Yunnan–Guizhou Plateau: a mycological hotspot

Abstract

Guizhou and Yunnan Provinces (Yungui Plateau) in Southwestern China are well known as biodiversity hotspots. We introduce two new species in this study viz., Mucispora hydei (in Fuscosporellaceae, Fuscosporellales, Sordariomycetes) and Tolypocladium cucullae (Ophiocordycipitaceae, Hypocreales, Sordariomycetes) and six new records based on morpho-molecular analyses. Full descriptions, color photographs and phylogenetic trees to indicate the placements of new species are provided.

Full text

Phytotaxa 523 (1): 001–031
https://www.mapress.com/j/pt/
Copyright © 2021 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Rajesh Jeewon: 20 Sept. 2021; published: 15 Oct. 2021
https://doi.org/10.11646/phytotaxa.523.1.1
1
Licensed under Creative Commons Attribution-N.C. 4.0 International https://creativecommons.org/licenses/by-nc/4.0/
Yunnan–Guizhou Plateau: a mycological hotspot
NALIN N. WIJAYAWARDENE1,2,3,18, LAKMALI S. DISSANAYAKE4,19, QI-RUI LI2,5,20*, DONG-QI DAI1,21*,
YUANPIN XIAO6,7,22, TING-CHI WEN4,7,8,23, SAMANTHA C. KARUNARATHNA9,10,11,24, HAI-XIA WU12,25,
HUANG ZHANG13,26, SAOWALUCK TIBPROMMA9,10,11,27, JI-CHUAN KANG4,28, YONG WANG14,29, XIANG-
CHUN SHEN2,5,30, LI-ZHOU TANG1,31, CHUN-YING DENG15,32*, YANXIA LIU16,33 & YINGQIAN KANG17,34
1 Center for Yunnan Plateau Biological Resources Protection and Utilization, College of Biological Resource and Food Engineering,
Qujing Normal University, Qujing, Yunnan 655011, P.R. China
2 State Key Laboratory of Functions and Applications of Medicinal Plants, Guizhou Medical University, Guiyang 550014, P.R. China
3 Section of Genetics, Institute for Research and Development in Health and Social Care No: 393/3, Lily Avenue, Off Robert
Gunawardane Mawatha, Battaramulla 10120, Sri Lanka
4 The Engineering Research Center of Southwest Bio–Pharmaceutical Resources Ministry of Education, Guizhou University, Guiyang
550025, Guizhou Province, China
5 The Key Lab of Optimal Utilization of Natural Medicine Resources, School of Pharmaceutical Sciences, Guizhou Medical University,
University Town, Guian New District, Guizhou 550025, P.R. China
6 Institute of Excellence in Fungal Research, School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand
7 State Key Laboratory Breeding Base of Green Pesticide and Agricultural Bioengineering, Key Laboratory of Green Pesticide and
Agricultural Bioengineering, Ministry of Education, Guizhou University, Guiyang 550025, China
8 The Mushroom Research Centre, Guizhou University, Guiyang 550025, China
9 Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Science,
Kunming 650201, Yunnan, P.R. China
10 CIFOR-ICRAF, World Agroforestry Centre, Kunming 650201, Yunnan, P.R. China
11 Centre for Mountain Futures (CMF), Kunming Institute of Botany, Kunming, Yunnan, 650201, P.R. China
12 International Fungal Research and Development Centre, The Research Institute of Resource Insects, Chinese Academy of Forestry,
Kunming 650224, PR China
13 Faculty of Agriculture and Food, Kunming University of Science & Technology, Kunming 650500, People’s Republic of China
14 Department of Plant Pathology, Agriculture College, Guizhou University, Guiyang 550025, P.R. China
15 Guizhou institute of biology, Guizhou academy of science, Guiyang, 550009, P.R. China
16 Guizhou Academy of Tobacco Science, No. 29, Longtanba Road, Huanshanhu District, Guiyang City, Guizhou province, P.R. China
17 Key Laboratory of Environmental Pollution Monitoring and Disease Control, Ministry of Education of Guizhou & Guizhou Talent
Base for Microbiology and Human Health, Key Laboratory of Medical Microbiology and Parasitology of Education Department of
Guizhou, School of Basic Medical Sciences, Guizhou Medical University, Guiyang, P.R. China
18
�
nalinwijayawardene@yahoo.com; https://orcid.org/0000-0003-0522-5498
19
�
dmlsdlakmali.ld@gmail.com; https://orcid.org/0000-0003-2933-3127
20
�
lqrnd2008@163.com; https://orcid.org/0000-0001-8735-2890
21
�
cicidaidongqin@gmail.com; https://orcid.org/0000-0001-8935-8807
22
�
emmaypx@gmail.com; https://orcid.org/0000-0003-1730-3545
23
�
tingchiwen@yahoo.com; https://orcid.org/0000-0003-1744-5869
24
�
samantha@mail.kib.ac.cn; https://orcid.org/0000-0001-7080-0781
25
�
aileen2008haixia@gmail.com; https://orcid.org/0000-0002-7169-6717
26
�
zhanghuang2002113@163.com; https://orcid.org/0000-0002-9464-2981
27
�
saowaluckfai@gmail.com; https://orcid.org/0000-0002-4706-6547
28
�
jckang@gzu.edu.cn; https://orcid.org/0000-0002-6294-5793
29
�
yongwangbis@aliyun.com; https://orcid.org/0000-0003-3831-2117
30
�
shenxiangchun@126.com; https://orcid.org/0000-0002-4333-9106
31
�
tanglizhou@163.com; https://orcid.org/0000-0002-6988-1876
32
�
171934233@qq.com; https://orcid.org/0000-0002-0960-0948
33
�
liuyanxia306@163.com; https://orcid.org/0000-0002-2634-0068
34
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449164105@qq.com; https://orcid.org/0000-0003-0189-9655
*Corresponding authors: Dong-Qin Dai,
�
cicidaidongqin@gmail.com, QI-RUI Li,
�
lqrnd2008@163.com,
Chun-Ying Deng,
�
171934233@qqcom
Abstract
Guizhou and Yunnan Provinces (Yungui Plateau) in Southwestern China are well known as biodiversity hotspots. We
introduce two new species in this study viz., Mucispora hydei (in Fuscosporellaceae, Fuscosporellales, Sordariomycetes)

