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337
Three new wood-inhabiting fungi of Botryobasidium
(Cantharellales, Basidiomycota) from subtropical forests
of Southwestern China
Lin-Jiang Zhou1,2* , Xue-Long Li3* , Hai-Sheng Yuan1
1 CAS Key Laboratory of Forest Ecology and Silviculture, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110164, China
2 University of the Chinese Academy of Sciences, Beijing 100049, China
3 Institute of Edible Fungi, Liaoning Academy of Agricultural Sciences, Shenyang 110161, Liaoning, China
Corresponding author: Hai-Sheng Yuan (hsyuan@iae.ac.cn)
Copyright: © Lin-Jiang Zhou et al.
This is an open access article distributed under
terms of the Creative Commons Attribution
License (Attribution 4.0 International – CC BY 4.0).
Research Article
Abstract
The basidiomycete genus Botryobasidium is a resupinate saprotrophic with a global dis-
tribution range from coniferous to broad-leaved forest ecosystems. Though numerous
species have been reported from Eurasia and North America, few have been described
from China. In the current work, phylogenetic analyses of Botryobasidium in China were
conducted based on the dataset of the internal transcribed spacer (ITS) regions and the
large subunit (LSU) of nuclear ribosomal RNA gene. Maximum likelihood and Bayesian
analyses were used to reconstruct the phylogenetic tree, and three new species, namely
Botryobasidium acanthosporum, B. leptocystidiatum and B. subovalibasidium, were de-
scribed from subtropical forests of Yunnan Province, Southwestern China. Botryobasidi-
um acanthosporum is characterized by having yellowish white to dark yellow basidiome,
clavate to tubular cystidia, and subglobose to globose basidiospores with obtuse spines.
Botryobasidium leptocystidiatum
white to ivory basidiome, generative hyphae with clamped, tubular cystidia, and subna-
vicular to navicular basidiospores. While, B. subovalibasidium is characterized by having
yellowish to ivory basidiome, subovoid basidia, navicular to suburniform basidiospores,
and thick-walled chlamydospores. These three new species are described and illustrat-
ed, and the discriminating characters between the new species and their closely related
species are discussed. A key to known species of Botryobasidium in China is provided.
Key words: Botryobasidiaceae, corticioid fungi, subtropical forests, taxonomy, wood-de-
caying fungi
Introduction
Botryobasidium Donk belongs to the order Cantharellales of phylum Basidio-
B. subcoronatum (Höhn. & Litsch.) Donk (Moncal-
vo et al. 2006). Many asexual morph generic names, such as Acladium Link,
Allescheriella Henn., Alysidium Kunze, Haplotrichum Link, Neoacladium P.N.
Singh & S.K. Singh, Physospora Fr., and Sporocephalium Chevall., are conge-
neric with Botryobasidium, and were re-combined in Botryobasidium (Stalpers
Academic editor:
Danushka Sandaruwan Tennakoon
Received:
29 July 2024
Accepted:
20 September 2024
Published:
22 October 2024
Citation: Zhou L-J, Li X-L, Yuan
H-S (2024) Three new wood-
inhabiting fungi of Botryobasidium
(Cantharellales, Basidiomycota)
from subtropical forests of
Southwestern China. MycoKeys 109:
337–354. https://doi.org/10.3897/
mycokeys.109.133325
MycoKeys 109: 337–354 (2024)
DOI: 10.3897/mycokeys.109.133325
This article is part of:
Exploring the Hidden Fungal Diversity:
Biodiversity, Taxonomy, and Phylogeny of
Saprobic Fungi
Edited by Samantha C. Karunarathna,
Danushka Sandaruwan Tennakoon,
Ajay Kumar Gautam
* These authors contributed equally to this work.
338
MycoKeys 109: 337–354 (2024), DOI: 10.3897/mycokeys.109.133325
Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
et al. 2021). The genus is characterized by resupinate, smooth, arachnoid, hy-
pochnoid, pellicular or grandinioid basidiomes, a monomitic hyphal system
with simple septate or nodose generative hyphae, clavate to cylindrical cystidia,
claviform to suburniform basidia with 2–8 sterigmata, and navicular to globose,
thin- or thick-walled, smooth or ornamented basidiospores (Binder et al. 2005;
Larsson 2007; Buyck et al. 2017; Bondartseva and Zmitrovich 2023). In macro-
morphology, species of Botryobasidium are easily confused with some genera,
e.g., Ceratobasidium D.P. Rogers, Sistotrema Fr., and Tulasnella J. Schröt. in the
Cantharellales (Donk 1956; Oberwinkler 1982), whereas Botryobasidium differs
from the others in absence of epibasidia, sturdy and long sterigmata, and oily
inclusions (Kotiranta and Saarenoksa 2005; Gorjón and Hallenberg 2008; Ober-
close relationships among Botryobasidium, Cantharellus, Clavulina, Hydnum
and Tulasnella (Jülich 1981; Hibbett et al. 1997; Bruns et al. 1998; Langer 1998;
Pine et al. 1999; Cao et al. 2021).
