Our current understanding of the genus Pseudocosmospora (Hypocreales, Nectriaceae) in China

Abstract

To explore species diversity of the genus Pseudocosmospora, collections from Beijing and Hubei Province of China were examined, and three undescribed species were encountered. Morphological features and DNA sequence analyses of the ITS, nc LSU rDNA, RPB1, and TUB regions support their placement in Pseudocosmospora and recognition of two new species. Pseudocosmospora curvispora sp. nov. encountered on fruiting bodies of a species of the Diatrypaceae has gregarious, subglobose to obpyriform perithecia seated directly on the host fungus clavate asci, with eight ellipsoidal, smooth ascospores, acremonium- to verticillium-like conidiophores, and strongly curved, allantoid, unicellular conidia. Pseudocosmospora shennongjiana sp. nov. is characterized by solitary to gregarious, subglobose to globose perithecia with a minute basal stroma clavate asci, containing eight ellipsoidal, spinulose ascospores, acremonium-like conidiophores, and rod-shaped, unicellular conidia. An unnamed species of the genus is also described. Distinctions between these species and their close relatives are compared in detail. The previously recorded species of the genus in China are briefly reviewed.

Full text

ORIGINAL ARTICLE
Our current understanding of the genus Pseudocosmospora
(Hypocreales, Nectriaceae) in China
Zhao-Qing Zeng
1
&Wen-Ying Zhuang
1
Received: 21 August 2020 /Revised: 12 January 2021 / Accepted: 14 January 2021
#German Mycological Society and Springer-Verlag GmbH Germany, part of Springer Nature 2021
Abstract
To explore species diversity of the genus Pseudocosmospora, collections from Beijing and Hubei Province of China were
examined, and three undescribed species were encountered. Morphological features and DNA sequence analyses of the ITS,
nc LSU rDNA, RPB1,andTUB regions support their placement in Pseudocosmospora and recognition of two new species.
Pseudocosmospora curvispora sp. nov. encountered on fruiting bodies of a species of the Diatrypaceae has gregarious,
subglobose to obpyriform perithecia seated directly on the host fungus clavate asci, with eight ellipsoidal, smooth ascospores,
acremonium- to verticillium-like conidiophores, and strongly curved, allantoid, unicellular conidia. Pseudocosmospora
shennongjiana sp. nov. is characterized by solitary to gregarious, subglobose to globose perithecia with a minute basal stroma
clavate asci, containing eight ellipsoidal, spinulose ascospores, acremonium-like conidiophores, and rod-shaped, unicellular
conidia. An unnamed species of the genus is also described. Distinctions between these species and their close relatives are
compared in detail. The previously recorded species of the genus in China are briefly reviewed.
Keywords Cosmospora-like fungi .Morphology .Multigene analyses .Taxonomy .Two new species
Introduction
The genus Pseudocosmospora C.S. Herrera & P. Chaverri,
typified by P. eutypellae C.S. Herrera & P. Chaverri, is char-
acterized by scattered to gregarious subglobose to obpyriform,
bluntly papillate perithecia that are collapsing laterally when
dry, scarlet, KOH+, and usually less than 250 μm in height;
cylindrical to narrowly clavate asci containing eight ellipsoi-
dal, yellow-brown, 1-septate and verrucose ascospores; and
producing acremonium- to verticillium-like asexual states, cy-
lindrical phialides, and ellipsoidal, ovoid or reniform, smooth,
non-septate conidia (Herrera et al. 2013). Some members pro-
duce bioactive secondary metabolites which play important
roles in biomedicine and agriculture (Lee et al. 2011; Shiono
et al. 2016; Nakamura et al. 2019). Fourteen species are
currently accepted in the genus. They are widely distributed,
and occur usually on stromata of diatrypaceous fungi, partic-
ularly species of Eutypa Tul.&C.Tul.andEutypella
(Nitschke) Sacc., and rarely on Biscogniauxia Kuntze and
Hypoxylon Bull. (Herrera et al. 2013; Zeng and Zhuang
2017;LechatandFournier2020).
Studies on Pseudocosmospora in China were initiated
by Teng (1934,1936) who described P. nummulariae
(Teng) W.Y. Zhuang & Z.Q. Zeng as Nectria
nummulariae Teng and P. effusa (Teng) Z.Q. Zeng &
W.Y. Zhuang as Phaeonectria manilensis var. effusa
Teng from Hainan Province. Seven species are currently
known from the country (Zhuang 2013; Zeng and Zhuang
2017,2020).
