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fmicb-13-818358 April 18, 2022 Time: 13:41 # 1
ORIGINAL RESEARCH
published: 25 April 2022
doi: 10.3389/fmicb.2022.818358
Edited by:
Qi Zhao,
University of Science and Technology
Liaoning, China
Reviewed by:
Angelina De Meiras-Ottoni,
Federal University of Pernambuco,
Brazil
Renata Dos Santos Chikowski,
Federal University of Pernambuco,
Brazil
Alexander Ordynets,
University of Kassel, Germany
*Correspondence:
Yuan Yuan
yuanyuan1018@bjfu.edu.cn
Specialty section:
This article was submitted to
Systems Microbiology,
a section of the journal
Frontiers in Microbiology
Received: 19 November 2021
Accepted: 28 February 2022
Published: 25 April 2022
Citation:
Liu Z-B, Wu Y-D, Zhao H,
Lian Y-P, Wang Y-R, Wang C-G,
Mao W-L and Yuan Y (2022) Outline,
Divergence Times, and Phylogenetic
Analyses of Trechisporales
(Agaricomycetes, Basidiomycota).
Front. Microbiol. 13:818358.
doi: 10.3389/fmicb.2022.818358
Outline, Divergence Times, and
Phylogenetic Analyses of
Trechisporales (Agaricomycetes,
Basidiomycota)
Zhan-Bo Liu1, Ying-Da Wu1,2 , Heng Zhao1, Ya-Ping Lian1, Ya-Rong Wang1,
Chao-Ge Wang1, Wei-Lin Mao1and Yuan Yuan1*
1School of Ecology and Nature Conservation, Beijing Forestry University, Beijing, China, 2Key Laboratory of Forest and
Grassland Fire Risk Prevention, Ministry of Emergency Management, China Fire and Rescue Institute, Beijing, China
Phylogenetic analyses inferred from the nuc rDNA ITS1-5.8S-ITS2 (ITS) data set and
the combined 2-locus data set [5.8S +nuc 28S rDNA (nLSU)] of taxa of Trechisporales
around the world show that Sistotremastrum family forms a monophyletic lineage within
Trechisporales. Bayesian evolutionary and divergence time analyses on two data sets
of 5.8S and nLSU sequences indicate an ancient divergence of Sistotremastrum family
from Hydnodontaceae during the Triassic period (224.25 Mya). Sistotremastrum family
is characterized by resupinate and thin basidiomata, smooth, verruculose, or odontoid-
semiporoid hymenophore, a monomitic hyphal structure, and generative hyphae bearing
clamp connections, the presence of cystidia and hyphidia in some species, thin-walled,
smooth, inamyloid, and acyanophilous basidiospores. In addition, four new species,
namely, Trechispora dentata,Trechispora dimitiella,Trechispora fragilis, and Trechispora
laevispora, are described and illustrated. In addition, three new combinations, namely,
Brevicellicium daweishanense,Brevicellicium xanthum, and Sertulicium limonadense,
are also proposed.
Keywords: Hydnodontaceae, phylogenetic analysis, Trechispora, taxonomy, wood-rotting fungi
INTRODUCTION
Trechisporales K.H. Larss. was established by Hibbett et al. (2007). Most species in this order are
corticioid fungi with smooth, grandinioid, odontioid, or hydnoid hymenophores, and others are
polypores. All species have a monomitic or dimitic hyphal system with generative hyphae bearing
clamp connections, and many species have rhizomorphs (mycelial cords) (Larsson, 2007).
At present, there is only an acknowledged and a named family belonging to Trechisporales,
i.e., Hydnodontaceae Jülich. Hydnodontaceae contains 11 genera now, namely, Brevicellicium K.H.
Larss. and Hjortstam, Dextrinocystis Gilb. and M. Blackw., Fibrodontia Parmasto, Pteridomyces
Jülich, Luellia K.H. Larss. and Hjortstam, Porpomyces Jülich, Scytinopogon Singer, Subulicystidium
Parmasto, Suillosporium Pouzar, Trechispora P. Karst., and Tubulicium Oberw (Larsson, 2007;
Spirin et al., 2021).
Trechispora is the genus type of Trechisporales and Hydnodontaceae. It is the largest genus in
this order, with more than 50 accepted species (Meiras-Ottoni et al., 2021;Zhao and Zhao, 2021).
Identification keys for Trechispora species recorded in China and Brazil have been provided by
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Liu et al. Outline of Trechisporales (Agaricomycetes, Basidiomycota)
some fungal taxonomists (Chikowski et al., 2020;Meiras-Ottoni
et al., 2021;Zong et al., 2021). Trechispora was typified with
Trechispora onusta P. Karst. [= Trechispora hymenocystis (Berk.
and Broome) K.H. Larss.] (Karsten, 1890). It is characterized by
the resupinate basidiomata (a few species have stipitate, flabellate,
and effused–reflexed basidiomata) with smooth grandinioid,
odontioid, hydnoid, or poroid hymenophores, a monomitic or
dimitic hyphal structure with clamped generative hyphae and
smooth to verrucose or aculeate basidiospores (Larsson, 1992;
Larsson et al., 2004). Most species in Trechispora are soil-dwelling
(Larsson et al., 2004). One remarkable character is the presence of
ampullate septa on the subicular and especially on some hyphae
of the mycelial cords. Above all, ampullate septa are only known
from Scytinopogon,Trechispora, and Porpomyces mucidus (Pers.)
Jülich within Trechisporales (Furtado et al., 2021;Meiras-Ottoni
et al., 2021).
Larsson (2007) used the term “Sistotremastrum family” for
the first time to accommodate Sistotremastrum suecicum Litsch.
ex J. Erikss. and Sistotremastrum niveocremeum [= Sertulicium
niveocremeum (Höhn. and Litsch.) Spirin and K.H. Larss.].
Since then, “Sistotremastrum family” has been adopted by some
taxonomists (Telleria et al., 2013;Liu et al., 2019). In this
work, the phylogeny of Trechisporales is carried out based
on combined 5.8S +nLSU sequences. In addition, Bayesian
evolutionary and divergence time analyses are also carried out to
indicate the divergence time of Trechisporales, Hydnodontaceae,
and Sistotremastrum family. We outline the Sistotremastrum
family and discuss the difference between Hydnodontaceae and
Sistotremastrum family.
During investigations on the diversity of wood-rotting fungi,
seven resupinate specimens were collected from China and
Malaysia. Their morphology corresponds to the concept of
Trechispora. To confirm their affinity, phylogenetic analyses
based on the ITS sequences are carried out. Both morphological
characteristics and molecular evidence demonstrate that
these seven resupinate specimens represent the four new
species of Trechispora.
In addition, we downloaded the type sequences of
Trechispora daweishanensis C.L. Zhao, Trechispora xantha
C.L. Zhao, and Sistotremastrum limonadense G. Gruhn
and P. Alvarado from GenBank. We also studied the
type specimens of T. daweishanensis and T. xantha. In
conclusion, T. daweishanensis and T. xantha were transferred
to Brevicellicium, while S. limonadense was transferred to
Sertulicium.
MATERIALS AND METHODS
Morphological Studies
Macro-morphological descriptions are based on field notes
and dry herbarium specimens. Microscopic structures are
photographed using a Nikon Digital Sight DS-L3 (Japan) or
Leica ICC50 HD (Japan) camera. Microscopic measurements
are made from slide preparations of dry tissues stained with
1% Phloxine B (C20H4Br4Cl2K2O5) (Fan et al., 2021). We
also use other reagents, such as Cotton Blue and Melzer’s
reagent following Dai’s (2010) study. Spore measurements
include both with ornamentation and without ornamentation.