The species of Botryobasidium are a group of saprotrophic fungi that cause
a white rot in fallen angiosperm and gymnosperm woods, which play a key role
al. 2000b; Bondartseva and Zmitrovich 2023). They can be commonly found on
various hosts or substrates from the litter, fallen trunk to stem of living trees,
including the macrophanerophytes, such as Abies Mill, Acer Linn., Alnus Mill.,
Betula L., Citrus L., Corylus L., Eucalyptus L. Herit, Fagus L., Magnolia Linn.,
Persea Mill., Picea Dietr., Pinus Linn, Populus L., Quercus L., Salix L. and Tsuga
Carr.; the shrubs, such as Bambusa Retz. corr. Schreber and Pandanus Linn.
f.; the pteridophytes, such as Pteris L. and Cibotium Kaulf. (Anon 1969; Hol-
ubová-Jechová 1969; Boidin and Gilles 1982; Langer 1994; Langer et al. 2000a,
2000b; Hjortstam et al. 2005). Additionally, some species also develop on ma-
ture basidiomes of Irpex lacteus (Boidin and Gilles 1982), as well as on soil and
underground timber (Anon 1969).
Up to now, the genus of about 84 species have been accepted globally in
Index Fungorum and MycoBank (Lentz 1967; Jung 1995; Langer et al. 2000a;
Hagara 2001; Bernicchia et al. 2010; Saitta et al. 2011; Bates et al. 2017; Ka-
linina et al. 2020; Ram et al. 2021). There have been recorded about 35 spe-
cies in Europe, 28 in North America, 26 in Latin America, 25 in Africa, 20 in
Oceania, and 23 in Asia (Dritter 1809; Anon 1969; Holubová-Jechová 1969;
Pouzar and Holubová-Jechová 1969; Holubová-Jechová 1980; Boidin and
Gilles 1982; Boidin and Gilles 1988; Langer 1994; Greslebin and Rajchenberg
2003; Parmasto et al. 2004; Hjortstam et al. 2005; Bates et al. 2017; Buyck et
al. 2017; Hyde et al. 2019; Vondrák et al. 2023). So far, 15 species of Botryo-
basidium have been reported from China, and most of them were distributed
in the north temperate to subtropical zones (Dai 2011; Liu et al. 2024; Zhou
et al. 2024).
During the surveys of lignicolous fungi in Yunnan Province, Southwestern Chi-
na, several Botryobasidium specimens were collected from the mixed forests.
The subsequent research by morphology and molecular phylogeny indicates
that these specimens represent several undescribed species. The phylogenetic
positions and the relationships of these species among Botryobasidium were
line drawings were provided in this study.
339
MycoKeys 109: 337–354 (2024), DOI: 10.3897/mycokeys.109.133325
Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
Materials and methods
Morphological study
The voucher specimens are deposited in the herbarium of the Institute of Ap-
plied Ecology, Chinese Academy of Sciences (IFP). Macromorphological char-
acteristics were examined using a Nikon SMZ 645 (Tokyo, Japan) stereo mi-
croscope and the color descriptions refer to Kornerup and Wanscher (1981).
Microscopical structures were checked using hand-cut sections stained with
Cotton blue, Melzer’s reagent, and 3% KOH, and line drawings were prepared
using a Nikon Eclipse 80i microscope (Nikon Corporation, Japan) with the
aid of a drawing tube. The surface morphology for the basidiospores was
CIQTEK Co., Ltd.) at an accelerating voltage of 3 kV. The working distance
was 9.62 mm. A thin layer of gold was plated on the sample to enhance the
conductivity. Basidiospores were measured based on the front and back side
view; the apex was excluded from the spore measurements. The following
abbreviations are used: L = mean spore length, W = mean spore width, Q = L/W
ratio, n (a/b) = number of spores (a) measured from number of specimens (b).