In connection with our studies on the Chinese fungus flora,
surveys of fungicolous species of Nectriaceae were carried
out, and three undescribed taxa are discovered based on mor-
phological characteristics and DNA sequence analyses of nu-
clear ribosomal DNA ITS1-5.8S-ITS2 (ITS), large subunit of
the nuclear ribosomal DNA (nc LSU rDNA), RNA polymer-
ase II subunit one (RPB1), and β-tubulin (TUB). Two are
introduced as new species and one is treated as an unnamed
species. The previously recorded species of the group in China
are briefly reviewed.
*Zhao-Qing Zeng
zengzq@im.ac.cn
*Wen-Ying Zhuang
zhuangwy@im.ac.cn
1
State Key Laboratory of Mycology, Institute of Microbiology,
Chinese Academy of Sciences, Beijing 100101, China
https://doi.org/10.1007/s11557-021-01672-1
Mycological Progress (2021) 20:419–429

Materials and methods
The specimens were collected from Beijing and Hubei
Province and deposited in the Herbarium Mycologicum
Academiae Sinicae (HMAS). The methods of Luo and
Zhuang (2010a) were followed for morphological observa-
tions. Perithecial wall reactions were tested in 3% potassium
hydroxide (KOH) and 100% lactic acid (LA). To observe
internal and microscopic characteristics of perithecial walls,
sections were made with a freezing microtome (YD-1508-III,
Jinhua, China) at a thickness of 6–8μm. Water and
lactophenol cotton blue solution were used as mounting media
for examinations of anatomic structures and measurements of
perithecia, asci, and ascospores. Photographs were taken with
a Leica DFC450 digital camera (Wetzlar, Germany) attached
to a Leica M125 stereomicroscope (Milton Keynes, UK) for
gross morphology and a Zeiss AxioCam MRc 5 digital cam-
era (Jena, Germany) attached to a Zeiss Axio Imager A2 mi-
croscope (Göttingen, Germany) for anatomical structures.
Measurements of individual structures were based on 30 units,
except as otherwise noted. Cultures were obtained from fresh
material using single ascospore isolation, and strains deposit-
ed in the China General Microbiological Culture Collection
Center (CGMCC). For colony features and growth rates,
strains were grown on potato dextrose agar [PDA, 20%
(w/v) potato + 2% (w/v) dextrose + 2% (w/v) agar] and syn-
thetic nutrient-poor agar (SNA; Nirenberg 1976)in90-mm
plastic Petri dishes at 25°C for 7 d with alternating periods
of light and darkness (12 h/12 h).
Genomic DNA was extracted from fresh mycelium follow-
ing the method of Wang and Zhuang (2004). Primer pair
ITS5/ITS4 (White et al. 1990) was used to amplify the ITS,
and LR0R/LR5 (Vilgalys and Hester 1990; Rehner and
Samuels 1994) was used for the partial nc LSU rDNA, and
Crpb1a/rpb1c (Castlebury et al. 2004) was used for the partial
RPB1, and T1/T2 (O’Donnell and Cigelnik 1997) was used
for the partial TUB. PCR reactions were performed using an
ABI 2720 Thermal Cycler (Applied Biosciences, Foster City,
USA) with a 25-μlreactionsystemconsistingof12.5-μlTaq
MasterMix, 1-μlofeachprimer(10μM), 1-μl template DNA,
and 9.5-μlddH
2
O. DNA sequencing was carried out in both
directions on an ABI 3730XL DNA Sequencer (Applied
Biosciences, Foster City, USA).
Newly acquired sequences and those retrieved from
GenBank are listed in Table 1. The sequences were assembled
and aligned, and the primer sequences were trimmed by using
BioEdit 7.0.5 (Hall 1999), and converted to NEXUS files by
ClustalX 1.83 (Thompson et al. 1997). A partition homoge-
neity test was performed with 1000 replicates in
PAUP*4.0b10 (Swofford 2002) to evaluate statistical congru-
ence among the four loci. The sequences of ITS, nc LSU
rDNA, RPB1,andTUB were combined and analyzed by
Bayesian inference (BI), maximum likelihood (ML), and
maximum parsimony (MP) methods to determine the phylo-
genetic positions of the studied taxa. The BI analysis was
conducted by MrBayes 3.1.2 (Ronquist and Huelsenbeck
2003) using a Markov Chain Monte Carlo algorithm.