The following abbreviations are used: KOH = 5% potassium
hydroxide; CB = Cotton Blue; CB(+) = weakly cyanophilous;
CB−= acyanophilous in Cotton Blue; IKI = Melzer’s reagent;
IKI−= neither amyloid nor dextrinoid in Melzer’s reagent;
L= mean spore length (arithmetic average of all spores including
ornamentation); W= mean spore width (arithmetic average of
all spores including ornamentation); Q= a variation in the L/W
ratios between the specimens studied; L0= mean spore length
(arithmetic average of all spores excluding ornamentation);
W0= mean spore width (arithmetic average of all spores
excluding ornamentation); Q0= a variation in the L0/W0ratios
between the specimens studied; n(a/b) = the number of
spores (a) measured from a given number of specimens (b).
When presenting spore size variation, 5% of measurements
are excluded from each end of the range and these values
are given in parentheses. Special color terms follow Petersen
(1996). Herbarium abbreviations follow Thiers (2018). The
studied specimens are deposited at the herbarium of the Institute
of Microbiology, Beijing Forestry University (BJFC), and the
herbarium of Southwest Forestry University (SWFC).
DNA Extraction, Polymerase Chain
Reaction Amplification, and Sequencing
Total genomic DNA from the dried specimens is extracted
by a CTAB rapid plant genome extraction kit (Aidlab
Biotechnologies Company Limited, Beijing, China) according to
the manufacturer’s instructions with some modifications (Liu
and Yuan, 2020;Du et al., 2021). The ITS regions are amplified
with the primers ITS4 and ITS5 (White et al., 1990). The nLSU
regions are amplified with the primers LR0R and LR7 (Vilgalys
and Hester, 1990).
The polymerase chain reaction (PCR) procedure for ITS is as
follows: initial denaturation at 95◦C for 3 min, followed by 35
cycles at 94◦C for 40 s, 58◦C for 45 s, and 72◦C for 1 min, and a
final extension of 72◦C for 10 min. The PCR procedure for nLSU
was as follows: initial denaturation at 94◦C for 1 min, followed by
35 cycles at 94◦C for 30 s, 48◦C for 1 min, and 72◦C for 1.5 min,
and a final extension of 72◦C for 10 min (Zhao et al., 2015;Liu
and Dai, 2021). The PCR products are purified and sequenced
in the Beijing Genomics Institute, China, with the same primers
used in the PCR reactions.
Phylogenetic Analyses
Two combined matrices, an ITS1-5.8S-ITS2 (ITS) data set and
a two-gene data set (5.8S +nLSU), are used for phylogenetic
analyses. Phylogenetic analyses are performed with maximum
likelihood (ML), maximum parsimony (MP), and Bayesian
inference (BI) methods in the ITS data set. Phylogenetic analyses
are performed with ML and BI methods in the combined two-
gene data set (5.8S +nLSU). Species and strain sequences
are adopted partly from 28S- and ITS-based tree topologies
established by Meiras-Ottoni et al. (2021) and Spirin et al.
(2021). New sequences generated in this study, along with
reference sequences retrieved from GenBank (Table 1), are
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Liu et al. Outline of Trechisporales (Agaricomycetes, Basidiomycota)
TABLE 1 | Information of taxa used in phylogenetic analyses.
Species Collector ID (herbarium ID) GenBank accession no.
ITS nLSU
Auricularia sp. PBM 2295 DQ200918 AY634277
Brevicellicium atlanticum LISU 178566 (holotype) NR_119820 HE963774
Brevicellicium atlanticum LISU 178590 HE963775 HE963776
Brevicellicium daweishanense CLZhao 18255 (SWFC) MW302338 MW293867
Brevicellicium daweishanense CLZhao 17860 (SWFC, holotype) MW302337 MW293866
Brevicellicium exile MA-Fungi 26554 (holotype) HE963777 HE963778
Brevicellicium olivascens KHL 8571 (GB) HE963792 HE963793
Brevicellicium olivascens MA-Fungi 23496 HE963787 HE963788
Brevicellicium xanthum CLZhao 17781 (SWFC) MW302340 MW293869
Brevicellicium xanthum CLZhao 2632 (SWFC, holotype) MW302339 MW293868
Dextrinocystis calamicola He 5700 (BJFC) MK204534 MK204547
Dextrinocystis calamicola He 5693 (BJFC) MK204533 MK204546
Exidia recisa EL 15-98 (GB) AF347112 AF347112
Exidiopsis calcea MW 331 AF291280 AF291326
Fibrodontia alba TNM F24944 (holotype) KC928274 KC928275
Fibrodontia gossypina AFTOL-ID 599 DQ249274 AY646100
Hyphodontia floccosa Berglund 150-02 (GB) DQ873618 DQ873617
Hyphodontia subalutacea GEL2196 (KAS) DQ340341 DQ340362
Porpomyces mucidus Dai 12692 (BJFC) KT157833 KT157838
Porpomyces submucidus Cui 5183 (BJFC) KU509521 KT152145
Pteridomyces galzinii GB0150230 LR694188 LR694210
Pteridomyces galzinii Bernicchia 8122 (GB) MN937559 MN937559
Scytinopogon angulisporus TFB13611 – JQ684661
Scytinopogon chartaceum FLOR56185 MK458775 –
Scytinopogon pallescens He 5192 (BJFC) – MK204553
Sertulicium chilense MA-Fungi 86368 (holotype) HG315521 –
Sertulicium granuliferum He 3338 MK204552 MK204540
Sertulicium jacksonii Spirin 10425 (H) MN987943 MN987943
Sertulicium lateclavigerum LY 13467 MG913225 –
Sertulicium limonadense LIP 0001683 (holotype) MT180981 MT180978
Sertulicium limonadense He 6276 (BJFC) OK298489*OK298947*
Sertulicium niveocremeum KHL13727 (GB) MN937563 MN937563
Sertulicium vernale Soderholm 3886 (H, holotype) MT002311 MT664174
Sistotremastrum aculeatum Miettinen 10380.1 (H) MN991176 MW045423
Sistotremastrum aculeatum Cui 8401 (BJFC) KX081133 KX081184
Sistotremastrum aculeocrepitans KHL 16097 (URM) MN937564 MN937564
Sistotremastrum confusum KHL 16004 (URM) MN937567 MN937567
Sistotremastrum denticulatum Motato-Vásquez 894 (SP, holotype) MN954694 MW045424
Sistotremastrum fibrillosum LIP 0001413 (holotype) NR_161047 NG_075239
Sistotremastrum fibrillosum s. l. GUY13-119 (GG) MG913224 MG913210
Sistotremastrum fibrillosum s. l. KHL 16988 (MG) MN937568 MN937568
Sistotremastrum geminum Miettinen 14333 (MAN, holotype) MN937568 MN937568
Sistotremastrum induratum Spirin 8598 (H, holotype) MT002324 MT664173
Sistotremastrum mendax KHL 12022 (O, holotype) MN937570 MN937570
Sistotremastrum rigidum Motato-Vásquez 833 (SP, holotype) MN954693 MW045435
Sistotremastrum suecicum Kunttu 5959 (H) MT075859 MT002335
Sistotremastrum suecicum Miettinen 14550.1 (H) MT075860 MT002336
Sistotremastrum suecicum KHL 11849 (GB) MN937571 MN937571
Sistotremastrum vigilans Fonneland 2011-78 (O, holotype) MN937572 MN937572
Sistotremastrum vigilans Spirin 8778 (H) MN991182 MN991182
Subulicystidium tropicum He 3968 (BJFC) MK204531 MK204544
Suillosporium cystidiatum Spirin 3830 (H) MN937573 MN937573
Trechispora alnicola AFTOL-ID 665 DQ411529 AY635768
Trechispora araneosa KHL8570 (GB) AF347084 AF347084
Trechispora bambusicola CLZhao 3302 (SWFC) MW544021 MW520171
Trechispora bispora CBS 142.63 (holotype) MH858241 MH869842
Trechispora cohaerens TU 110332 UDB008249 –
Trechispora cohaerens TU 115568 UDB016421 –
Trechispora confinis KHL11064 (GB) AF347081 AF347081
(Continued)
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Liu et al. Outline of Trechisporales (Agaricomycetes, Basidiomycota)
TABLE 1 | (Continued)
Species Collector ID (herbarium ID) GenBank accession no.