Potassium hydroxide solution (KOH) was used to detect changes in hyphae,
gloeocystidia, and encrusted. Melzer’s reagent (IKI) was used to determine
amyloidity and dextrinoidity.
DNA extraction, PCR amplication, and DNA sequencing
According to the manufacturer’s instructions, the Fungal Fast Non-Toxic DNA
Extraction Kit (Demeter Biotech Co., Ltd, Beijing, China) was employed to ex-
primers ITS1 and ITS4 (White et al. 1990), and the procedure was an initial
denaturation at 95 °C for 3 min, followed by 34 cycles at 95 °C for 30 s, 58 °C
LR0R and LR7 (Vilgalys and Hester 1990), and the procedure involved an initial
denaturation at 95 °C for 3 min, followed by 34 cycles at 95 °C for 30 s, 50 °C
for 30 s, and 72 °C for 1 min, the procedure ended with an extension at 72 °C
for 5 min.
DNA sequencing was conducted at the Beijing Genomics Institute (BGI), and
the sequences were assembled using Geneious v.9.0.2 (Kearse et al. 2012).
and integrity, and uploaded to GenBank (Table 1).
Phylogenetic analyses
Suillosporium cystidiatum (D.P. Rogers) Pouzar (Botryobasidiaceae) was cho-
sen as the outgroup according to the result of sequence BLAST in NCBI data-
base, ensuring that it has suitable phylogenetic distances from other species
in Botryobasidium. The concatenated datasets of ITS and LSU sequences of
the species in Botryobasidiaceae were used to infer the molecular phylogeny.
340
MycoKeys 109: 337–354 (2024), DOI: 10.3897/mycokeys.109.133325
Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
Table 1. Species and GenBank numbers used in phylogenetic analysis in this study.
Species name ITS LSU Specimen No. Substrate Country References
Botryobasidium acanthosporum
L.J. Zhou & H.S. Yuan
PP229497 / Yuan16326 on fallen angiosperm branch China Present study
B. acanthosporum PP229511 / Yuan17989 on bark of angiosperm China Present study
B. acanthosporum PP229512 PP218361 Yuan18083 on fallen trunk of Abies China Present study
B. acanthosporum PP229517 / Yuan18128 on fallen trunk of Abies China Present study
Botryobasidium aureum
Parmasto
AJ389783 / GEL 2910 / Germany Langer et al. 2000b
B. botryosum (Bres.) J. Erikss. DQ267124 DQ089013 AFTOL-ID 604 / USA AFTOL Database
B. candicans J. Erikss. KP814200 / UC2022893 on litter or well decayed wood in
pinaceous forest
USA Rosenthal et al.
2017
B. cf. subcoronatum KP814216 / UC2022856 on litter or well decayed wood in
pinaceous forest
USA Rosenthal et al.
2017
B. cf. subcoronatum KP814322 / UC2022917 on litter or well decayed wood in
pinaceous forest
USA Rosenthal et al.
2017
B. coniferarum S.L. Liu & L.W.
Zhou
PP209210 PP218367 Yuan18440 on fallen gymnosperm trunk China Present study
B. coniferarum OR557262 OR527286 LWZ20171016-15 on fallen branch of Pinus China Liu et al. 2024
B. coniferarum OR557259 OR527282 LWZ20210928-3 on fallen branch of Pinus China Liu et al. 2024
B. conspersum J. Erikss. DQ911612 DQ521414 PBM 2747 (CUW) / USA AFTOL Database
B. conspersum OP163274 / FLAS-F-69114 / USA NCBI Database
B. conspersum /AY586657 GB/KHL11063 / Sweden Larsson et al. 2004
B. curtisii (Berk.) Hol.-Jech. EU118629 EU118629 KHL 12950GB / Costa
Rica
Larsson 2007
B. gossypirubiginosum Q. Zhou
& C.L. Zhao
OR668924 OR708665 CLZhao 26052 on fallen angiosperm branch China Zhou et al. 2024
B. incanum Q. Zhou & C.L. Zhao OR668923 OR708664 CLZhao 26697 on fallen angiosperm branch China Zhou et al. 2024
B. incanum PP209201 PP218357 Yuan17803 on fallen angiosperm branch China Present study
B. indicum (P.N. Singh & S.K.
Singh) R. Kirschner & G. Langer
PP209209 PP218363 Yuan18250 on root of Quercus China Present study
B. indicum ON406471 / CLZhao 21791 / China NCBI Database
B. indicum NR171230 NG070816 AMH:10054 dead bark of Leucaena leucocephala India Hyde et al. 2019
B. indicum MK391496 MK391493 AMH:10054 dead bark of Leucaena leucocephala India Hyde et al. 2019
B. intertextum (Schwein.) Jülich
& Stalpers
KP814540 / UC2022959 on litter or well decayed wood in
pinaceous forest
USA Rosenthal et al.