MrModeltest 2.3 was used to determine the nucleotide substi-
tution models (Nylander 2004). Four Markov chains were run
simultaneously for 1,000,000 generations with the trees sam-
pled every 100 generations. A 50% majority rule consensus
tree was computed after excluding the first 2500 trees as
“burn-in.”Bayesian inference posterior probability (BIPP)
was determined from the remaining trees. The ML analysis
was performed via IQ-Tree 1.6.12 program (Nguyen et al.
2015) using the best model for each locus chosen by
ModelFinder (Chernomor et al. 2016). The MP analysis was
performed with PAUP 4.0b10 (Swofford 2002) using heuris-
ticsearcheswith1000replicatesofrandomadditionofse-
quences and subsequent TBR (tree bisection and reconnec-
tion) branch swapping. The topological confidence of the
resulting trees, and statistical supports of branches were tested
in maximum parsimony bootstrap analyses (MPBP) with
1000 replications and each with 10 replicates of random ad-
dition of taxa. Trees were examined by TreeView 1.6.6 (Page
1996). BIPP greater than 90% and maximum likelihood boot-
strap proportion (MLBP) and MPBP greater than 70% were
shown at the nodes.
Results
The sequences of ITS, nc LSU rDNA, RPB1,andTUB from
25 strains of 17 Pseudocosmospora species were analyzed.
The partition homogeneity test (P= 0.01) indicated that the
individual partitions were not highly incongruent
(Cunningham 1997); thus, these four loci were combined for
the phylogenetic analyses. In the MP analysis, the datasets
included 2587 nucleotide characters, of which 1574 bp were
constant, 503 were variable and parsimony-uninformative,
and 510 were parsimony-informative. The MP analysis result-
ed in a single most parsimonious tree (tree length = 2170,
consistency index = 0.6498, homoplasy index = 0.3502, re-
tention index = 0.6649, rescaled consistency index = 0.4320)
(Fig. 1). The topologies of ML and BI trees were similar with
topology of the MP tree. The final matrix was deposited in
TreeBASE with accession no. S26403. The isolates of
CGMCC 3.20176, 3.20177, and 3.20178 grouped with other
members of Pseudocosmospora, and the genus clade received
high statistical support values (MPBP/MLBP/BIPP = 91%/
100%/100%). The isolate CGMCC 3.20176 grouped together
with Pseudocosmospora sp. GJS 96-216 (MPBP/MLBP/
BIPP = 100%/100%/100%). The isolate CGMCC 3.20177
was shown as a separate lineage and grouped with
P. rogersonii C.S. Herrera & P. Chaverri, P. eutypellae,and
three unnamed strains. The isolate CGMCC 3.20178 clustered
420 Mycol Progress (2021) 20:419–429

with the unidentified strain MAFF 241531 (Herrera et al.
2013) (MPBP/MLBP/BIPP = 100%/100%/100%) and further
grouped with P. rogersonii, which formed a supported termi-
nal branch in Pseudocosmospora (MPBP/MLBP/BIPP =
99%/94%/100%).
Taxonomy
Pseudocosmospora curvispora Z.Q. Zeng & W.Y. Zhuang,
sp. nov. Fig. 2
Fungal names FN570727
Etymology: The specific epithet refers to the strongly
curved conidia.
Mycelium not visible on natural substratum.
Perithecia superficial, gregarious, non-stromatic,
subglobose to globose, with a truncate apex, laterally
collapsed upon drying, orange-red, reddish-brown to dark
brown, turning brownish red in KOH, becoming slightly
yellow in LA, 167–235 × 108–167 μm(n=14).
Perithecial wall of two layers, 15–30 μm thick; outer
layer of textura globulosa to textura angularis, with sur-
face slightly warted, 10–18 μm thick, cells 5–8×2.5–5
μm, walls 1–1.2 μm thick; inner layer of textura
prismatica, 5–12.5 μm thick, cells 5–10 × 2.5–3.5 μm,
walls 0.8–1μmthick.Asci clavate, 8-spored, with a sim-
ple apex, 53–68 × 3–5μm. Ascospores ellipsoidal, 1-
septate, light yellow-brown, smooth, uniseriate, and over-
lapping obliquely, 8–10 × 3–5μm.