ITS nLSU
Trechispora copiosa AMO456 MN701019 MN687976
Trechispora copiosa AMO422 (holotype) MN701013 MN687971
Trechispora cyatheae FR-0219442 UDB024014 UDB024014
Trechispora cyatheae FR-0219443 (holotype) UDB024015 UDB024015
Trechispora dentata Dai 22565 (BJFC) OK298491*OM049408*
Trechispora dimitiella Dai 21931 (BJFC) OK298492*OK298948*
Trechispora dimitiella Dai 21181 (BJFC) OK298493*OK298949*
Trechispora echinocristallina FR-0219445 (holotype) UDB024018 UDB024019
Trechispora echinocristallina FR-0219448 UDB024022 –
Trechispora echinospora MA-Fungi 82485 (holotype) JX392845 JX392846
Trechispora farinacea KHL 8793 (GB) AF347089 AF347089
Trechispora farinacea KHL 8451 (GB) AF347082 AF347082
Trechispora fimbriata CLZhao 7969 (SWFC) MW544024 MW520174
Trechispora fimbriata CLZhao 4154 (SWFC, holotype) MW544023 MW520173
Trechispora fissurata CLZhao 4571 (SWFC, holotype) MW544027 MW520177
Trechispora fissurata CLZhao 995 (SWFC) MW544026 MW520176
Trechispora fragilis Dai 20535 (BJFC) OK298494*OK298950*
Trechispora gelatinosa AMO1139 (holotype) MN701021 MN687978
Trechispora gelatinosa AMO824 MN701020 MN687977
Trechispora havencampii SFSU DED8300 (holotype) NR_154418 NG_059993
Trechispora hymenocystis TL 11112 (holotype) UDB000778 UDB000778
Trechispora hymenocystis KHL 8795 (GB) AF347090 AF347090
Trechispora incisa GB0090648 KU747095 KU747087
Trechispora incisa GB0090521 KU747093 –
Trechispora kavinioides KGN 981002 (GB) AF347086 AF347086
Trechispora laevispora Dai 21655 (BJFC) OK298495*OM108710
Trechispora minispora MEXU 28300 (holotype) MK328886 MK328894
Trechispora minispora MEXU 28301 MK328886 MK328895
Trechispora mollis URM 85884 (holotype) MK514945 MH280003
Trechispora mollusca DLL2011-186 (CFMR) KJ140681 –
Trechispora mollusca DLL2010-077 (CFMR) JQ673209 –
Trechispora nivea GB0102694 KU747096 AY586720
Trechispora nivea MA-Fungi 74044 JX392832 JX392833
Trechispora papillosa AMO713 MN701022 MN687979
Trechispora papillosa AMO795 (holotype) MN701023 MN687981
Trechispora regularis KHL10881 (GB) AF347087 AF347087
Trechispora rigida URM 85754 MT406381 MH279999
Trechispora sp. AMO799 MN701008 MN687969
Trechispora sp. AMO440 MN701006 MN687967
Trechispora sp. KHL16968 (O) MH290763 MH290763
Trechispora sp. Dai 22173 (BJFC) OK298496*OK298951*
Trechispora sp. Dai 22174 (BJFC) OK298497*OK298952*
Trechispora stevensonii TU 115499 UDB016467 UDB016467
Trechispora stevensonii MA-Fungi 70669 JX392841 JX392842
Trechispora subsphaerospora KHL 8511 (GB) AF347080 AF347080
Trechispora termitophila AMO396 (holotype) MN701025 MN687983
Trechispora termitophila AMO390 MN701024 MN687982
Trechispora torrendii URM 85886 (holotype) MK515148 MH280004
Tubulicium raphidisporum He 3191 (BJFC) MK204537 MK204545
*Newly generated sequences for this study. New species and new combinations or putatively new species are in bold.
aligned by MAFFT 7 (Katoh et al., 20191) using the “G-
INS-i” strategy and manually adjusted in BioEdit (Hall, 1999).
Unreliably aligned sections are removed before analyses and
attempts are made to manually inspect and improve alignment.
The data matrix is edited in Mesquite v3.70 software (Maddison
and Maddison, 2021). The sequence alignment is deposited
1http://mafft.cbrc.jp/alignment/server/
at TreeBase (submission ID 29141 and 29142). Sequences of
Auricularia sp., Exidia recisa (Ditmar) Fr., and Exidiopsis calcea
(Pers.) K. Wells are included in phylogenetic analyses. They
belong to another order, Auriculariales Bromhead. The order is
close to Trechisporales (Sulistyo et al., 2021). We add these three
sequences in the combined two-gene data set (5.8S +nLSU)
to demonstrate that Trechisporales forms a strongly supported
sister clade to Auriculariales. Sequences of Hyphodontia floccosa
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Liu et al. Outline of Trechisporales (Agaricomycetes, Basidiomycota)
(Bourdot and Galzin) J. Erikss. and Hyphodontia subalutacea
(P. Karst.) J. Erikss. in Hymenochaetales Oberw. obtained from
GenBank are used as outgroups to root trees in the 5.8S +nLSU
analysis. Two sequences of Brevicellicium atlanticum Melo,
Tellería, M. Dueñas and M.P. Martín obtained from GenBank are
used as outgroups to root trees in the ITS analysis.
The MP analysis is applied to the ITS data set sequences.
Approaches to phylogenetic analysis follow Liu and Dai (2021),
and the tree construction procedure is performed in PAUP∗
version 4.0 beta 10 software (Swofford, 2002). All characters
are equally weighted, and gaps are treated as missing data.
Trees are inferred using the heuristic search option with tree
bisection and reconnection (TBR) branch swapping, and 1,000
random sequence additions maxtrees are set to 5,000, branches
of zero length are collapsed, and all parsimonious trees are saved.
Clade robustness is assessed using a bootstrap (BT) analysis with
1,000 replicates (Felsenstein, 1985). Descriptive tree statistics tree
length (TL), consistency index (CI), retention index (RI), rescaled
consistency index (RC), and homoplasy index (HI) are calculated
for each maximum parsimonious tree (MPT) generated.
Maximum likelihood research is conducted with RAxML-
HPC v. 8.2.3 (Stamatakis, 2014) and RAxML-HPC through the
CIPRES Science Gateway (Miller et al., 20092). Statistical support
values (BS) are obtained using nonparametric bootstrapping with
1,000 replicates. The BI analysis is performed with MrBayes
3.2.7a (Ronquist and Huelsenbeck, 2003). Four Markov chains
are run for two runs from random starting trees for 4 million
generations (ITS) and 8 million generations (5.8S +nLSU) until
the split deviation frequency value reaches <0.01, and trees are
sampled every 1,000 generations. The first 25% of the sampled
trees are discarded as burn-in, and the remaining ones are used
to reconstruct a majority rule consensus tree and to calculate
Bayesian posterior probabilities (BPP) of the clades.