2017
B. intertextum AJ389782 / DAOM 197881 / Canada Langer et al. 2000b
B. isabellinum (Fr.) D.P. Rogers MZ159478 / K(M):181602 / UK NCBI Database
B. leptocystidiatum L.J. Zhou &
H.S. Yuan
PP209211 PP218178 Yuan17548 on fallen branch of Pinus China Present study
B. leptocystidiatum PP204173 PP218180 Yuan17557 on dead tree of Pinus China Present study
B. leptocystidiatum PP209200 PP218353 Yuan17706 on fallen angiosperm trunk China Present study
B. leptocystidiatum PP209197 PP218354 Yuan17708 on bark of living angiosperm tree China Present study
B. leptocystidiatum PP209198 PP218355 Yuan17709 on fallen angiosperm trunk China Present study
B. robustius Pouzar & Hol.-Jech. MH859491 MH871272 CBS:945.69 / Czech Vu et al. 2019
B. robustius PP436446 / HAY-F-004374 / USA NCBI Database
B. subcoronatum (Höhn. &
Litsch.) Donk
EU118607 EU118607 KHL s.n. (GB) / Sweden Larsson 2007
B. subcoronatum MH211720 FLAS-F-61064 / USA NCBI Database
B. subcoronatum DQ200924 AY647212 AFTOL-ID 614 / USA Matheny et al. 2007
B. subovalibasidium. L.J. Zhou &
H.S. Yuan
PP209199 PP218152 Yuan16439 on fallen trunk of Hippophae
rhamnoides
China Present study
B. subovalibasidium PP209196 PP218362 Yuan18179 on fallen trunk of Abies China Present study
B. tubulicystidium G. Langer OL436769 / DK14_139 / USA NCBI Database
B. vagum (Berk. & M.A. Curtis)
D.P. Rogers
OR680661 / personal:Alden
Dirks:ACD0672
/ USA Zhou et al. 2024
B. vagum OR471092 / TENN:075258 on Pinus USA Zhou et al. 2024
B. yunnanense Q. Zhou & C.L.
Zhao
OR668925 OR708666 CLZhao 24877 on fallen angiosperm branch China Zhou et al. 2024
Suillosporium cystidiatum (D.P.
Rogers) Pouzar
MN937573 MN937573 VS3830 On Picea jezoensis var. jezoensis Russia NCBI Database
341
MycoKeys 109: 337–354 (2024), DOI: 10.3897/mycokeys.109.133325
Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
The ITS and LSU sequences were aligned separately using MEGA v.7.0 (Kumar et al. 2016).
Maximum likelihood (ML) analysis was done using RAxML v.1.5b2 (Silvestro
and Michalak 2012) with non-parametric bootstrapping of 500 replicates un-
der the GTRGAMMA model. A Bayesian inference (BI) was also performed for
the same data sets using MrBayes 3.2.6 (Ronquist et al. 2012). A substitution
model was selected in PhyloSuite v1.2.2 (Zhang et al. 2020). The Bayesian
information criterion (BIC) values under each model were compared and the
model with the lowest BIC value was selected. Two parallel analyses were then
run in MrBayes for 2 million generations, with 4 chains each, sampling every
500 generations. Burn-in trees (initial 25%) were discarded for each run and
posterior probabilities of the matrix were determined by calculating a major-
ity-rule consensus tree generated from the post-burnin trees by the MCMC
runs using the sump of MrBayes. The phylogenetic trees were visualized using
FigTree v1.4.3 (Rambaut 2016). Branches that received bootstrap support for
-
ed, respectively. The datasets were deposited in TreeBASE (www.treebase.org/
treebase-web/, study no. 31569).