Table 1 List of species, herbarium/strain numbers, and GenBank accession numbers of materials used in this study
Species Strain number GenBank numbers
ITS nc LSU
rDNA
RPB1 TUB
Corallomycetella repens (Berk. & Broome) Rossman & Samuels AR 4547 JF832594 JF832679 JF832763 JF832838
Microcera larvarum (Fuckel) Gräfenhan, Seifert & Schroers AR 4580 KC291751 KC291759 KC291894 KC291935
Pseudocosmospora curvispora Z.Q. Zeng & W.Y. Zhuang CGMCC
3.20176
MT592897
*
MT592879 MT606153 MT606156
Pseudocosmospora eutypae C.S. Herrera & P. Chaverri CH 11-01 KC291735 KC291766 KC291884 KC291925
IMI 73016 KC291736 KC291786 KC291885 -
AR 4527 KC291720 KC291756 KC291870 KC291909
Pseudocosmospora eutypellae C.S. Herrera & P. Chaverri AR 4562 KC291721 KC291757 KC291871 KC291912
GJS 10-248 KC291722 KC291772 KC291872 KC291911
Pseudocosmospora henanensis (Y. Nong & W.Y. Zhuang) W.Y. Zhuang &
Z.Q. Zeng
HMAS 183528 GU075856 GU075863 - HM054103
Pseudocosmospora hypoxylicola Lechat & J. Fourn. CLL 19020 MN886606 MN886608 - -
Pseudocosmospora joca (Samuels) C.S. Herrera & P. Chaverri AR 4779 KC291746 KC291762 KC291887 KC291924
Pseudocosmospora metajoca C.S. Herrera & P. Chaverri AR 4576 KC291745 KC291758 KC291886 KC291923
Pseudocosmospora rogersonii C.S. Herrera & P. Chaverri GJS 90-56 KC291729 KC291780 KC291878 KC291915
GJS 10-296 KC291727 KC291774 KC291876 KC291917
GJS 09-1384 KC291726 KC291770 KC291875 KC291914
Pseudocosmospora shennongjiana Z.Q. Zeng & W.Y. Zhuang CGMCC
3.20177
MT592898 MT592880 MT606154 MT606157
Pseudocosmospora vilior (Starbäck) C.S. Herrera & P. Chaverri AR 4771 KC291734 KC291761 KC291898 KC291926
AR 4810 KC291737 KC291763 KC291900 KC291928
PC 1246 KC291738 KC291791 KC291899 KC291927
Nectriaceae sp. GJS 95-143 KC291750 KC291782 KC291880 KC291920
MAFF 241499 KC291739 KC291789 KC291874 KC291913
MAFF 241531 KC291730 KC291790 KC291879 KC291918
Pseudocosmospora sp. CGMCC
3.20178
MT592899 MT592881 MT606155 MT606158
AR 4768 KC291724 KC291760 KC291881 KC291922
CH 11-02 KC291725 KC291767 KC291882 KC291919
GJS 95-141 KC291749 KC291781 KC291883 KC291921
GJS 96-216 KC291733 KC291783 KC291889 KC291930
*Italicized numbers indicate the newly provided sequences
421Mycol Progress (2021) 20:419–429

Colony on PDA 62 mm in diam. after 1 week at 25°C,
whitish with a pinkish tint, surface cottony, with dense aerial
mycelium. Colony on SNA 23 mm diam. after 1 week at
25°C, with sparse whitish aerial mycelium. Conidiophores
acremonium-like to verticillium-like, arising from aerial hy-
phae, septate, branches more or less verticillate, with terminal
whorl of 2–6phialides.Phialides subulate, tapering toward
apex, 10–55 μm long, 0.9–1.2 μm wide at base, 0.2–0.3 μm
wide at tip. Conidia allantoid, mainly strongly curved, unicel-
lular, smooth, hyaline, 3–5 × 0.8–1.2 μm.
Type: China, Beijing, Mentougou, Tanzhesi, on unidenti-
fied Diatrypaceae, 12 Jun 2018, W.Y. Zhuang, Z.Q. Zeng,
H.D. Zheng 11855 (HMAS 271239). Ex-type strain,
CGMCC 3.20176. Sequences obtained from ex-type strain:
ITS, MT592897; nc LSU rDNA, MT592879; RPB1,
MT606153; TUB, MT606156.
Distribution: Known only from type location.