WIJAYAWARDENE ET AL.
2 • Phytotaxa 523 (1) © 2021 Magnolia Press
and Tolypocladium cucullae (Ophiocordycipitaceae, Hypocreales, Sordariomycetes) and six new records based on morpho-
molecular analyses. Full descriptions, color photographs and phylogenetic trees to indicate the placements of new species
are provided.
Keywords: 2 new species, polyphasic approach, six new records, species diversity, taxonomy
Introduction
Guizhou and Yunnan Provinces (Yungui Plateau) in Southwestern China are known as biodiversity hotspots with, high
floral, faunal and microbial diversity (Xu et al. 2017). Due to its temperate climate, beautiful scenic spots such as
waterfalls and caves, and variety of ethnic groups, Guizhou is one of the most environmentally and culturally diverse
provinces in China (Liu et al. 2013; Chi et al. 2017; Liu et al. 2018). It is a mountainous province home to several rare
animal species, like the Kuankuoshui salamander (Pseudohynobius kuankuoshuiensis) that is not found anywhere else
in the world (Sparreboom 2014). Guizhou owns an average 61.92% karst landforms out of all the landforms and the
main vegetation types in Guizhou are broadleaf and mixed forests (Liu et al. 2018). Domestically, Yunnan Province
is contiguous with Guizhou, Sichuan, Guangxi, and Tibet in China and shares international borders with Vietnam,
Laos and Myanmar (Qian et al. 2020). Natural resources and biodiversity in Yunnan are abundant, and approximately
19,333 plant species belong to 3,084 genera, and 440 families can be found of which 17,000 are endemic (Qian et
al. 2020). Yunnan has three climatic areas viz. the tropical area at the southwest, south, and southeastern border; the
subtropical zone in the west, middle and east; and the temperate zone in the high-elevation area in the northwest (Yang
et al. 2008; Qian et al. 2020). Main vegetation types in Yunnan are Tropical Rainforest, Monsoon Forest, Evergreen
Broadleaf Forest, Sclerophyllous Evergreen Broadleaf Forest, Deciduous Broadleaf Forest, Subtropical Needleleaf
Forest, Temperate Needleleaf Forest, Bamboo Forest, Savanna-like Shrubby Grassland, Scrub and Meadow (Qian et
al. 2020). During the last decade (2010–2020), a considerable number of mycology studies have been carried out in
this region. Taxonomical studies of micro-fungi based on morpho-molecular analyses, wild mushroom cultivation and
domestication, fungal secondary metabolite analyses and ethnomycological surveys are some of the major research
areas.
Taxonomic research based on DNA sequences of non-pathogenic fungi is one of the popular topics among research
groups on the Yungui Plateau. Researchers focus on micro-fungi based on their life modes (e.g., saprobic, epiphytic
and endophytic) or habitat/niche (e.g., freshwater fungi, mycorrhizal fungi, karst fungi, air fungi). Fungal pathology
(agricultural pathogens, clinical pathogens) is another well-developed field on the Yungui Plateau. Pathogens of
agricultural crops, timber plants, ornamental plants and medicinal plants have been broadly studied. Entomopathogenic
fungi on the Yungui Plateau is also a popular discipline.
In the course of our fieldwork on the Yungui Plateau, we encountered several interesting fungal specimens, and
morpho-molecular analyses confirmed that these taxa comprise two new species and six new records (country, host or
new record in Yunnan or Guizhou).
Materials and methodology
Sample collection and incubation
Living plant materials with disease symptoms and dead plant materials were randomly collected from Yunnan and
Guizhou provinces. Temperature, date, time, elevation, and humidity of the collection sites were recorded. Samples
were sealed in Ziploc plastic bags and returned to the laboratory. Samples were incubated using a moist chamber,
sealed them and incubated at room temperature.
Isolation, morphological examination and maintain specimens and cultures
Single spore isolation was followed to isolate fungi (Senanayake et al. 2020). Ascomata/ conidiomata were sectioned
with a razor blade, centrum tissue containing ascospores were removed using a sterile needle and placed in sterile
water. A water drop, which contained the ascospore /conidial suspension, was placed on Water Agar (WA) (15 g agar,
1000 mL sterile distilled water) and incubated overnight at room temperature. Germinated spores were transferred