The optimal substitution models for the combined data set
are determined using the Akaike information criterion (AIC)
implemented in MrModeltest 2.3 (Posada and Crandall, 1998;
Nylander, 2004) after scoring 24 models of evolution by PAUP∗
version 4.0 beta 10 software (Swofford, 2002). The selected
model applied in the BI analyses and ML analyses is the model
GTR +I+G.
Branches that received BT support for ML (BS), MP (BP),
and BPP greater than 65% (BS), 70% (BP), and 0.9 (BPP) are
considered as significantly supported, respectively. Additionally,
the ML analysis results in the best tree, and only the ML tree
is presented along with the support values from the MP and
BI analyses. FigTree v1.4.4 (Rambaut, 2018) is used to visualize
the resulting tree.
Divergence Time Estimation
Divergence time is estimated with the BEAST v2.6.5 software
package (Bouckaert et al., 2019) with 5.8S and nLSU sequences
representing all main lineages in Basidiomycota (Table 2).
Sequences of the species are adopted partly from the topology
established by Wang et al. (2021).Neurospora crassa Shear and
B.O. Dodge from Ascomycota are designated as outgroup taxon
2http://www.phylo.org
TABLE 2 | Information of taxa used in molecular clock analysis.
Species Specimen no. ITS nLSU
Amylocorticium cebennense HHB-2808 GU187505 GU187561
Anomoloma myceliosum MJL-4413 GU187500 GU187559
Athelia arachnoidea CBS 418.72 GU187504 GU187557
Auricularia heimuer Xiaoheimao LT716074 KY418890
Auricularia sp. PBM 2295 DQ200918 AY634277
Australovuilleminia coccinea MG75 HM046875 HM046931
Boletopsis leucomelaena AFTOL-ID 1527 DQ484064 DQ154112
Bondarzewia montana AFTOL-ID 452 DQ200923 DQ234539
Brevicellicium atlanticum LISU 178566 NR_119820 HE963774
Brevicellicium atlanticum LISU 178590 HE963775 HE963776
Brevicellicium daweishanense CLZhao 17860 MW302337 MW293866
Brevicellicium daweishanense CLZhao 18255 MW302338 MW293867
Brevicellicium exile MA-Fungi 26554 HE963777 HE963778
Brevicellicium olivascens KHL8571 HE963792 HE963793
Brevicellicium olivascens MA-Fungi 23496 HE963787 HE963788
Brevicellicium xanthum CLZhao 17781 MW302340 MW293869
Brevicellicium xanthum CLZhao 2632 MW302339 MW293868
Bridgeoporus sinensis Cui 10013 KY131832 KY131891
Calocera cornea AFTOL-ID 438 AY789083 AY701526
Coltricia perennis Cui 10319 KU360687 KU360653
Coltriciella dependens Dai 10944 KY693737 KY693757
Corticium roseum MG43 GU590877 AY463401
Craterocolla cerasi TUB 020203 KF061265 KF061265
Cryptococcus humicola AFTOL-ID 1552 DQ645516 DQ645514
Dacryopinax spathularia AFTOL-ID 454 AY854070 AY701525
Dextrinocystis calamicola He 5700 MK204534 MK204547
Dextrinocystis calamicola He5693 MK204533 MK204546
Exidia recisa EL 15-98 AF347112 AF347112
Exidiopsis calcea MW 331 AF291280 AF291326
Fasciodontia brasiliensis MSK-F 7245a MK575201 MK598734
Fasciodontia bugellensis MSK-F 5548 MK575204 MK598736
Fibrodontia alba TNMF 24944 KC928274 KC928275
Fibrodontia gossypina AFTOL-ID 599 DQ249274 AY646100
Fomitiporia hartigii MUCL 53551 JX093789 JX093833
Fomitiporia mediterranea AFTOL 688 AY854080 AY684157
Gloeophyllum sepiarium Wilcox-3BB HM536091 HM536061
Gloeophyllum striatum ARIZAN 027866 HM536092 HM536063
Grifola frondosa AFTOL-ID 701 AY854084 AY629318
Gymnopilus picreus ZRL2015011 LT716066 KY418882
Hymenochaete rubiginosa He1049 JQ716407 JQ279667
Hyphodontia densispora LWZ 20170908-5 MT319426 MT319160
Hyphodontia zhixiangii LWZ 20170818-13 MT319420 MT319151
Jaapia argillacea CBS 252.74 GU187524 GU187581
Gomphidius roseus MB 95-038 DQ534570 DQ534669
Kneiffiella barba-jovis KHL 11730 DQ873609 DQ873610
Kneiffiella subalutacea LWZ 20170816-9 MT319407 MT319139
Lepiota cristata ZRL20151133 LT716026 KY418841
Leptosporomyces raunkiaeri HHB-7628 GU187528 GU187588
Leucophellinus hobsonii Cui 6468 KT203288 KT203309
Lyomyces macrosporus LWZ20170817-2 MT319459 MT319194
Multiclavula mucida AFTOL-ID 1130 DQ521417 AY885163
Neoantrodiella gypsea Cui 10372 KT203290 MT319396
Neoantrodiella thujae Dai 5065 KT203293 MT319397
Neurospora crassa OR74A HQ271348 AF286411
(Continued)
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TABLE 2 | (Continued)
Species Specimen no. ITS nLSU
Nigrofomes melanoporus JV 1704/39 MF629835 MF629831
Nigrofomes sinomelanoporus Cui 5277 MF629836 MT319398
Porodaedalea chinensis Cui 10252 KX673606 MH152358
Porpomyces mucidus Dai 12692 KT157833 KT157838
Porpomyces submucidus Cui 5183 KU509521 KT152145
Pteridomyces galzinii GB0150230 LR694188 LR694210
Pteridomyces galzinii Bernicchia8122 MN937559 MN937559
Ramaria rubella AFTOL-ID 724 AY854078 AY645057
Rigidoporus corticola ZRL20151459 LT716075 KY418899
Rigidoporus ginkgonis Cui 5555 KT203295 KT203316
Scytinopogon angulisporus TFB13611 – JQ684661
Scytinopogon pallescens He 5192 – MK204553
Sertulicium chilense MA-Fungi 86368 HG315521 –
Sertulicium granuliferum He 3338 MK204552 MK204540
Sertulicium jacksonii Spirin 10425 MN987943 MN987943
Sertulicium lateclavigerum LY 13467 MG913225 –
Sertulicium limonadense LIP 0001683 MT180981 MT180978
Sertulicium niveocremeum KHL13727 MN937563 MN937563
Sertulicium vernale Soderholm 3886 MT002311 MT664174
Sistotremastrum aculeatum Cui 8401 KX081133 KX081184
Sistotremastrum aculeatum Miettinen 10380.1 MN991176 MW045423
Sistotremastrum aculeocrepitans KHL 16097 MN937564 MN937564
Sistotremastrum confusum KHL 16004 MN937567 MN937567
Sistotremastrum denticulatum MV894 MN954694 MW045424
Sistotremastrum fibrillosum LIP 0001413 NR_161047 NG_075239
Sistotremastrum fibrillosum s. l. GUY13-119 MG913224 MG913210
Sistotremastrum fibrillosum s. l. KHL 16988 MN937568 MN937568
Sistotremastrum geminum Miettinen 14333 MN937568 MN937568
Sistotremastrum induratum Spirin 8598 MT002324 MT664173
Sistotremastrum mendax KHL12022 MN937570 MN937570
Sistotremastrum rigidum MV833 MN954693 MW045435
Sistotremastrum suecicum Kunttu 5959 MT075859 MT002335
Sistotremastrum suecicum Miettinen 14550.1 MT075860 MT002336
Sistotremastrum suecicum KHL 11849 (GB) MN937571 MN937571
Sistotremastrum vigilans Fonneland 2011-78 MN937572 MN937572
Sistotremastrum vigilans Spirin 8778 MN991182 MN991182
Subulicystidium tropicum He3968 MK204531 MK204544
Suillosporium cystidiatum VS3830 MN937573 MN937573
Suillus pictus AFTOL 717 AY854069 AY684154
Thelephora ganbajun ZRL20151295 LT716082 KY418908
Trametes versicolor ZRL20151477 LT716079 KY418903
Trechispora hymenocystis KHL8795 AF347090 AF347090
Tremellodendron sp. PBM2324 DQ411526 –
Tubulicium raphidisporum He 3191 MK204537 MK204545
Ustilago maydis AFTOL 505 AY854090 AF453938
Xylodon heterocystidiatus LWZ 20171015-33 MT319518 MT319264
(Wang et al., 2021). A BEAST XML input file is generated with
BEATUti v2. The estimation of rates of evolutionary changes at
nuclear acids is using ModelTest 3.7 with the GTR substitution
model (Posada and Crandall, 1998). A log-normal distribution
is employed for molecular clock analysis (Drummond and
Rambaut, 2007). A Yule speciation model is selected as prior
assuming a constant speciation rate per lineage. Three fossil
fungi, viz. Paleopyrenomycites devonicus (Taylor et al., 1999,
2005), Archaeomarasmius leggetti (Hibbett et al., 1995, 1997), and
Quatsinoporites cranhamii (Smith et al., 2004;Berbee and Taylor,
2010) are taken from Wang et al.’s (2021) study. An XML file is
conducted for 10 billion generations, producing log files and trees
files. The log file is analyzed in Tracer 1,3and a maximum clade
credibility (MCC) tree is interpreted in TreeAnnotator by trees
file, removing the first 10% of the sampled trees as burn-in, and
viewed in FigTree v1.4.2.