Notes: Among the known species of Pseudocosmospora,
P. curvispora resembles P. rogersonii in having gregarious,
subglobose perithecia with a truncate apex, smooth asco-
spores, pinkish colony on PDA, unbranched conidiophores,
and non-guttulate conidia (Herrera et al. 2013). However,
P. rogersonii has much wider asci (5.7–8.4 μmvs.3–5μm)
and conidia (1.1–2.6 μm vs. 0.8–1.2 μm) that are straight
rather than strongly curved (Herrera et al. 2013). Both ITS
and nc LSU rDNA sequences of the ex-type strains of
P. curvispora (CGMCC 3.20176) and P. rogersonii (GJS
90-56) differ at 14 positions, while 77 differing bp positions
were found in TUB sequence comparisons.
Pseudocosmospora shennongjiana Z.Q. Zeng & W.Y.
Zhuang, sp. nov. Fig. 3
Fungal names FN570728
Etymology: The specific epithet refers to the type locality.
Mycelium not visible on natural substratum. Perithecia
superficial, solitary to gregarious, with a minute basal stroma,
subglobose to pyriform, laterally collapsed upon drying, or-
ange to orange-red, turning purplish red in KOH, becoming
slightly yellow in LA, 157–186 × 108–147 μm. Perithecial
wall of two layers, 18–30 μm thick; outer layer of textura
globulosa to textura angularis, with surface slightly rough-
ened, 13–22 μm thick, cells 3–6×2–3μm, walls 0.8–
1.2 μm thick; inner layer of textura prismatica, 5–8μmthick,
cells 6–13 × 2–3μm, walls 0.6–0.8 μm thick. Asci clavate, 8-
spored, with a simple apex, 45–65 × 4.5–6μm. Ascospores
ellipsoidal with ends narrower, 1-septate, light yellow-brown,
spinulose, uniseriate and overlapping obliquely, 7.5–12 × 3–5
μm.
Fig. 1 Single most parsimonious
tree generated from analyses of
combined ITS, nc LSU rDNA,
RPB1 and TUB sequences of
Pseudocosmospora species.
MPBP (left) and MLBP (middle)
greater than 70% and BIPP (right)
greater than 90% are shown at the
nodes
422 Mycol Progress (2021) 20:419–429

Colony on PDA 46 mm in diam. after 1 week at 25°C,
cream to pale yellow, surface somewhat cottony, with white
aerial mycelium. Colony on SNA 50 mm diam. after 1 week at
25°C, with sparse whitish aerial mycelium. Conidiophores
acremonium-like, arising from aerial hyphae, septate, un-
branched. Phialides subulate, tapering toward apex, 10–65
μm long, 0.6–0.8 μm wide at base, 0.1–0.2 μm wide at tip.
Conidia rod-shaped, unicellular, smooth, hyaline, 3–6 × 0.9–
1.3 μm.
Type: China, Hubei, Shennongjia, Muchengshaoqia, asso-
ciated with or on old fruiting bodies of an unidentified fungus,
22 Sept 2014, Z.Q. Zeng, H.D. Zheng, W.T. Qin & K. Chen
10058 (HMAS 273904). Ex-type strain, CGMCC 3.20177.
Sequences obtained from ex-type strain: ITS, MT592898; nc
Fig. 2 Pseudocosmospora curvispora.a–cPerithecia on natural
substratum. d Colony after 1 week at 25°C on PDA. eLongitudinal
section through perithecium in lactophenol cotton blue. fLongitudinal
section through perithecium in water. g, h Asci with ascospores. i-k
ascospore. l, n Conidiophores and conidia. m, o Conidia. Scale bars: a–
c1mm;e, f 50 μm; g–o10 μm
423Mycol Progress (2021) 20:419–429

LSU rDNA, MT592880; RPB1, MT606154; TUB,
MT606157.
Distribution: Known only from type location.
Notes: Among the known species of the genus,
P. shennongjiana resembles P. metajoca C.S. Herrera & P.
Chaverri in ellipsoidal, verrucose, and similarly sized asco-
spores, and the acremonium-like conidiophores. However,
P. metajoca differs in having larger perithecia (222–251 ×
204–213 μm vs. 157–186 × 108–147 μm), larger asci
(62.2–69.2 × 5.9–7.2 μmvs.45–65 × 4.5–6μm), and wider
conidia (1.6–3.1 μm vs. 0.9–1.3 μm) (Herrera et al. 2013).
The 4-locus phylogeny indicates the two species are remotely
related (Fig. 1).