YUNNAN–GUIZHOU PLATEAU: A MYCOLOGICAL HOTSPOT Phytotaxa 523 (1) © 2021 Magnolia Press • 3
to potato dextrose agar (PDA Difco; 39 g/L sterile distilled water). Dried specimens were deposited at Herbaria of
Guizhou Medical University, Qujing Normal University and Kunming Institute of Botany. Cultures were deposited at
culture collections at Guizhou Medical University, Kunming Institute of Botany.
Morphological characteristics were captured by using a digital camera fitted on to a Nikon ECLIPSE 80i compound
microscope. Squash mount preparations (Sutton 1980) were used to observe micro-morphological characteristics
such as asci, ascospores and pseudoparaphyses in sexual morph; conidiophores, conidiogenous cells, conidia in
asexual morphs. Free hand sections were taken to observe ascoma and peridium structures and shape of conidiomata.
Melzer’s reagent was used to stain the asci and apical rings, whereas Indian ink was used to stain mucilaginous sheaths
surrounding the ascospores. Observed characteristics were presented as photo plates which were edited and combined
using Adobe Photoshop version CS5 (Adobe Systems Inc., United States) and micro-morphological structures were
measured in a Tarosoft (R) Image Frame Work version 0.9.7 program. Index Fungorum identifiers were obtained after
registered new names as outlined in Index Fungorum (2021). New species are established as per guidelines established
by Jeewon & Hyde (2016).
Phylogeny
DNA extraction, PCR amplification and sequencing
Genomic DNA of microfungi was extracted from fresh mycelia grown on PDA at 25–27°C using the Biospin Fungus
Genomic DNA Extraction Kit (BioFlux®, Hangzhou, and P.R. China) according to the manufacturer’s instructions.
DNA was also extracted from fruiting bodies of Mucispora sp. Surface of fruiting bodies was sterilized by
75% alcohol and rinsed three times by sterile water. Conidia and conidiophores were picked up by sterilized forceps
and ground in a mortar into powder with liquid nitrogen. OMEGA E.Z.N.A. Forensic DNA Kit was used following
manufacturer’s instructions.
Amplification of LSU, SSU, ITS genes were performed by using LR0R/ LR5, NS1/ NS4 and ITS5/ ITS4 primers
respectively (White et al. 1990). PCR products were sent for sequencing at Shanghai Sangon Biological Engineering
Technology & Services Co. (Shanghai, P.R. China). All newly generated sequences were deposited in GenBank and
accession numbers were obtained.
TABLE 1. Genes/loci used in the study with PCR primers, references and protocols.
Gene region Primers Thermal cycles Reference
ITS ITS5/ ITS4 (95 °C: 30 s, 55 °C:50 s, 72 °C: 90 s) × 35 cycles White et al. (1990)
LSU LR0R/ LR5 (95 °C: 30 s, 55 °C:50 s, 72 °C: 90 s) × 35 cycles Vilgalys & Hester (1990), Rehner et al. (1994)
SSU NS1/ NS4 (95 °C: 30 s, 55 °C:50 s, 72 °C: 90 s) × 35 cycles White et al. (1990)
Phylogenetic analyses
Phylogenetic analyses were conducted based on the combined relevant genes (Table 1). Single gene alignment was
carried out for prior comparable tree topologies. The combined gene sequence matrix was built based upon taxa
generated in this study and related sequences retrieved from GenBank. Sequences were combined and aligned in Mega
6.0.5 (Tamura et al. 2013) and MAFFT: multiple sequence alignment software version 7.215 (Katoh et al. 2019) and
manually improved where necessary. Sequence alignment was converted to NEXUS file for maximum parsimony
analysis using ClustalX2 v. 1.83 (Thompson et al. 1997) and PHYLIP-compatible for maximum likelihood analysis
using ALTER (alignment transformation environment: http://sing.ei.uvigo.es/ALTER/). Phylogenetic analyses were
performed by maximum likelihood (ML), maximum parsimony (MP) and Bayesian Inference (BI) analyses as outlined
below:
a. Maximum Likelihood (ML)
Maximum likelihood analysis (ML) was performed in RaxmlGUI v.1.3 (Silvestro & Michalak 2012) with 1000
thorough bootstrap replicates. The available substitution models comprised a generalized time reversible (GTR) for
nucleotides was applied with a discrete gamma distribution (Silvestro & Michalak 2012). A discrete GAMMA (Yang

