*
M. e ucosa CBS 181.73
M.his oplasma oides CBS 321.78*
M.cogi ans CBS 879.97*
M.scle o iella CBS 529.68*
M.gamsii CBS 749.68*
M.gamsii CBS 551.73
M.sa nyensis CBS 122.72*
M.nan ahalensis CBS 610.70*
M.wuyishanensis CBS 370.95*
M.bi amosa CBS 370.95
M.schmucke i CBS 295.59*
M.cama gensis CBS 221.58*
M.zychae CBS 316.52*
M.exigua CBS 655.68*
M.ac o ona CBS 386.71*
M.elonga a FSU823
M.elonga a FSU822
M. ishikesha CBS 652.68*
81/0.90
99/1
96/1
100/1
100/1
92/1
96/1
97/0.99
100/1
100/0.99
100/1
100/1
81/1
93/1
100/1
100/0.99
100/1
100/1
73/1
81/0.99
91/1
100/1
100/1
100/1
70/1
98/1
100/1
99/0.99
100/0.99
95/1
100/1
82/0.99
100/0.99
95/1
71/1
98/1
100/1
0.1
Figu e 2. The Maximum Likelihood phylogene ic ee o he genus Mo ie ella based on combined ITS, LSU, SSU, RPB1,
and Ac sequences wi h Umbelopsis au o ophica as an ou g oup. Node suppo s a e indica ed by wo me ics: Maximum
Likelihood Boo s ap Values (le , MLBV ≥ 70%) and Bayesian In e ence Pos e io P obabili ies ( igh , BIPP ≥ 0.90),
sepa a ed by a slash (/). No el species a e emphasized in ed. Bold en ies wi h as e isks (*) deno e ex- ype o ex-holo ype
s ains. The scale ba in he lowe le ep esen s 0.1 subs i u ions pe si e.
252
MycoKeys 125: 245–262 (2025), DOI: 10.3897/mycokeys.125.168474
Xin-Yu Ji e al.: Th ee new species in Linnemannia and one in Mo ie ella
Desc ip ion. Colonies on PDA a 16 °C o 5 d, eaching 70 mm diame e , as
g owing wi h a a e o 14 mm/d, ga lic smell, annual ing-like. Hyphae hyaline,
2.0–7.8 µm wide (n = 15, x
– = 4.8). Spo angia mos ly sphe ical, smoo h, hyaline,
9.0–12.7 µm in diame e (n = 15, x
– = 11.2). Spo angiospo es hyaline, smoo h,
mos ly ound, 5.8–12.7 µm in diame e (n = 15, x
– = 9.6). Chlamydospo es abun-
dan , o al, ound, and i egula , 11.2–33.6 µm long and 6.8–21.7 µm wide (n =
15, x
– =17.4 × 13.0 µm). Zygospo es no ound.
Figu e 3. Linnemannia chlamydospo a ex-holo ype CGMCC 3.28892. a, b. Colonies on PDA (a. ob e se; b. e e se);
c, d. Spo angia; e–i. Chlamydospo es; j–m. Spo angiospo es. Scale ba s: 10 µm (c–m).
253
MycoKeys 125: 245–262 (2025), DOI: 10.3897/mycokeys.125.168474
Xin-Yu Ji e al.: Th ee new species in Linnemannia and one in Mo ie ella
Tempe a u e equi emen s. Minimum g ow h empe a u e 4 °C and maximum
g ow h empe a u e 28 °C.
Addi ional s ains examined. China • Xizang, Nyingchi Ci y, Bayi Dis ic
(29°33'40"N, 94°33'13"E, al i ude 3713 m), om soil, 29 Augus 2024, X.Y. Ji,
li ing cul u e XG10460-10-2.
No es. The phylogene ic analysis showed ha he new species
L. chlamydospo a is closely ela ed o L. longigemma a (Fig. 2). I is dis in-
guished om L. longigemma a by 55/621 cha ac e s in ITS sequences. Mo pho-
logically, compa ed o L. longigemma a, he new species has la ge spo angio-
spo es (5.8–12.7 µm s 5.0–9.0 µm) and smalle chlamydospo es (11.2–33.6
× 6.8–21.7 µm s up o 60.0 µm).
Linnemannia o alispo a X.Y. Ji, H. Zhao & X.Y. Liu, sp. no .
Fig. 4
Fungal Names: FN 572920
Type. China • Yunnan P o ince, Yuxi Ci y, Resea ch and se ice a ea o Hong a
Dis ic (24°15'41"N, 102°28'58"E, al i ude 1632.18 m), om soil, 15 Ma ch
2024, X.Y. Ji, holo ype HMAS 354072, ex-holo ype li ing cul u e CGMCC
3.28891 (=XG09524-11-1).
E ymology. The epi he o alispo a (La .) e e s o he o al spo angiospo es.
Figu e 4. Linnemannia o alispo a ex-holo ype CGMCC 3.28891. a, b. Colonies on MEA (a. ob e se; b. e e se);
c, d. Chlamydospo es; e–g. Spo angiospo es. Scale ba s: 10 µm (c–g).
260
MycoKeys 125: 245–262 (2025), DOI: 10.3897/mycokeys.125.168474
Xin-Yu Ji e al.: Th ee new species in Linnemannia and one in Mo ie ella
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Supplemen a y ma e ial 1
GenBank accession numbe s
Au ho s: Xin-Yu Ji, Zi-Ying Ding, Wen-Xiu Liu, Fei Li, Heng Zhao, Shi Wang, Xiao-Yong Liu
Da a ype: docx
Explana ion no e: GenBank accession numbe s o Linnemannia and Mo ie ella genus
used in his s udy. The newly disco e ed species iden i ied in he p esen s udy a e in
bold. Ex- ype s ains a e ma ked wi h a s a ma ke “*”. NA s ands o “no a ailable”.
Copy igh no ice: This da ase is made a ailable unde he Open Da abase License
(h p://openda acommons.o g/licenses/odbl/1.0/). The Open Da abase License
(ODbL) is a license ag eemen in ended o allow use s o eely sha e, modi y, and
use his Da ase while main aining his same eedom o o he s, p o ided ha he
o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/mycokeys.125.168474.suppl1