Pseudocosmospora sp. (close to P. rogersonii C.S. Herrera
& P. Chaverri, Mycologia 105: 1299, 2013.) Fig. 4
Fig. 3 Pseudocosmospora shennongjiana.a–cPerithecia on natural
substratum. dColony after 1 week at 25°C on PDA. eLongitudinal
section through perithecium in lactophenol cotton blue. fLongitudinal
section through perithecium in water. g–jAsci with ascospores. k–n
Ascospore. o, p Conidiophores and conidia. Scale bars: a–c1mm;e, f
50 μm; g–p10 μm

Fig. 4 Pseudocosmospora sp. a–cPerithecia on natural substratum. d
Perithecia on SNA medium. eColony after 1 week at 25°C on PDA. f
Longitudinal section through perithecium in lactophenol cotton blue. g–i
Asci with ascospores. j–lAscospore. mConidia. n, o Conidiophores and
conidia. Scale bars: a–d1mm;f–o10 μm
424 Mycol Progress (2021) 20:419–429

425Mycol Progress (2021) 20:419–429

Mycelium not visible natural substratum. Perithecia
superficial, solitary to gregarious, non-stromatic,
subpyriform, laterally collapsed upon drying, orange-
red, turning brown in KOH, becoming slightly yellow
in LA, 98–196 × 69–118 μm(n= 8). Perithecial wall
of two layers, 15–28 μm thick; outer layer of textura
globulosa to textura angularis, with smooth surface, 10–
20 μm thick, cells 5–9×4–6μm, walls 0.8–1μmthick;
inner layer of textura prismatica, 5–8μm thick, cells 5–
12 × 2–3.5 μm, walls 0.6–0.8 μmthick.Asci
subcylindrical, 8-spored, with a simple apex, 58–75 ×
5–6μm. Ascospores ellipsoidal with ends slightly taper-
ing, 1-septate, light yellow-brown, smooth, uniseriate and
obliquely overlapping within asci, 7.5–10 × 3–4.5 μm.
Colony on PDA 42 mm in diam. after 1 week at
25°C, pale yellow to cream, surface velvet, with dense
aerial mycelium. Colony on SNA 40 mm diam. after 1
week at 25°C, surface with sparse aerial mycelium.
Conidiophores acremonium- to verticillium-like, arising
from aerial hyphae, septate, branches more or less verti-
cillate, with terminal whorl of 2–5 phialides. Phialides
subulate, tapering towards apex, 12–30 μm long, 0.9–
1.2 μmwideatbase,0.15–0.25 μm wide at tip.
Conidia elliptic-fusoidal, unicellular, smooth, hyaline,
2.5–6×1–1.5 μm.
Specimen examined: China, Hubei, Shennongjia,
Muchengshaoqia, on unidentified ?Xylariaceae, 22
Sept 2014, Z.Q. Zeng, H.D. Zheng, W.T. Qin & K. Chen
10048 (HMAS 271240). Strain, CGMCC 3.20178.
Sequences: ITS, MT592899; nc LSU rDNA, MT592881;
RPB1, MT606155; TUB, MT606158.
Notes: The fungus is most similar to P. rogersonii in having
gregarious perithecia, cylindrical to clavate asci and similarly
sized, ellipsoidal ascospores. However, P. rogersonii occurs on
fruiting bodies of Eutypella sp. other than ?Xylariaceae, and has
wider conidia (1.1–2.6 μmvs.1–1.5 μm) (Herrera et al. 2013).
Additionally, there are 1 bp, 8 bp, and 18 bp divergences in the
nc LSU rDNA, RPB1 and TUB regions between sequences of
CGMCC 3.20178 and GJS 90-56 (ex-holotype culture of
P. rogersonii). The phylogenetic analyses shows that it is close-
ly related to and quite possibly conspecific with MAFF 241531
from Japan (Herrera et al. 2013). We treated the Chinese col-
lection as “Pseudocosmospora sp.”awaiting more collections
to be investigated.
Other Pseudocosmospora species recorded in China
Pseudocosmospora effusa (Teng) Z.Q. Zeng & W.Y. Zhuang,
Mycosystema 36(3): 280, 2017
≡Phaeonectria manilensis var. effusa Teng, Sinensia 7:
497, 1936.
≡Cosmospora effusa (Teng) W.Y. Zhuang & X.M. Zhang,
Nova Hedwigia 74: 279, 2002.
Distribution: China (Zhuang and Zhang 2002).
Pseudocosmospora eutypellae C.S. Herrera & P. Chaverri,
Mycologia 105(5): 1293, 2013
Distribution: France, China and USA (Herrera et al. 2013;
Zeng and Zhuang 2020).