RESULTS
Phylogenetic Analyses
The concatenated 5.8S +nLSU data set contains 50 5.8S and
50 nLSU sequences from 52 fungal specimens representing
35 taxa in Trechisporales. The data set has an aligned
length of 1,528 characters, of which 1,126 are constant,
89 are variable but parsimony-uninformative, and 313 are
parsimony-informative. The average standard deviation (SD)
of split frequencies is 0.005271 (BI). Three new combinations,
namely, Brevicellicium daweishanense,Brevicellicium xanthum,
and Sertulicium limonadense, are proposed based on the
examination of type materials and phylogenetic analyses of type
sequences (Figure 1).
The ITS data set contains sequences from 58 fungal specimens
representing 36 Trechispora taxa (4 new species and another
32 Trechispora taxa). The data set has an aligned length of
753 characters, of which 284 are constant, 72 are variable but
parsimony-uninformative, and 397 are parsimony-informative.
MP analysis yields 13 equally parsimonious trees (TL = 2,318,
CI = 0.398, RI = 0.638, RC = 0.254, and HI = 0.602). The
average SD of split frequencies in BI analyses is 0.006959 (BI). The
phylogenetic tree (Figure 2) reveals four new and independent
lineages represented by our specimens, indicating that they are
phylogenetically distinct from the species currently known in the
genus. In addition, another taxon (Dai 22173 and Dai 22174) is
treated as Trechispora sp.
The combined data set for the molecular clock analysis
includes 100 collections, of which 47 belonged to Trechisporales.
This data set results in a concatenated alignment of 1,588
characters with GTR as the best-fit evolutionary model. The
MCC tree is used to study divergence time. The tree shows that
Trechisporales occurs in a mean stem age of 270.85 Mya with
a 95% highest posterior density (HPD) of 234.1–307.93 Mya
(Figure 3). The tree also shows that the Sistotremastrum family
and Hydnodontaceae occur in a mean stem age of 224.25 Mya
[posterior probabilities (PP) = 0.8] with a 95% HPD of 182.47–
266.75 Mya.
Taxonomy
Sistotremastrum family
“Type genus”: Sistotremastrum J. Erikss.
Habitat: It grows on rotten angiosperm and
gymnosperm wood.
Basidioma are resupinate, thin, pruinose, or waxy.
Hymenophores are smooth, verruculose, or odontioid-
semiporoid. The hyphal structure is monomitic; generative
hyphae bear clamp connections, CB(+). Cystidia and hyphidia
3http://beast.community/tracer
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Liu et al. Outline of Trechisporales (Agaricomycetes, Basidiomycota)
FIGURE 1 | Phylogeny of Trechisporales generated by maximum likelihood (ML) analyses based on combined 5.8S + nLSU sequences. Branches are labelled with
ML bootstrap (BT) >65%, and Bayesian posterior probabilities (BPP) >0.90, respectively. New combinations, the sequence origin from holotype and the type status
of the species in the genus are indicated in bold.
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FIGURE 2 | Phylogeny of Trechispora generated by ML analyses based on combined ITS sequences. Branches are labelled with ML BT >65%, Parsimony
Bootstrap Proportions >70%, and BPP >0.90, respectively. Putatively new species, the sequence origin from the holotype, and the type status of the species in the
genus are indicated in bold.
are present in some species. Basidia are clavate or cylindrical,
often with a median constriction, mostly with 2–4 or 4–6
sterigmata, and rarely with 6–8 sterigmata. Basidiospores are
narrowly ellipsoid, ovoid, or cylindrical, thin-walled (but the wall
is distinct), smooth, inamyloid, and acyanophilous.
Notes:Sistotremastrum family accommodates the genera
Sistotremastrum and Sertulicium in the order Trechisporales
based on its distinct lineage in the phylogenetic analysis. The
combined phylogeny of two-gene data (Figure 1) demonstrates
that Sistotremastrum family forms a supported sister clade to
Hydnodontaceae. Basidia of most species in the Sistotremastrum
family have more than four sterigmata, and basidiospores are
smooth, while basidia of species in Hydnodontaceae have four
sterigmata and their basidiospores are smooth to verrucose
or aculeate. In addition, ampullate septa are only present in
Scytinopogon,Trechispora, and P. mucidus in Hydnodontaceae.
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FIGURE 3 | Maximum clade credibility (MCC) chronogram and estimated
divergence times of all main lineages in Basidiomycota inferred from the
combined data set of 5.8S and LSU regions. The estimated divergence times
of 95% highest posterior density (HPD) for all clades are indicated as node
bars. The colored dots refer to the positions of the mean stem age of
Sistotremastrum family, Hydnodontaceae, Trechisporales, and
Hymenochaetales. The BPP above 0.8 and the mean divergence times of
clades are labelled above and below the branches, respectively, at the nodes.
Trechispora dentata Z.B. Liu and Yuan Yuan, sp. November
Figure 4
MycoBank number: MB 842865.
Type: China, Yunnan province, Sipsongpanna, Mengla
County, XiShuangBanNa Tropical Botanical Garden, on soil, in
southwestern China, ca. E 101◦250, N 21◦410, alt. 570 m. The
vegetation is a natural tropical forest. 4 July 2021, Y.C. Dai 22565
(holotype BJFC 037139).
Etymology:Dentata (Lat.): It refers to the species having a
dentate hymenophore.
Basidioma: They are annual, resupinate, soft when fresh,
fragile when dry, easily separable from the substratum, up
to 2.5-cm long, 2-cm wide, and less than 1-mm thick
at the center; hymenial surface irpicoid, white when fresh,
becoming cream (4A2/3) when dry; margin indistinct and
fimbriate, mycelial cords absent; pores or aculei 3–4/mm;
hymenophore lacerate to dentate; subiculum very thin to almost
absent; tubes or aculei concolorous with a hymenial surface,
less than 1 mm long.