WIJAYAWARDENE ET AL.
4 • Phytotaxa 523 (1) © 2021 Magnolia Press
1994) was complemented for each substitution model. Rapid bootstrap analysis (Stamatakis et al. 2014) and search for
a best-scoring ML tree were applied (Silvestro & Michalak 2012).
b. Maximum Parsimony (MP)
MP analysis was carried out with stepwise additions of sequences by using PAUP v. 4.0b10 (Swofford 2002). The
heuristic search option with 1000 random sequences addition and tree-bisection reconnection (TBR) of branch-
swapping algorithm were performed. Maxtrees were setup at 1000. A zero of maximum branch length was collapsed
and gaps were treated as missing data. Calculating of consistency index (CI), retention index (RI), rescaled consistency
index (RC) and homoplasy index (HI) were included in the analysis. The robustness of the most parsimonious tree was
estimated based on 1000 bootstrap replications with each 100 replicates of random stepwise addition of taxa.
c. Bayesian Inference (BI) analysis
BI analysis was performed by MrBayes v. 3.1.2 (Huelsenbeck & Ronquist 2001) with the best-fit model of sequences
evolution estimated with MrModeltest 2.2 (Nylander 2004). Markov Chain Monte Carlo sampling (MCMC) was used
to determine the posterior probabilities (PP) (Rannala & Yang 1996, Zhaxybayeva & Gogarten 2002) in MrBayes v.
3.0b4 (Huelsenbeck & Ronquist 2001). Six simultaneous Markov chains were run for 1000000 to 5000000 generations
based on the standard deviation of split frequencies less than 0.01. Trees were sampled every 1000th generations). First
20% trees were the burn-in phase and were discarded. Remaining trees were used to calculate the posterior probability
(PP).
Taxonomy
In this section, we introduce two new species and five new records.
Karst fungi
Guizhou and Yunnan Provinces are well known destination for its karst formation. During the last five years, Chen et
al. (2017), Zhang et al. (2017, 2018, 2019, 2020) broadly discussed and introduced over 50 new species from caves
in Yungui Plateau. We collected a taxon inhabiting on decaying wood from a cave in Guizhou. Morphologically, it
resembles Melanocephala and Mucispora. Megablast search in NCBI GenBank confirmed that it bears high sequence
similarity to Mucispora. Morphological characteristics of the new taxon is distinct from all the other known taxa in the
genus and phylogenetic analyses based on combined genes, LSU, SSU and ITS also confirmed that it is a new species
of Mucispora.
Mucispora hydei Wijayaw., Q.R. Li, Y.C. Deng, L.S. Dissan & D-Q Dai sp. nov. (FIGURE 1)
Index Fungorum number: IF558463
Etymology:—Named in honour of British mycologist, K.D. Hyde for his immense contributions to mycology
Holotype:—GMB0028
Saprobic on decaying wood. Asexual morph Hyphomycetous. Conidiophores 60–110 × 8–12 µm (x̅ = 78.6 × 9.8
µm, n = 30), macronematous, mononematous, erect, solitary or in small groups on compactly aggregated cells, simple,
cylindrical, smooth, brown, straight or slightly flexuous, percurrently proliferate 2–3 times, 1–2-septate. Conidiogenous
cells holoblastic, integrated, terminal, cylindrical, smooth, pale brown. Conidia 35-50 × 20-30 µm (x̅ = 41.2 × 25.5 µm,
n = 30), acrogenous, solitary, simple, smooth, ellipsoidal to obovoid, hyaline to subhyaline when young, dark brown
when mature, with obvious septa in young conidia, paler at basal cell, truncate at base, sometimes covered by a hyaline
mucilaginous sheath. Sexual morph Undetermined.
Material examined:—CHINA, Guizhou Province, Guiyang, Gaopo Township, Raorao village (106°48’6.54”E,
26°19’3.46”N), on decaying submerged wood, 9th December 2019, Nalin N. Wijayawardene, Q.R Li, (GMB0028,
holotype, NNW56, isotype).
LSU: MW797122, SSU MW800164, ITS MW797039 (Supplementary Table 1)
Known distribution:—Guizhou Province, China

YUNNAN–GUIZHOU PLATEAU: A MYCOLOGICAL HOTSPOT Phytotaxa 523 (1) © 2021 Magnolia Press • 5
Notes:—Yang et al. (2016) introduced the genus Mucispora Jing Yang et al. with M. obscuriseptata J. Yang et
al. as the type species. Besides the type species, the genus comprises two species viz. M. phangngaensis J. Yang &
K.D. Hyde (Yang et al. 2017) and M. infundibulata J. Yang & K.D. Hyde (Hyde et al. 2020). All these species have
been reported from submerged plant materials in Southern Thailand (Prachuap Khiri Khan Province and Phang Nga
Province). In morphology, Mucispora closely resembles Melanocephala but it is specific in its cupulate proliferating
conidiogenous cells and its conidia bearing a central downwardly directed collar with a fimbriate margin’ (Hughes
1979; Yang et al. 2017).
Our new collection did not germinate in different media (WA, PDA, MEA) and in different temperatures thus
we extracted DNA directly from the fruiting body (Zeng et al. 2018). PCR amplification of ITS (ITS4/ ITS5), LSU
(primers: LR5/ LROR) and SSU (primers: NS1/NS4) were successful.
Phylogenetic analyses of combined LSU and ITS genes (Fig. 2) that our new strain is distinct from other taxa.
However, the separation value is medium (69% in ML) and PP value is low. Nevertheless, morphological characters,
of our collection is well-distinct from other Mucispora species (Table 2). Hence, we introduce the fourth species of the
genus, Mucispora hydei. This is the first record of the genus outside Thailand.
TABLE 2. Morphological comparison of Mucispora species
Species Conidiophore Conidia Location
M. phangngaensis 170–305 × 5–7 35–45 × 16.5–25 Phang Nga Province, Thailand
M. obscuriseptata 80–170 × 5–7.5 29–41 × 16–22 Prachuap Khiri Khan Province, Thailand
M. infundibulata 50–65 × 4–6 29–34 × 19–21 Phang Nga Province, Thailand
M. hydei 60–110 × 8–12 35–50 × 20–30 Guizhou Province, China
FIGURE 1. Mucispora hydei (GMB0028, holotype). a. Decaying wood. b, c. Colony on wood. d, e, g. Conidiophores with conidia. f.
Matured conidia. h–j. Conidiophore. Scale bars: b =100 µm, c =200 µm, d, e, h–j = 20 µm, f, g = 40 µm.

WIJAYAWARDENE ET AL.
6 • Phytotaxa 523 (1) © 2021 Magnolia Press
FIGURE 2. RAxML tree based on a combined dataset of partial LSU and ITS sequence analyses. Bootstrap support values for ML equal to
or greater than 60 %, Bayesian posterior probabilities (BYPP) equal to or greater than 0.95 are shown as ML/ BYPP above the nodes. New
isolates are in red bold. The tree is rooted to Conioscypha lignicola and Conioschypha minutispora (FMR11245) and Conioscyphascus
varius. The scale bar represents the expected number of nucleotide substitutions per site.
Saprobic micro fungi on plants
Saprobic fungi are a vital component in the ecosystems since they are crucial in decaying organic matter, recycling
minerals and nutrients (Hyde et al. 2018). In here, we report three new records from Yungui Plateau.
Immersidiscosia eucalypti (Pat.) Kaz. Tanaka, Okane & Hosoya, Persoonia 26: 94 (2011) (FIGURE 3)
Index Fungorum Number: IF519747
Foliicolous, the host plant is Quercus palustris. Sexual morph: Undetermined. Asexual morph: coelomycetous.
Conidiomata 354–522 μm (
x
= 427 μm, n = 5) diameter, 287 μm high, conspicuous, pycnidial, subglobose to sometimes
lenticular in section view, semi-immersed, scattered, unilocular, with relatively thin stromatic base, black, glabrous.
Beak of conidiomata long, 384 μm long, 13 – 61 μm wide. Peridium 18–42 μm wide (upper wall 25–42 μm (
x
= 33
μm, n = 7) wide; basal wall 18–26 μm (
x
= 27 μm, n = 7) wide), composed of 4 – 7 layers, with outer 3–5 layers light
brown and inner layer hyaline, composed of thin-walled cells of textura angularis. Conidiophores up to 45 μm long,