Pseudocosmospora henanensis (Y. Nong & W.Y.
Zhuang) W.Y. Zhuang & Z.Q. Zeng, Mycosystema 36(3):
280, 2017
≡Cosmospora henanensis Y. Nong & W.Y. Zhuang,
Fungal Diversity 19: 96, 2005.
Distribution: China and Japan (Hirooka et al. 2008; Nong
and Zhuang 2005).
Pseudocosmospora joca (Samuels) C.S. Herrera & P.
Chaverri, Mycologia 105: 1296, 2013
≡Nectria joca Samuels, in Samuels, Rossman, Lowen &
Rogerson, Mycol. Pap. 164: 21, 1991.
≡Cosmospora joca (Samuels) Rossman & Samuels, in
Rossman, Samuels, Rogerson & Lowen, Stud. Mycol. 42:
122. 1999.
Distribution: Argentina, Brazil and China (Guu et al. 2007;
Samuels et al. 1991).
Pseudocosmospora nummulariae (Teng) W.Y. Zhuang &
Z.Q. Zeng, Mycosystema 36(3): 280, 2017
≡Nectria nummulariae Teng, Sinensia 4: 274, 1934.
≡Cosmospora nummulariae (Teng) W.Y. Zhuang & X.M.
Zhang, Nova Hedwigia 74: 280, 2002.
≡Dialonectria nummulariae (Teng) Lar.N. Vassiljeva,
Nizshie Rasteniya, Griby i okhoobraznye Dalnego Vostoka
Rossii, Griby. Tom 4. Pirenomitsety i Lokuloaskomitsety
(Sankt-Peterburg): 171, 1998.
Distribution: China (Zhuang and Zhang 2002).
Pseudocosmospora triqua (Samuels) C.S. Herrera & P.
Chaverri, Mycologia 105: 1300, 2013
≡Nectria triqua Samuels, in Samuels, Rossman, Lowen &
Rogerson, Mycol. Pap. 164: 40, 1991.
≡Cosmospora triqua (Samuels) Rossman & Samuels, in
Rossman, Samuels, Rogerson & Lowen, Stud. Mycol. 42:
125, 1999.
Distribution: French Guiana and China (Samuels et al.
1991; Guu et al. 2007; Herrera et al. 2013).
Pseudocosmospora vilior (Starbäck) C.S. Herrera & P.
Chaverri, Mycologia 105: 1301, 2013
≡Nectria vilior Starbäck, Bih. K. svenska VetenskAkad.
Handl., Afd. 3, 25(no. 1): 28, 1899.
426 Mycol Progress (2021) 20:419–429

≡Cosmospora vilior (Starbäck) Rossman & Samuels, in
Rossman, Samuels, Rogerson & Lowen, Stud. Mycol. 42:
126, 1999.
Distribution: Argentina, Brazil, China and Mexico (Teng
1934; Guu et al. 2007; Herrera et al. 2013;Raymundoetal.
2017).
Discussion
For a long time, nectriaceous fungi possessing small, reddish,
smooth, thin-walled perithecia were assigned to Cosmospora
Rabenh. (Rossman et al. 1999). Morphological and phyloge-
netic data suggest that the genus was polyphyletic (Zhang and
Zhuang 2006; Luo and Zhuang 2008,2010b; Samuels et al.
2009) and narrowly defined monophyletic generic concepts
were suggested by Gräfenhan et al. (2011) and Schroers et al.
(2011). Cosmospora vilior and its allies on Eutypa and
Eutypella with acremonium- to verticillium-like asexual states
were moved to Pseudocosmospora (Herrera et al. 2013)com-
prising so far 17 species.
The phylogenetic overview of Pseudocosmospora based
on multilocus sequence analyses showed that the genus is
monophyletic (Herrera et al. 2013). Our work inferred from
sequences of the ITS, nc LSU rDNA, RPB1 and TUB regions
including the newly added taxa had a similar tree topology;
and formed a well-supported monophyletic genus
(MPBP/MLBP/BIPP = 91%/100%/100%) (Fig. 1).
Pseudocosmospora shennongjiana, as a separate lineage, is
grouped with P. rogersonii,P. eutypellae, and the two un-
named species represented by strains of MAFF 241499 and
CGMCC 3.20178 (MPBP/MLBP/BIPP = 94%/100%/100%).