Hyphal structure: Hyphal system is monomitic; generative
hyphae bear clamp connections; ampullate septa occasionally
present in subiculum and trama, up to 5-µm wide; all hyphae
IKI−, CB−are unchanged in KOH; rhomboidal calcium oxalate
crystals are scattered.
Subiculum: Generative hyphae hyaline, thin- to thick-walled,
frequently branched, loosely interwoven, 2–4 µm in diameter.
Tubes or aculei: Generative hyphae in trama hyaline, thin- to
thick-walled, frequently branched, loosely interwoven, 2–3 µm in
diameter; cystidia and cystidioles are absent; basidia are clavate or
barrel-shaped, hyaline, bearing four sterigmata and a basal clamp
connection, 10–15 ×4–5 µm; basidioles are similar to basidia in
shape but slightly shorter.
Basidiospores: They are ellipsoid, hyaline, thick-walled,
aculeate, occasionally with one guttule, IKI−, CB−, (4−)4.1–
5×(3−)3.2–4(−4.1) µm (including ornamentation),
L= 4.46 µm, W= 3.66 µm, Q= 1.22 (n= 60/1); (2.2−)2.6–
3.7(−3.8) ×2–2.5 µm (excluding ornamentation), L0= 3.17 µm,
W0= 2.23 µm, and Q0= 1.42 (n= 60/1).
Notes:T. dentata was discovered in the Yunnan Province
of China. Phylogenetically, T. dentata is close to Trechispora
regularis (Murrill) Liberta with strong support (96% BS, 96%
BP, 1.00 BPP; Figure 2). However, T. regularis is strictly
poroid (Liberta, 1973), and basidiospores of T. dentata are
smaller than that of T. regularis [4.1–5 ×3.2–4 µm vs. 4–
5.5 ×3.5–5 µm in T. regularis (including ornamentation);
Liberta, 1973].
Trechispora dimitiella Z.B. Liu and Yuan, sp. November
Figure 5
MycoBank number: MB 842866.
Type: China, Hainan Province, Haikou, Jinniuling Park, on
a rotten leaf, in southwestern China, ca. E 110◦190, N 20◦10,
alt. 17 m. The vegetation is a plantation in tropical China. 7
November 2020, Y.C. Dai 21931 (holotype BJFC 035830).
Etymology:Dimitiella (Lat.): It refers to the species having a
dimitic hyphal system.
Basidioma: They are annual, resupinate, soft when fresh,
fragile when dry, easily separable from the substratum, up to
6-cm long, 4-cm wide, and approximately 3-mm thick at the
center; the hymenial surface is poroid, pore surface white to
cream (4A2/3) when fresh, becoming white to buff-yellow (4A4)
when dry; margin indistinct, often with emerging mycelial cords;
pores angular, 5–6/mm; dissepiments thin, lacerate; subiculum
up to 1 mm thick; tubes concolorous with a poroid surface,
up to 2 mm long.
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FIGURE 4 | Trechispora dentata (holotype, Dai 22565). (A) A basidioma, (B) hyphae from subiculum, (C) hyphae from trama, (D) hyphae with ampullate septa (black
arrow), (E) basidia and basidioles, and (F) basidiospores. Photo by Ya-Ping Lian and Zhan-Bo Liu.
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Liu et al. Outline of Trechisporales (Agaricomycetes, Basidiomycota)
FIGURE 5 | Trechispora dimitiella (holotype, Dai 21931). (A) A basidioma, (B) hyphae with ampullate septa from subiculum (black arrow), (C) hyphae from tubes, (D)
basidia, (E) basidioles, and (F) basidiospores. Photo by Ya-Ping Lian and Zhan-Bo Liu.
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Hyphal structure: Hyphal system is dimitic; generative hyphae
bear clamp connections; ampullate septa occasionally present
in subiculum and trama, up to 4.5 µm wide; all hyphae IKI−,
CB−are unchanged in KOH; rhomboidal calcium oxalate
crystals are scattered.
Subiculum: Generative hyphae hyaline, thin-walled, rarely
branched, 2–3 µm in diameter; skeletal hyphae thick-walled with
a wide lumen, unbranched, loosely interwoven, 2–4 µm diameter.
Tubes: Generative hyphae hyaline, thin-walled, rarely
branched, 1.5–2.5 µm in diameter; skeletal hyphae thick-walled
with a wide lumen, unbranched, loosely interwoven, 2–3 µm
in diameter; cystidia and cystidioles are absent; basidia are
barrel-shaped, hyaline, bearing four sterigmata and a basal clamp
connection, 9.5–12 ×4–5 µm; basidioles are similar to basidia in
shape but slightly shorter.
Basidiospores: They are ellipsoid, hyaline, thick-walled,
aculeate, IKI−, CB−, (3.5−)3.6–4(−4.2) ×(2.5–)2.7–3.1(−3.2)
µm (including ornamentation), L= 3.84 µm, W= 2.92 µm,
Q= 1.31–1.33 (n= 60/2); (2.6−)2.7–3.4(−3.7) ×2–2.6(−2.9) µm
(excluding ornamentation), L0= 3.04 µm, W0= 2.18 µm, and
Q0= 1.38–1.4 (n= 60/2).
Additional specimen examined (paratypes): China, Yunnan
Province, Jinghong, Primeval Forest Park, on soil, 7 July
2021, Y.C. Dai 22601 (BJFC), Dai 22602 (BJFC). Malaysia,
Selangor, Kota Damansara, Community Forest Reserve,
on rotten angiosperm wood, 7 December 2019, Y.C. Dai
21181 (BJFC 032835).
Notes:T. dimitiella was discovered in China and Malaysia.
Most species in Trechispora are corticioid fungi with a monomitic
hyphal structure, but T. dimitiella is different. Morphologically,
T. dimitiella and Trechispora brasiliensis (Corner) K.H. Larss.
share the poroid hymenophore with a dimitic hyphal system
and aculeate basidiospores. However, the basidiospores of
T. dimitiella are smaller than that of T. brasiliensis [3.6–4 ×2.7–
3.1 µm vs. 4–4.5 ×3–4 µm in T. brasiliensis (including
ornamentation), Larsson, 1992]. Phylogenetically, T. dimitiella
is close to Trechispora incisa K.H Larss. (80% BS, 0.99 BPP;
Figure 2), but T. dimitiella can be easily distinguished from
T. incisa due to its poroid hymenophore with a dimitic
hyphal system because T. incisa has arachnoid to farinose or
minutely granulose hymenophore with a monomitic hyphal
system (Larsson, 1996).
Trechispora fragilis Z.B. Liu and Yuan Yuan, sp. November
Figure 6
MycoBank number: MB 842867.
Type: China, Yunnan Province, Sipsongpanna, Mengla
County, XiShuangBanNa Tropical Botanical Garden, on the
ground of the forest, in southwestern China, ca. E 101◦250, N
21◦410, alt. 570 m. The vegetation is a natural tropical forest. 18
August 2019, Y.C. Dai 20535 (holotype BJFC 032203).
Etymology:Fragilis (Lat.): It refers to the species having
fragile basidiocarps.
Basidioma: They are annual, resupinate, soft when fresh,
fragile when dry, easily separable from the substratum, up
to 3 cm long, 2 cm wide, and less than 1 mm thick at
the center; the hymenial surface is odontoid, white when
fresh, becoming cream (4A2/3) to buff-yellow (4A4) when
dry; margin is indistinct and fimbriate, often with emerging
mycelial cords; aculei sparse, 4–6/mm; subiculum very thin
to almost absent; aculei concolorous with a hymenial surface,
less than 1 mm long.