YUNNAN–GUIZHOU PLATEAU: A MYCOLOGICAL HOTSPOT Phytotaxa 523 (1) © 2021 Magnolia Press • 7
cylindrical, branched. Conidia 15.4 – 17 × 2.6 – 3.3 μm (
x
= 16.1 × 3 μm, n = 10), cylindrical to subcylindrical, slightly
curved, 3-septate, hyaline, with an appendage at both ends; basal cell 2–2.8 μm long (
x
= 2.5 μm, n = 10), obconic,
truncate at the base; 2 median cells 10.5–12.2 μm long (
x
= 11.3 μm, n = 10), cylindrical (second cell from the base
4.7–6.6 μm long (
x
= 5.6 μm, n = 10), third cell 4.6–6.7 μm long (
x
= 5.7 μm, n = 10)); apical cell 1.7–3.1 μm long (
x
= 2.7 μm, n = 10). Appendage single, cellular, unbranched, filiform, flexuous or straight appendage; apical appendage
7.9–9.1 × 0.8–1.1 μm (
x
= 8.7 × 1 μm, n = 6); basal appendage 7.8–9.3 × 0.7–1.1 μm (
x
= 8.5 × 0.9 μm, n = 6).
Material examined:—CHINA, Yunnan Province, Dali; 25°43′27″N 100°6′54″E, 2260 m alt.; 11 August 2019;
Hai-Xia Wu leg; collected on a fallen leaf of Quercus palustris (IFRD 500-20) (new country record).
Known hosts and distribution (based on molecular data) :—Thailand, Yunnan China
Notes:—The genus, Immersidiscosia Kaz. Tanaka et al. (2011) was introduced by Tanaka et al. (2011) with
I. eucalypti as the type species. The genus, morphologically resembles Discosia but phylogenetically distinct.
Immersidiscosia eucalypti was reported from both temperate and tropical countries such as France, Italy, Japan and
Tunisia (Tanaka et al. 2011; Hyde et al. 2017; Wijayawardene et al. 2017; Farr & Rossman 2021). This is the first
report of I. eucalypti in China. Further collections are essentially required to study whether this taxon is pathogenic on
Quercus species.
FIGURE 3. Immersidiscosia eucalypti (IFRD 500-20) a. Host leaves. b. Specimen with conidiomata. c. Conidiomata. d. Section of
conidiomata. e. Peridium of conidiomata. f–j. Conidia. Scale bars: b = 300 µm, c, e = 100 µm, d = 200 µm, f–j = 10 µm.

WIJAYAWARDENE ET AL.
8 • Phytotaxa 523 (1) © 2021 Magnolia Press
Helminthosporium velutinum Link [as ‘Helmisporium’], Mag. Gesell. naturf. Freunde, Berlin 3(1–2): 10, tab. 1:9
(1809) (FIGURE 4)
Index Fungorum Number: IF250075
FIGURE 4. Helminthosporium velutinum (HKAS 107064, new host record and a new record from Guizhou Province) a–c. Colony on the
substrate. d. Conidiophores. e–g. Conidiophore and conidia. h–l. Conidia. m. Germinating conidium. n, o. Culture on PDA from. n. above
o. below after 4 weeks. Scale bars: d = 100 μm, e–g =50 μm, h–m = 20 μm.