Results
Phylogeny
The ITS dataset consists of 39 sequences representing 20 taxa of Botryoba-
sidium, and a sample of Suillosporium cystidiatum as the outgroup. The ITS se-
quence had an aligned length of 661 base pairs (bp), of which 321 were parsimo-
ny-informative, 75 were singleton sites, 265 were constant sites. The Bayesian
analysis had an average standard deviation of split frequencies = 0.004148,
and a 50% majority-rule consensus phylogram was generated. The best model
-
-
resenting 20 taxa of Botryobasidium, and a sample of Suillosporium cystidiatum
-
cluding 663 bp of ITS and 839 bp of LSU), of which 434 were parsimony-infor-
mative, 158 were singleton sites, 910 were constant sites. The Bayesian anal-
ysis had an average standard deviation of split frequencies = 0.005929, and a
50% majority-rule consensus phylogram was generated. The best model was
-
freqpr = dirichlet (1, 1, 1, 1)].
In the phylogenetic tree based on ITS dataset (Fig. 1), four specimens of
B. acanthosporum formed a clade (ML 100%/BPP 1), and grouped with B. in-
canum, B. vagum, and B. isabellinum with full support (ML 100%/BPP 1). Two
specimens of B. leptocystidiatum formed a clade with full support (ML 100%/
BPP 1). The remaining two specimens of B. subovalibasidium formed a clade,
and clustered with B. aureum, B. botryosum and B. candicans with strong sup-
port (ML 95%/BPP 1).
to which two new species belong swapped positions, but the taxonomic posi-
tions of these three new species and the relationships with their sibling species
are no discrepancy. Moreover, the support of the branches to which two new
342
MycoKeys 109: 337–354 (2024), DOI: 10.3897/mycokeys.109.133325
Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
Figure 1. Phylogram of Botryobasidium resulting from a maximum likelihood analysis based on ITS sequence. Maximum
nodes. New species are in yellow background.
Figure 2. Phylogram of Botryobasidium-
New species are in yellow background.
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Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
species belong, B. acanthosporum and B. subovalibasidium, was strengthened
(ML 98%/BPP 1). Thus, the phylogenetic analyses revealed the taxonomic posi-
tions of these three new species.
Taxonomy
Botryobasidium acanthosporum L.J. Zhou & H.S. Yuan, sp. nov.
Fungal Names: FN 572031
Figs 3A, B, 4, 5
Diagnosis. Differed from other Botryobasidium species in having arachnoid ba-
sidiome with attached granules, clavate to subcylindrical cystidia, and subglo-
bose to globose basidiospores with blunt spines up to 4 µm long.
Type. • Yunnan Province, Diqing Prefecture, Pudacuo National Park,
27°53'54"N, 99°57'04"E, on fallen trunk of Abies, 14 August 2023, Yuan 18083
(IFP 19972).
Etymology. acanthosporum (Lat.), referring to the spore with spines.
Description. Basidiomes:
and separates easily when wet. Hymenophoral surface smooth, greyish white
to yellowish white (1B2–4B2) when fresh, pale yellow to dark yellow (3A3–4C8)
when dry. Sterile margin often indeterminate and not differentiated.
Hyphal structure: hyphal system monomitic; generative hyphae simple sep-
tate, thin- to slightly thick-walled; tissues unchanged in KOH.
Subiculum: subicular hyphae colorless, thick-walled, frequently branched at
-
Cystidia: clavate to tubular, infrequent, smooth, thin-walled, colorless, simple
septate, apically obtuse, acyanophilous, inamyloid, unchanged in KOH and dis-
Basidia: clavate to subcylindrical, smooth, thin-walled, with 2 sterigmata,
simple septate, acyanophilous, inamyloid, unchanged in KOH and distilled wa-
Basidiospores: subglobose to globose, aculeate, slightly thick- to thick-
walled, colorless, cyanophilous, inamyloid, unchanged in KOH and distilled
Q = 1.0–1.13 (n = 60/2); spines with apically obtuse, usually isolated, some-
times grouped in 2, up to 4 µm long.
Chlamydospores absent and anamorph not seen.
Ecology and distribution. Growing in mixed forests dominated by Abies and
a small number of Picea, Quercus, and other angiosperm trees. So far, known
from Yunnan Province and Xizang Autonomous Region, China.
Additional specimens examined. • Xizang Autonomous Region, Bomi
County, Yigong Tea Farm, 30°07'55"N, 95°01'05"E, on fallen angiosperm branch,
24 October 2021, Yuan 16326 (IFP 19970; paratype) • Yunnan Province, Diqing
Prefecture, Baimaxueshan National Nature Reserve, 28°18'19"N, 99°08'57"E, on
bark of angiosperm, 13 August 2023, Yuan 17989 (IFP 19971) • Pudacuo Na-
tional Park, 27°53'56"N, 99°57'16"E, on fallen trunk of Abies, 14 August 2023,
Yuan 18128 (holotype IFP 19973).