Hyphal structure: Hyphal system monomitic; generative
hyphae bear clamp connections; ampullate septa occasionally
present in subiculum and aculei, up to 7 µm wide; all hyphae
IKI−, CB−are unchanged in KOH; rhomboidal calcium oxalate
crystals are scattered.
Subiculum: Generative hyphae hyaline, thin- to thick-walled,
frequently branched, loosely interwoven, 1.5–4 µm in diameter.
Aculei: Generative hyphae in trama hyaline, thin- to thick-
walled, frequently branched, loosely interwoven, 1.5–3 µm in
diameter; cystidia and cystidioles are absent; basidia are clavate
shaped, hyaline, bearing four sterigmata, and a basal clamp
connection, 12–14 ×3.5–4 µm; basidioles are similar to basidia
in shape but slightly shorter.
Basidiospores: Ellipsoid, hyaline, thick-walled, aculeate,
IKI−, CB−, (3.2−)3.8–4(−4.2) ×(2.4−)2.5–3 µm (including
ornamentation), L= 3.53 µm, W= 2.79 µm, Q= 1.27 (n= 60/1);
(2.6−)2.8–3.7(−4) ×(1.9−)2–2.7(−3.1) µm (excluding
ornamentation), L0= 3.16 µm, W0= 2.26 µm, and Q0= 1.40
(n= 60/1).
Notes:T. fragilis was discovered in the Yunnan Province
of China. Phylogenetically, T. fragilis groups with Trechispora
termitophila Meiras-Ottoni and Gibertoni and Trechispora
havencampii (Desjardin and B.A. Perry) Meiras-Ottoni and
Gibertoni (69% BS, 0.92 BPP; Figure 2). T. termitophila can
be easily distinguished from T. fragilis due to its coralloid
basidioma. In addition, the basidiospores of T. fragilis are
smaller than that of T. termitophila [6.5–7.5 µm vs. 4.5–5 µm
in T. termitophila (including ornamentation), Meiras-Ottoni
et al., 2021]. T. havencampii can also be easily distinguished
from T. fragilis due to its coralloid basidioma. In addition,
basidiospores of T. fragilis are smaller than that of T. havencampii
[3.8–4 ×2.5–3 µm vs. 5.2–6.5 ×3.5–4.2 µm in T. havencampii
(including ornamentation), Desjardin and Perry, 2015].
Trechispora laevispora Z.B. Liu, Y.D. Wu and Yuan Yuan, sp.
November Figure 7
MycoBank number: MB 842868.
Type: China, Inner Mongolia Autonomous Region, Arxan,
Bailang Feng Scenic Spot, on the charred trunk of Larix, in
southwestern China, ca. E 119◦560, N 47◦100, alt. 1,511 m. The
vegetation is a natural boreal forest. 25 August 2020, Y.C. Dai
21655 (holotype BJFC 035556).
Etymology:Laevispora (Lat.): It refers to the species having
smooth basidiospores.
Basidioma: They are annual, resupinate, soft when fresh and
dry, up to 8 cm long, 3 cm wide, and less than 1 mm thick at the
center; the hymenial surface is smooth, white when fresh and dry;
margin is indistinct and fimbriate, often with emerging mycelial
cords; subiculum very thin to almost absent.
Hyphal structure: Hyphal system monomitic; generative
hyphae bear clamp connections; ampullate septa frequently
present in subiculum and hymenium, up to 7 µm wide; all hyphae
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FIGURE 6 | Trechispora fragilis (holotype, Dai 20535). (A) A basidioma, (B) hyphae with ampullate septa from subiculum (black arrows), (C) hyphae from trama, (D)
hymenium with basidioles, (E) basidia, and (F) basidiospores. Photo by Ya-Ping Lian and Zhan-Bo Liu.
IKI−, CB−are unchanged in KOH; rhomboidal calcium oxalate
crystals are abundant.
Subiculum: Generative hyphae hyaline, thin-walled, frequently
branched, loosely interwoven, 1.5–3 µm in diameter.
Hymenium: Generative hyphae in subhymenium hyaline,
thin-walled, frequently branched, 1.5–3 µm in diameter;
cystidia and cystidioles are absent; basidia are clavate shaped,
hyaline, bearing four sterigmata and a basal clamp connection,
11.5–15 ×4–5 µm; basidioles are similar to basidia in shape but
slightly shorter.
Basidiospores: Ellipsoid, hyaline, thin-walled, smooth,
IKI−, CB−, (2.5−) 2.6–3.2(−3.3) ×(1.8−)1.9–2.2(−2.5) µm,
L= 2.92 µm, W= 2.04 µm, and Q= 1.43 (n= 60/1).
Notes:T. laevispora was discovered in the Inner Mongolia
Autonomous Region of China. Phylogenetically, T. laevispora
groups with Trechispora cohaerens (Schwein.) Jülich and Stalpers
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FIGURE 7 | Trechispora laevispora (holotype, Dai 21655). (A) A basidioma, (B,C) hyphae from subiculum, (D) subicular hyphae with ampullate septa (black arrow)
and a piece of hymenium, (E) basidia and basidioles, and (F) basidiospores. Photo by Ya-Ping Lian and Zhan-Bo Liu.
with strong support (94% BS, 96% BP, 1.00 BPP; Figure 2).
Both species share a smooth hymenophore, a monomitic hyphal
system with smooth basidiospores. However, basidiospores
of T. cohaerens are thick-walled and larger than that of
T. laevispora (3.5–4 ×2.2–2.5 µm in T. cohaerens;Larsson,
1992).
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B. daweishanense (C.L. Zhao) Z.B. Liu and Yuan Yuan, comb.
November
MycoBank number: MB 842869.
Basionym:T. daweishanensis C.L. Zhao, Phytotaxa
479(2): 153 (2021).
Type: China. Yunnan Province, Honghe, Pingbian County,
Daweishan National Nature Reserve, on the fallen branch of
angiosperms, 1 August 2019, CLZhao 17860 (holotype SWFC).
Description: See Zong et al. (2021, as T. daweishanensis).
B. xanthum (C.L. Zhao) Z.B. Liu and Yuan Yuan, comb.
November
MycoBank number: MB 842870.
Basionym:T. xantha C.L. Zhao, Phytotaxa 479(2): 155 (2021).
Type: China. Yunnan Province, Yuxi, Xinping County,
Mopanshan National Forestry Park, on the trunk of Albizia
julibrissin, 20 August 2017, CLZhao 2632 (holotype SWFC).
Description: See Zong et al. (2021, as T. xantha).
Notes:Zong et al. (2021) described T. daweishanensis and
T. xantha as new species. However, in our phylogeny, they
belong to the genus Brevicellicium (98% BS, 1.00 BPP; Figure 1).
The type specimens of abovementioned species are studied
[CLZhao 17860 (SWFC); CLZhao 2632 (SWFC)]. We do not
observe ampullate hyphae from type materials as mentioned
by Zong et al. (2021). We suppose that Zong et al. (2021)
confused basidioles with ampullate hyphae (ampullate septa
on some generative hyphae), which are remarkable characters
of Trechispora. In fact, T. daweishanensis and T. xantha have
a smooth hymenophore, a monomitic hyphal structure with
clamped generative hyphae, and the absence of ampullate septa.
They fit Brevicellicium well. Herein, we combine these two species
in Brevicellicium based on morphological and phylogenetic
evidence (Figure 1).
S. limonadense (G. Gruhn and P. Alvarado) Z.B. Liu and Yuan
Yuan, comb. November
MycoBank number: MB 842871.
Basionym:S. limonadense G. Gruhn and P. Alvarado,
Phytotaxa 498(1): 36 (2021).
Type: French Guiana. On the bark of an unidentified
dead trunk lying on the ground, October 22, 2013, LIP
0001683 (holotype).