YUNNAN–GUIZHOU PLATEAU: A MYCOLOGICAL HOTSPOT Phytotaxa 523 (1) © 2021 Magnolia Press • 9
Description. Saprobic on dead twigs, dark brown, effuse, velvety. Sexual morph: Undetermined. Asexual
morph: Mycelium immersed, composed of branched, septate, thick-walled hyphae. Conidiophores mononematous,
macronematous, mostly unbranched, proliferating, dark brown, 96–296 × 5–7 µm (x̅ = 153 × 6 µm, n = 10), 7–12
septate, erect or flexuous, tapering towards apex, bulbous at base with cells near apex of conidiophore guttulate and
fertile. Conidiogenous cells polytretic integrated, intercalary and terminal. Conidia 99–131 × 20–36 µm (x̅ = 115 × 28
µm, n = 20)single, obclavate, pale brown to brown, 6–9 distoseptate, smooth, straight or curved, base slightly truncate,
cicatrized and wider than apex, dark brown, apical cell paler than other cells, rounded at apex, guttulate when young,
non-guttulate at maturity.
Culture characteristics: Colonies on PDA, reaching 21 mm diam., after 2 weeks at 20–25 oC, medium dense,
circular to slightly irregular, slightly raised and cottony surface, colony from above: at first white, becoming buff; from
below: blackish white at the margin, black to ash at the center; mycelium blackish.
Material examined: CHINA, Guizhou Province, Huaxi District, Guizhou university garden (South), on a dead
branch of Platanus sp., 05 October 2019, Nalin N. Wijayawardene, NWGUP01 (HKAS 107064, new host record, a
new record from Guizhou Province), ex-type living culture, KUMCC 20–0029
Known hosts and distribution: Guizhou province, China (this study), Yunnan Province, Dali, WanHua stream,
China (Zhu et al. 2016).
Known hosts: Platanus sp. (this study), saprobic on decaying wood submerged in stream (Zhu et al. 2016).
GenBank Numbers: LSU: MW273148, SSU: MW273295, ITS: MW273144
Notes: Helminthosporium velutinum, the type species of Helminthosporium was re-visited by Voglmayr
& Jaklitsch (2017) and designated the epitype and the ex-epitype. The genus was reported with the sexual morph
however, Helminthosporium velutinum lacks the sexual morph (Voglmayr & Jaklitsch 2017). According to Voglmayr
& Jaklitsch (2017), distribution of the species was reported as ‘Widespread and common in temperate Eurasia and
America, probably almost cosmopolitan’. Zhu et al. (2016) reported Helminthosporium velutinum from submerged
wood from Yunnan Province, China. In this study, we collected Helminthosporium velutinum on dead branches of
Platanus sp. from Guizhou Province, China. According to Farr & Rossman (2021), a taxon named Helminthosporium
spiciferum (Nicot 1953) (current name: Curvularia spicifera Index Fungorum 2021) was reported from Platanus
occidentalis. Besides this record, as far as we know, Helminthosporium species have not been reported from Platanus
species. Moreover, this is the first record of this genus from terrestrial habitats from China.
Roussoella pseudohysterioides D.Q. Dai & K.D. Hyde, Fungal Diversity 82: 37 (2016) (FIGURE 5)
Index Fungorum Number: IF552026
Saprobic on decaying bamboo culms. Sexual morph: Ascostromata forming under black area, including 3–5 locules,
up to 3–5 mm long and 0.5–2 mm wide, slightly raised at maturity, irregular, black, coriaceous. Locules in vertical
section 220–280 μm high, 180–330 μm diam., gregarious, subglobose to ellipsoidal, dark brown, with ostiolate
opening. Peridium composed of dark brown cells comprising host and fungal tissues. Hamathecium comprising dense,
2–3.5 μm wide, cellular pseudoparaphyses, indistinctly septate, embedded in a gelatinous matrix. Asci 85–290 ×
7.5–17.5 μm (
x
= 165×10.5 μm, n=30), 8-spored, bitunicate, cylindrical, with a short furcate pedicel, with an apical
ocular chamber. Ascospores 11–19.5 × 4–6.5 μm (
x
= 16.5×5.5 μm, n=30), uniseriate, fusiform-ellipsoidal, 1-septate,
constricted at the septum, narrow at both ends, with striate wall ornamentation, some with obvious verrucose. Asexual
morph: Undetermined.
Material examined:—CHINA, Guizhou Province, Leigong Mountain National Nature Reserve, on dead culm of
bamboo, July 2019, Q.R. Li 2019LGS13 (GMB0009), living cultures, GMBC0009 (new country record).
Known hosts and distribution:—Guizhou, China, Thailand
Known hosts:—Bamboo
GenBank Numbers:—ITS: MW881445; LSU: MW881451; RPB2: MW883345
Notes:—Roussoella, typified by Roussoella nitidula Sacc. & Paol. was introduced by Saccardo & Paoletti (1888).
Most species of Roussoella were observed from monocotyledon, such as bamboo and palms (Dai et al. 2017; Hyde et
al. 2018). Roussoella pseudohysterioides was originally introduced by Dai et al. (2017) isolated from Thailand. This
is the first report of Roussoella pseudohysterioides discovered from China.

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FIGURE 5. Roussoella pseudohysterioides (GMB0009). a–d. Ascostromata developing on bamboo culm. e, f. Vertical sections of
ascostromata. g–j. Asci containing eight ascospores. k. Fragment of ascostromata in KOH without stromatal pigments. l–m. Ascus apex in
Melzer’s reagent. n–r. Dark brown ascospores. Scale bars: j–r = 10 μm.