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Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
Botryobasidium leptocystidiatum L.J. Zhou & H.S. Yuan, sp. nov.
Fungal Names: FN 571970
Figs 3C, D, 6
Diagnosis. Differed from other Botryobasidium species in having tubular cystid-
ia and clamped in all hyphae.
Type. • Yunnan Province, Lincang City, Wulaoshan National Forest
Park, 23°54'47"N, 100°10'53"E, on bark of living angiosperm tree, 9 August
2023, Yuan 17708 (holotype IFP 019955).
Etymology. leptocystidiatum (Lat.), referring to the leptocystidia.
Description. Basidiomes:
-
ed. Hymenophoral surface smooth, greyish white (1B1–30B1) to smoky grey
(3C2) when fresh, greyish white (1B1–30B1) to ivory (4B3) when dry; margin
often indeterminate and not differentiated.
Figure 3. The habitats and basidiomes of three new species of Botryobasidium A, B B. acanthosporum (holotype Yuan
18083) C, D B. leptocystidiatum (holotype Yuan 17708) E, F B. subovalibasidium (holotype Yuan 18179).
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MycoKeys 109: 337–354 (2024), DOI: 10.3897/mycokeys.109.133325
Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
Hyphal structure: hyphal system monomitic; generative hyphae clamped,
thin- to slightly thick-walled; tissues unchanged in KOH.
Subiculum: subicular hyphae colorless, slightly thick-walled, sparsely
diam. Subhymenial hyphae colorless, thin-walled, frequently branched at right
Cystidia: tubular, infrequent, smooth, thin-walled, colorless, apically ob-
tuse, basal clamped, without additional septate, acyanophilous, inamyloid, un-
Basidia: ordered by botryose cluster, subcylindrical, smooth, thin-walled, usu-
ally with 6 sterigmata, occasionally with 7 sterigmata, basal clamped, acyano-
Basidiospores: subnavicular to navicular, smooth, thin-walled, colorless, oc-
casionally a few stuck together, acyanophilous, inamyloid, unchanged in KOH
Chlamydospores absent and anamorph not seen.
Ecology and distribution. Growing in mixed forests dominated by Pinus and a
small number of Fagaceae trees. So far only known from Yunnan Province, China.
Figure 4. Microscopic features of Botryobasidium acanthosporum (drawn from holotype Yuan 18083) A a section through
basidiome B basidiospores C basidia D basidioles E
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Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
Figure 6. Microscopic features of Botryobasidium leptocystidiatum (drawn from holotype Yuan 17708) A a section
through basidiome B basidiospores C basidioles D basidia E
Figure 5. SEM of basidiospores of Botryobasidium acanthosporum species (holotype Yuan 18083).
Additional specimens examined. • Yunnan Province, Lincang City, Wu-
laoshan National Forest Park, 23°54'47"N, 100°10'53"E, on fallen branch of Pi-
nus, 8 August 2023, Yuan 17548 (IFP 019952; paratype) • on dead tree of Pinus,
8 August 2023, Yuan 17557 (IFP 019953) • on fallen angiosperm trunk, 9 August
2023, Yuan 17706 (IFP 019954), Yuan 17709 (IFP 019956).
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MycoKeys 109: 337–354 (2024), DOI: 10.3897/mycokeys.109.133325
Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
Botryobasidium subovalibasidium L.J. Zhou & H.S. Yuan, sp. nov.
Fungal Names: FN 571974
Figs 3E, F, 7
Diagnosis. Differed from other Botryobasidium species in having effuse, yellow-
ish to ivory basidiomes, subovoid to ovoid basidia, ellipsoid chlamydospores.
Type. • Yunnan Province, Diqing Prefecture, Pudacuo National Park,
27°83'67"N, 99°95'76"E, Alt. 3655 m, on fallen trunk of Abies, 15 August 2023,
Yuan 18179 (holotype IFP 019957).
Etymology. subovalibasidium (Lat.), referring to the subovoid basidia.
Description. Basidiomes:
thick, adherent to the substrate and not easily separated. Hymenophoral sur-
face smooth, greyish white (1B1–30B1) to ivory (4B3) when fresh, pale yellow
(4A3) to greyish yellow (4B5) when dry; margin not differentiated, distinct.
Hyphal structure: hyphal system monomitic; generative hyphae simple sep-
tate, thin- to slightly thick-walled; tissues unchanged in KOH.