Pseudocosmospora curvispora clustering with another
Pseudocosmospora sp. (GJS 96-216) formed a terminal
branch (MPBP/MLBP/BIPP = 100%/100%/100%).
Although sequence data of some species, such as P. effusa,
P. metepisphaeria (Samuels) C.S. Herrera & P. Chaverri,
P. nummulariae,P. pithoides (Ellis & Everh.) C.S. Herrera
&P.Chaverri,P. pseudepisphaeria (Samuels) C.S. Herrera &
P. Chaverri and P. triqua are not available, P. shennongjiana
and P. curvispora can be easily distinguished from them by
sizes of perithecia, asci, and ascospores (Samuels et al. 1991;
Zhuang and Zhang 2002; Herrera et al. 2013).
Host specificity and substratum preference have been
considered as important taxonomic criteria at generic level
for Cosmospora and cosmospora-like genera. For exam-
ples, Cosmospora is usually found on basidiomycetes
(Fomitopsis P. Karst., Inonotus P. Karst. and Stereum
Hill ex Pers.) and xylariaceous fungi (e.g. Hypoxylon),
while Geejayessia Schroers, Gräfenhan & Seifert and
Volutella Fr. occur on woody substratum (Gräfenhan
et al. 2011; Schroers et al. 2011). Dialonectria (Sacc.)
Cooke, Macroconia (Wollenw.) Gräfenhan, Seifert &
Schroers and Stylonectria Höhn. inhabit ascomata of other
ascomycetes on deciduous trees. However, Microcera
Desm. prefers scale insects (Gräfenhan et al. 2011).
Some Pseudocosmospora spp. favor Diatrypaceae, espe-
cially Eutypa and Eutypella, with the exceptions of
P. joca and P. nummulariae commonly found on
Xylariaceae, and P. hypoxylicola Lechat & J. Fourn on
Hypoxylaceae (Zhuang and Zhang 2002; Herrera et al.
2013; Lechat and Fournier 2020). Many fungal hosts of
Pseudocosmospora are hardly identified to species due to
their decayed condition. Herrera et al. (2015)attemptedto
recognize the host species of Cosmospora through DNA
amplification from ascomata, but most of the obtained
sequences were not the hosts but associated fungi.
Although the known host fungi of Pseudocosmospora
are so far in Xylariales, host specificity has not been
treated as critical for species identification.
Pseudocosmospora is worldwide distributed and expects to
show high species diversity in warm temperate and tropical
regions (Herrera et al. 2013). Among them, P. effuse,
P. hypoxylicola,P. metajoca,P. metepisphaeria,
P. nummulariae,P. pithoides,P. pseudepisphaeria,and
P. rogersonii are probably rare species and only known from
their type locations (Zhuang and Zhang 2002; Herrera et al.
2013; Lechat and Fournier 2020). Since P. nummulariae was
first reported in the 1930s, additional taxa of the genus were
successively discovered from different regions of China. The
country is extremely diverse in its climate, vegetation, geo-
graphic structures, multiple niches, and fungal resources.
Large-scale surveys covering diverse climates, geographic re-
gions, and ecosystems will update our understanding of the
diversity of Pseudocosmospora.
Supplementary Information The online version contains supplementary
material available at https://doi.org/10.1007/s11557-021-01672-1.
Acknowledgements The authors would like to thank the anonymous
reviewers and the editor for their valuable suggestions and linguistic
corrections. We also thank Drs. Huan-Di Zheng, Wen-Tao Qin, and
Kai Chen for collecting specimens for this study.
Author contribution Conceived and designed the experiments: Wen-
Ying Zhuang. Performed the experiments: Zhao-Qing Zeng. Analyzed
the data: Zhao-Qing Zeng and Wen-Ying Zhuang. Wrote the paper:
Zhao-Qing Zeng and Wen-Ying Zhuang.
Funding This work was supported by the National Natural Science
Foundation of China (31750001, 31870012) and Frontier Key Program
of Chinese Academy of Sciences (QYZDY-SSW-SMC029).
Data availability All specimens and strains are deposited in registered
herbaria and culture collection center. Relevant data have been deposited
in GenBank, Fungal Names, and TreeBASE.
427Mycol Progress (2021) 20:419–429

Declarations
Ethics approval and consent to participate Not applicable.
Consent for publication Not applicable.
Conflict of interest The authors declare that they have no conflict of
interest.Supplementary Information The online version contains
supplementary material available at https://doi.org/10.1007/s11557-021-
01672-1.
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