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Entomopathogenic fungi
Studying entomopathogenic fungi is one of the popular research areas in China since they are medicinal importance.
Beauveria, Cordyceps and Metarhizium are some important genera which have widely been studied. Here, we introduce
one new species and one new record of entomopathogenic fungi.
Tolypocladium W. Gams, Persoonia 6(2): 185 (1971)
Tolypocladium used to be known as an asexually genus since it was described (Gams 1971) until Hodge et al. (1996)
linked one sexual species to this genus. This genus was transferred in the family Ophiocordycipitaceae based on
phylogenetic analyses (Sung et al. 2007). Many species of Elaphocordyceps and Chaunopycnis have been transferred
to Tolypocladium, which was protected in the International Code of Nomenclature for algae, fungi, and plant (Kirk et
al. 2013, Quandt et al. 2014).
Tolypocladium cucullae Y.P. Xiao & T.C. Wen sp. nov. (FIGURE 6)
Index Fungorum Number: 558265
Etymology:—The specific epithet refers to the feature of the capitate stromata.
Holotype:—HKAS 55588
Parasitic in an unidentified host buried in the upper 1 cm of soil, forming brown to dark stromata. Sexual morph:
Ascomata 8–13 cm long, 5–10 mm wide, stromatic, brown to olive when fresh, dark when dry, tough, capitate, mostly
solitary, stipitate, inside hollow when mature. Stipe 8–12 × 0.5–0.7 cm, cylindrical, yellow to brown when fresh, dark
brown when dry, with green scales on the surface when fresh, with dark furfuraceous when dry, fibrous, hollow, with
stromata on the top. Fertile head 8-10 mm in diam, hemispherical, minutely mammilate, bracken green to dark olive
when fresh, dark when dry, distinctly separated from the stipe, tough, solitary, with a cortex of closely interwoven
hyaline hyphae pseudoparenchymatous in section. Perithecia 500–600 × 340–420 μm (
x
= 560 × 380 µm, n = 30),
subglobose to ovoid, immersed in stroma with slightly protruding ostiolar papilla. Ostiole lined with paraphyses.
Peridium 20–25 µm (
x
= 22 µm, n = 60) wide, of brown pigmented cells of textura porrecta to paler textura prismatica.
Asci 320–400 × 10–15 um (
x
= 360 × 13 µm, n = 60), 8-spored, unitunicate, narrow cylindrical, hyaline, with thick
apex. Apical cap 5.5–7.5 × 5–7.5μm (
x
= 6.5 × 6 µm, n = 60) μm diam, hyaline. Ascospores as long as asci, filiform,
hyaline break into secondary spores. Secondary spores 25–35 × 3–4.5 μm (
x
= 30 × 3.8 µm, n = 60), cylindrical to
fusoid with truncated ends, smooth, hyaline, with or without septa. Asexual morph: Undetermined.
Material examined:—CHINA, Yunnan Province, Lijiang City, Laojun Mountain. 15 July 2008, Yun Ting Huang
(HKAS 55588, holotype), (GZU A-77, isotype).
LSU: MW798786 MW7987877, SSU MW798784 MW798785, ITS MW798788 MW798789 (Supplementary
Table 1)
Notes:—We identified this species after we inspected the unidentified specimens in the Herbarium of Kunming
Institute of Botany, Chinese Academy of Sciences (HKAS). According to morphology and phylogenetic analysis
(Fig. 7), the new species Tolypocladium cucullae is close to T. capitatum, T. delicatistipitatum, T. fumosum and T.
longisegmentatum. Tolypocladium cucullae is distinct from T. capitatum by producing hollow, furfuraceous stipe,
smaller perithecia and smaller asci, while T. capitatum produces tough stipe, bigger perithecia and longer asci (Mains
1957, Table 3). Tolypocladium cucullae is distinct from T. fumosum in having larger and brown to olive when fresh,
dark when dry stromata; larger, hemispherical and bracken green to dark olive when fresh, dark when dry fertile
head; smaller perithecia; longer and cylindrical to fusoid secondary spores. Tolypocladium fumosum has smaller, pale
chalcedony yellow at the base to dark gull grey at the apex stromata; ellipsoidal when young and capitate when mature
fertile head; larger perithecia; shorter and cylindrical to cubic secondary spores. The phylogenetic tree also supports
that T. cucullae is distinct from T. capitatum and T. fumosum (Fig. 7).
The morpho-characters of T. cucullae are similar to T. delicatistipitatum, but the latter has no DNA sequence data.
Both of them formed stipitate stromata, subglobose to ovoid perithecia, cylindrical asci and cylindrical secondary
spores with truncate ends. Tolypocladium cucullae is different from Tolypocladium delicatistipitatum in producing
stromata with a hemispherical, dark (when dry) fertile part, with a thinner (5.5–6 μm in diam) apical cap and longer
(25–35 μm long) secondary spores, while T. delicatistipitatum produces stromata with a spherical or oval fertile part,
a thicker (8 μm in diam) apical cap and shorter (18–28 μm long) secondary ascospores.

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Molecular data have been supplemented by four strains, including OSC 110992 (Sung et al. 2007), HMJAU6903
(Yan & Bau 2014), MHHNU 8699 (Chen & Zhang 2019) and 2731.S (Stensrud et al. 2005). Furthermore, HMJAU6903
(Yan & Bau 2014) and MHHNU 8699 (Chen & Zhang 2019) were reported molecular data with descriptions and
illustrations among these four strains. Tolypocladium cucullae is distinct from T. longisegmentatum (DAOM 137162,
Ginns 1988; HMJAU6903, Yan & Bau 2014; MHHNU 8699, Chen & Zhang 2019) in having a hemispherical fertile
head, brown perithecia and shorter secondary spores (Table 3). Molecular data indicated that the new species has 31 bp
in ITS that differ from HMJAU 6903, 36 bp in ITS that is different from MHHNU 8699, 38 bp in ITS is different from
2731.S, 26 bp in LSU that are different from OSC 110992. In conclusion, we propose T. cucullae as a new species.
FIGURE 6. Tolypocladium cucullae (HKAS 55588, holotype). a, b. Material of Tolypocladium cucullae. c. Ascostromata. d. Fertile head
of ascostroma. e. Vertical section of stroma. f. Peridium. g–j. Asci. k. Apical cap of asci. l–o. Secondary ascospores. Scale Bars: d = 5 mm,
e = 100 µm, f = 50 µm, g–j = 200 µm, k, l–o = 20 µm.

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TABLE 3. Morphological comparison of closely related species of T. cucullae.
Strain Location Stromata Stipe (cm) Fertile head (mm) Perithecia (μm) Asci (μm) Secondary spores (μm) Reference
DAOM
137162 America Single,
rarely doulbe
13 × 0.7, greyish yellow
above and the lower
one-third a deep yellow,
cylindrical, olive, some with
the basal part dark olive or
black, glabrous, hollow
Broadly rounded, above
the stipe apex, brown,
dark brown to olive
brown, glabrous, 13 diam
Imbedded, ellipsoid
500 × 300
Cylindrical to
narrowly ellipsoid
440 × 10–15
(12–)40–65 × 3–5 Ginns (1988)
HMJAU6903 China Single
2.5–6.5 × 0.2 wide, yellow
brown, with black scales on
the surface,
Broadly rounded, dark
brown, 4–9 diam
Imbedded, ellipsoid or
flask-shaped, 632–681 ×
273–292
Cylindrical,
341–428 × 12
(12.2–) 29.3–48.8
(–73.3) × 3.7–4.9 Yan & Bau (2014)
HKAS 55588 China Single
8–12 × 0.5–0.7, cylindrical,
yellow to brown when fresh,
dark when dry, with green
scales on the surface, hollow
Hemispherical, bracken
green to dark green when
fresh, dark when dry, 8-10
in diam
Immersed, subglobose to
ovoid, 500–600 × 340–420
Cylindrical to
fusoid 320–400 ×
10–15
25–35 × 3–4.5 This study