Subiculum: subicular hyphae colorless, slightly thick-walled, frequently
Subhymenial hyphae colorless, thin-walled, moderately branched, acyanophi-
Cystidia: absent.
Basidia: subovoid to ovoid, smooth, thin-walled, with 4–6 sterigmata, basal
simple septate, acyanophilous, inamyloid, unchanged in KOH and distilled wa-
ter, (12–)14–18 × 9–10 µm.
Basidiospores: navicular to suburniform, smooth, thin-walled, colorless, oc-
casionally stuck together, acyanophilous, inamyloid, unchanged in KOH and dis-
Figure 7. Microscopic features of Botryobasidium subovalibasidium (drawn from holotype Yuan 18179) A a section through
basidiome B basidiospores C basidioles D basidia E secondary spores F chlamydospores G
348
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Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
tilled water, (5.7–)7–9.8(–10) × (3.2–)3.7–5(–5.1) µm, L = 8.3 µm, W = 4.2 µm,
Q = 1.53–2.5 (n = 60/1).
Chlamydospores: orange-yellow, ellipsoid, smooth, thick-walled, cyanophi-
lous, inamyloid, unchanged in KOH, unchanged in distilled water, 17–21(–22) ×
(9–)10–11 µm, L = 18.5 µm, W = 10.3 µm, Q = 1.50–2.1 (n = 60/2).
Ecology and distribution. Growing in mixed forests dominated by Abies and
a small number of Picea, Quercus, and other angiosperm trees. So far, known
from Yunnan Province and Xizang Autonomous Region, China.
Additional specimen examined. • Xizang Autonomous Region, Bomi
County, on fallen trunk of Hippophae rhamnoides, 26 October 2021, Yuan 16439
(IFP 019951; paratype).
Discussion
In this study, three new species of Botryobasidium collected from Southwestern
China are described based on morphological characteristics and phylogenetic
analyses combining ITS and LSU sequences. The molecular phylogenetic analy-
ses showed moderate to high support in the deeper nodes and at the species lev-
el which is consistent with the previous study (Cao et al. 2021; Zhou et al. 2024).
The phylogenetic trees show that B. acanthosporum is closely linked to B. inca-
num, B. isabellinum and B. vagum (Figs 1, 2). B. acanthosporum resembles B. inca-
num and B. vagum in having pellicular and greyish basidiomes. However, the new
species is unique by having spine-ornamented basidiospores. B. acanthosporum
is similar to B. isabellinum in having yellowish basidiomes and spine-ornamented
basidiospores, but B. isabellinum differs from the new species by having narrower
In morphology, B. bondarcevii resembles B. acanthosporum in having pellicular
and slightly yellow to dark yellow basidiomes, and spine-ornamented basidio-
spores, but B. bondarcevii can be distinguished by having bigger basidia (18–23
In the phylogenetic trees (Figs 1, 2), B. leptocystidiatum grouped together with
B. subcoronatum. Morphologically, they share similar characteristics in having
thin and whitish to pale yellow basidiomes, clamped generative hyphae with
frequently vertical branches, and subnavicular basidiospores. Nevertheless,
B. subcoronatum differs from B. leptocystidiatum by the absence of cystidia,
longer basidia and slightly narrower basidiospores (6–8 × 2.5–3 µm vs. 6.5–
7.8 × 2.9–3.7 µm). Moreover, B. leptocystidiatum and B. sassofratinoense are
similar in having greyish-white to yellow basidiomes and clamped hyphae. But
B. sassofratinoense can be differentiated by having wider subhymenial hyphae
Botryobasidium subovalibasidium has an adjacent phylogenetic relationship
with B. aureum, B. botryosum and B. candicans in the phylogenetic trees (Figs 1,
2). In morphology, they exhibit some similarities in having whitish to yellowish
basidiomes and absence of cystidia (Breitenbach and Kränzlin 1986; Bernic-
chia and Gorjón 2010). However, B. aureum is distinguished from the new spe-
349
MycoKeys 109: 337–354 (2024), DOI: 10.3897/mycokeys.109.133325
Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
cies by having narrower subhymenial hyphae (4–6 µm) and subicular hyphae
(5–10 µm) and smaller basidia (12–18 × 6–8 µm). Botryobasidium candicans
differs from the new species by having narrower subhymenium and subicular
hyphae, slightly smaller basidiospores (6–8 × 3–4 µm vs. 7–9.8 × 3.7–5.0 µm),
and smaller chlamydospores (15–17 × 9–12 µm vs. 17–21 × 10–11 µm) (Ber-
nicchia and Gorjón 2010). Botryobasidium botryosum can be separated from
vs. 7–9.8 × 3.7–5.0 µm), and absence of conidiospores (Jülich 1978). More-
over, B. subovalibasidium and B. danicum are similar in having greyish to yellow-
ish basidiomes, absence of cystidia, and basidia with 4–6 sterigmata. However,
B. danicum is distinct from the new species by longer basidiospores (12–14 ×
3.0–5.0 µm vs. 7–9.8 × 3.7–5.0 µm), bigger basidia (15–20 × 8–12 µm vs. 14–18
× 9–10 µm) and absence of anamorphic spores (Bernicchia and Gorjón 2010).