Description: See Gruhn and Alvarado (2021, as
S. limonadense).
Notes:Gruhn and Alvarado (2021) described S. limonadense
as a new species. However, at the same time, Spirin et al.
(2021) segregated the species around S. niveocremeum (Höhn.
and Litsch.) J. Erikss. into the new genus Sertulicium. In our
phylogeny, S. limonadense groups with Sertulicium granuliferum
(Hallenb.) Spirin and Volobuev Sertulicium lateclavigerum
(Boidin and Gilles) Spirin and Viner (Figure 1). We did not
study specimens, but S. limonadense is characterized by smooth
to tuberculate hymenophore and basidia have 6–8 sterigmata
(Gruhn and Alvarado, 2021) and fits Sertulicium better. Hence,
we transfer S. limonadense to Sertulicium.
DISCUSSION
Larsson (2007) showed that S. suecicum and S. niveocremeum
(= S. niveocremeum) formed a strongly supported sister clade
(94% BS, 1.00 BPP) to Hydnodontaceae within Trechisporales.
However, in his phylogenetic analysis of 5.8S +nLSU, there
were a few species in Hydnodontaceae and Sistotremastrum to
establish a new family for S. suecicum and S. niveocremeum.
Hence, Larsson (2007) named this clade Sistotremastrum family.
The same strongly supported topology was recovered by Telleria
et al. (2013); Gruhn et al. (2018), and Meiras-Ottoni et al. (2021)
by the nLSU phylogenetic analysis. Spirin et al. (2021) presented
a comprehensive study of Sistotremastrum and Sertulicium with
17 species. They used the nLSU region to perform phylogenetic
analyses of 16 species in the two genera (Figure 1 in Spirin et al.,
2021), except for Sertulicium chilense (Telleria, M. Dueñas and
M.P. Martín) Spirin and Volobuev because the nLSU sequences
of S. chilense were absent. However, they were not able to generate
high support values for the node connecting Sistotremastrum and
Sertulicium (87% BS, 0.87 BPP, Figure 1 in Spirin et al., 2021). As
a result, they gave up establishing a new family too.
ITS1-5.8S-ITS2 is an important marker used for the barcoding
of fungal species (Liu et al., 2021;Wangsawat et al., 2021).
However, the difficulty in aligning ITS sequences for fungi in
Trechisporales is evident because it is a data set covering taxa in
distinct taxonomic levels (Larsson, 2007). Therefore, it is not a
good idea to run combined analyses of ITS +nLSU, so we use
the most stable and conservative portion of ITS (5.8S) and nLSU
to our phylogenetic analyses of Sistotremastrum and Sertulicium
(5.8S +nLSU) (Figure 1). We add S. chilense and S. limonadense
to phylogenetic analyses. Our results of the Sistotremastrum are
the same as phylogenetic analyses by Spirin et al. (2021,Figure 1).
However, our phylogenetic analyses of Sertulicium are a bit
different from that by Spirin et al. (2021,Figure 1) because the
data sets used in both studies are different. Above all, we generate
high support values for the node connecting Sistotremastrum
and Sertulicium from ML analysis (93% BS) based on 5.8S and
nLSU sequences; however, BI fails to provide support for the
node (0.76 BPP).
Divergence time is estimated with 5.8S and nLSU sequences
representing all main lineages in Basidiomycota (Figure 3). The
MCC tree shows that Basidiomycota occurs in a mean stem
age of 509.57 Mya. Trechisporales occurs in a mean stem age
of 270.85 Mya. The tree also shows that the Sistotremastrum
family and Hydnodontaceae occur in a mean stem age of
224.25 Mya (PP = 0.8). Zhao et al. (2017) indicate that the
divergence times of Basidiomycota are 530 Mya (the mean stem
age). He et al. (2019) indicate that the divergence times of
Trechisporales and Hydnodontaceae are 259 Mya (the mean
stem age). Our experimental results agree with them. In
this paper, we update the divergence times of Trechisporales
and Hydnodontaceae and define the divergence time of the
Sistotremastrum family.
Bayesian phylogenetic inference fails to provide support for
the node of Sistotremastrum and Sertulicium, so we use the term
“Sistotremastrum family” for the two genera without a formal
description of the new family. In the future, we will sequence
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Liu et al. Outline of Trechisporales (Agaricomycetes, Basidiomycota)
additional DNA regions or whole genomes, for a more robust
phylogenetic analysis.
At present, there are only two species in the Sistotremastrum
family ever been recorded from China, i.e., Sistotremastrum
aculeatum Miettinen and Viner (Cui 8401) and S. granuliferum
(He 3338; CLZhao 5531, 9771). Recently, we collected a specimen
from the Yunnan Province of China (He 6276), and its
morphological and DNA data demonstrated the specimen is
S. limonadense. The species is a new record in China, and
we have uploaded ITS and nLSU sequences of the specimen
(He 6276) to GenBank. Above all, we study all the Chinese
specimens of species in the Sistotremastrum family seriously, and
their morphology fits the descriptions of Gruhn and Alvarado
(2021) and Spirin et al. (2021). We also collected a specimen
from the Hainan Province of China (Dai 17696). The ITS
(OK298490) region is different from Sistotremastrum fibrillosum
G. Gruhn and P. Alvarado by 6%, and morphologically it
is similar to S. fibrillosum. However, we only have a single
specimen, so for the time being we regard Dai 17696 as
Sistotremastrum sp.
In this article, we use the whole ITS region in analyses of
Trechispora to visualize the genetic distances among new taxa
and those already described. T. dentata,T. dimitiella,T. fragilis,
and T. laevispora are described as new to science based on
morphological characteristics and molecular evidence (Figure 2).
Most of these new species are found in subtropical or tropical
Asia and conform to the phenomenon that subtropical or
tropical Asia harbors high taxonomic diversity for all wood-
decaying fungi (Dai, 2012;Cui et al., 2019). We also collected
two resupinate specimens (Dai 22173 and Dai 22174) from
the Hainan Province of China. The morphology of the two
specimens corresponds to the concept of Trechispora and forms a
distinct lineage within the Trechispora clade (100% BS, 1.00 BPP;
Figure 2). However, these specimens are sterile, so we regard Dai
22173 and Dai 22174 as Trechispora spp. temporarily here.
Molecular phylogenetic analyses in the present study show
that Brevicellicium forms a monophyletic clade in which
all Brevicellicium species are included (98% BS, 1.00 BPP;
Figure 1). However, when we add sequences of T. xantha and
T. daweishanensis, we find sequences of a two-species cluster with
Brevicellicium with high support (100% BS, 1.00 BPP; Figure 1).
We request and examine type specimens from Zhao and find
T. xantha and T. daweishanensis corresponding to the concept
of Brevicellicium and they should be transferred to the genus
Brevicellicium (see the notes of B. daweishanense).
DATA AVAILABILITY STATEMENT
The datasets presented in this study can be found in online
repositories. The names of the repository/repositories and
accession number(s) can be found in the article/supplementary
material.
AUTHOR CONTRIBUTIONS
Z-BL: design of the research, performance of the research, and
writing and revising this manuscript. Z-BL, HZ, Y-PL, Y-RW,
C-GW, and W-LM: data analysis and interpretation. Z-BL, YY,
and Y-DW: a collection of the materials. All authors contributed
to the article and approved the submitted version.
FUNDING
The research is supported by the National Natural Science
Foundation of China (Project Nos. 31870007 and 32011540380).
ACKNOWLEDGMENTS
We thank Prof. Dr. Chang-Lin Zhao (SWFC, China) and
Prof. Yu-Cheng Dai for allowing us to study their specimens.
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