Key to known 18 species of Botryobasidium in China
1 Basidiospores with spines ............................................................................ 2
– Basidiospores smooth .................................................................................. 4
2 Basidiospores ellipsoid, 7–9 × 5–6.3 µm .............................. B. bondarcevii
– Basidiospores globose .................................................................................3
3 Basidiospores 7–10 µm, spines up to 1–3 µm, basidia with 4 sterigmata .
.................................................................................................. B. isabellinum
– Basidiospores 8–10 µm, spines up to 4 µm, basidia with 2 sterigmata ......
........................................................................................... B. acanthosporum
4 Conidia absent ............................................................................................... 5
– Conidia present ...........................................................................................15
5 Hyphae with clamps at least in a part of basidiome ...................................6
– Hyphae without clamps ................................................................................9
6 Clamps present on all septa ......................................................................... 7
– Both clamps and simple septa present .......................................................8
7 Basidiospores navicular, 6–7 × 2.5–3 µm; cystidia absent ... B. subcoronatum
– Basidiospores subnavicular to navicular, 6.5–7.8 × 2.9–3.7 µm; cystidia
present .............................................................................B. leptocystidiatum
8 Clamps often present in subiculum and subhymenium ..... B. angustisporum
– Clamps often absent in subiculum ........................................ B. intertextum
9 Basidiospores navicular ..............................................................................10
– Basidiospores not navicular .......................................................................12
10 Basidiospores 7–8 × 3–3.5 µm ............................................. B. coniferarum
– Basidiospores more than 8 µm long ..........................................................11
11 Basidiospores 9–10 × 3.5–5 µm; basidia cylindrical, 9–16 × 7–9 µm ........
............................................................................................. B. subbotryosum
– Basidiospores 8–12 × 4.5–6 µm; basidia clavate to subcylindrical, 20–25
× 8–12 µm ........................................................................................ B. vagum
12 Basidiospores obliquely ovoid, apically obtuse .........................................13
– Basidiospores not ovoid .............................................................................14
13 Basidiospores 7.5–12 × 3.5–5 µm ..................................... B. obtusisporum
– Basidiospores 5–8 × 2.5–3.5 µm ............................................. B. pruinatum
14 Basidiospores subglobose, 14–17.5 × 13–15.5 µm ....B. gossypirubiginosum
– Basidiospores ellipsoid, 6.5–8.5 × 3.5–5 µm ............................. B. incanum
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Lin-Jiang Zhou et al.: Three new Botryobasidium from Southwestern China
15 Conidia ellipsoid .......................................................................................... 16
– Conidia not ellipsoid ....................................................................................17
16 Conidia 13–22 × 9–12 µm; basidia ellipsoid to obovate, 12–15 × 6–8 µm
................................................................................................. B. conspersum
– Conidia 17–21 × 10–11 µm; basidia subovoid to ovoid, 14–18 × 9–10 µm
........................................................................................ B. subovalibasidium
17 Conidia subglobose to citriform, 15–20 × 8–10 µm ................B. candicans
– Conidia subglobose to globose, 11.5–14.5 × 9.5–10.5 µm .... B. yunnanense
Additional information
Conict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
Funding
This research was supported by the National Natural Science Foundation of China (Proj-
ect Nos. U2102220 & 31970017) and the CAS Key Laboratory Annual Project.
Author contributions
Investigation and writing draft: LJZ. Data measurement and analysis: XLL. Conceptu-
alization and supervision: HSY. All authors contributed to the article and approved the
submitted version.
Author ORCIDs
Lin-Jiang Zhou https://orcid.org/0000-0003-2665-6959
Xue-Long Li https://orcid.org/0009-0006-3948-0234
Hai-Sheng Yuan https://orcid.org/0000-0001-7056-140X
Data availability
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