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FIGURE 7. Phylogram of Tolypocladium generated from Maximum likelihood analysis of ITS, SSU and LSU sequence data.
Purpureocillium lilacinum (CBS 284.36) was selected as an outgroup taxon. The tree topology of the ML analysis was similar to the BI.
Maximum likelihood bootstrap values greater than 75 and Bayesian posterior probabilities over 0.90 were indicated above the nodes. The
scale bar indicates 0.006 changes. The new species was in blue.
Metarhizium guizhouense Q.T. Chen & H.L. Guo, Acta Mycol. Sin. 5(3): 181 (1986) (FIGURE. 8)
Index Fungorum Number: 130206
Specimen found on stick insects (Phasmatodea). Host’s internodes between abdominal segments were covered with
white to pale green mycelium and sporulating conidiophores. Conidiophores arising from hyphae, smooth-walled.
Phialides cylindrical, solitary, smooth-walled, 8–18 × 1–1.5 μm. Conidia smooth-walled, pale green to colorless (6.5–
9.5 × 2.5–3 μm), cylindrical, slightly constricted in the middle, round at both ends or tapered at one end. Bi-celled
conidium was not observed.
Culture characteristics:—Colonies on PDA were relatively slow-growing, fluffy, beginning to white, and the
spores appear green, started to produce conidia after 3 days in culture at 25 °C in the laboratory, 17 mm diam. after 10
days. Mature conidia chains are often spread on the surface of the colony in small granular clumps. Hyphae hyaline,
separated, branched, about 3 um wide.
Material examined:—China, Guizhou Province, Guiyang, on dead stick insects, July 2019, Q.R. L, 2019GY03
(GMB0010), living cultures, GMBC0010 (new host record).
Known hosts and distribution:—Guizhou
Known hosts:—larvae of Noctuidae sp., stick insects
GenBank Numbers:—ITS: MW881444, LSU: MW881450, RPB2: MW883344
Note:—Metarhizium guizhouense, isolated on Hepialus sp. in Guizhou China, was introduced by Guo et al.
(1986). In 1991, Liang et al. reported a M. taii Z.Q. Liang & A.Y. Liu on larvae of Noctuidae sp. (Lepidoptera).
Metacordyceps taii was recognized to be the sexual morph of M. guizhouense by Bischoff et al. (2009). Qu et al. also

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reported that M. taii should be treated as a synonym of M. guizhouense based on molecular data. This is the first report
of M. guizhouense isolated on stick insects (Phasmatodea).
FIGURE 8. Metarhizium guizhouense (GMB0010) (new host record). a, b. Fungus on stick insects (Phasmatodea) c, d. Green mycelium
and sporulating conidiophores covered on the surface of inscect. e, f, g. Conidiophores h, i. Conidia on insect host. Scale bars: a, b = 5 mm,
c = 2 mm, d = 500 μm, j–r = 10 μm, e–i = 5μm
Fresh water fungi
In Yungui region, freshwater fungi are mainly found in lotic (rivers, streams and waterfalls) habitats. They play
important ecological roles since they are decomposers of submerged substrates (mainly from riparian vegetation), and
participate crucial biogeochemical cycles, such as carbon cycling (Wurzbacher et al. 2010; Gulis & Barlocher 2017).
In this study we provide a new country record of Myrmecridium schulzeri.
Myrmecridium schulzeri (Sacc.) Arzanlou, W. Gams & Crous, Stud. Mycol. 58: 84 (2007) (FIGURE. 9)
Index Fungorum Number: IF504560
Saprobic on submerged decaying wood. Sexual morph undetermined. Asexual morph Colonies on natural substrata
effuse, superficial, scattered, hairy, solitary or in small groups, black, with a mass of visible whitish to grayish conidia on
middle to upper part of conidiophores. Mycelium partly superficial, partly immersed. Conidiophores macronematous,
mononematous, straight to slightly curve, unbranched, medium brown to brown at base part, pale towards top part,
thin-walled, septate, 172–304 × 2–3 μm ( = 212 × 2.6 μm, n = 15). Conidiogenous cells holoblastic, polyblastic,

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integrated, terminal and intercalary, cylindrical, subhyaline to pale brown, forming a rachis with scattered pimple-
shaped denticles which are less than 1 µm long and approx. 0.5 µm in diameter. Conidia solitary, fusoid or ellipsoidal
to obovoidal, rounded at the apex, obtuse and tapering towards base, hyaline, aseptate, thin-walled, smooth, without
guttule, some with a small protuberance, 5–6.5 × 2.3–3.6 μm ( = 5.8 × 2.9 μm, n = 35).
FIGURE 9. Myrmecridium schulzeri (IFRD500–012) a, b. Colonies on natural substrate. c–e. Conidiophores with conidia. f, g.
Conidiogenous cells with conidia. h–l. Conidia. m Germinating conidia on PDA. n, o. Culture on PDA, n. from front, o. from reverse.
Scale bars: c–e, h = 20 μm, m = 10 μm, f, g = 5 μm, i–l = 2 μm.

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Culture characteristics:—Conidia germinating on PDA within 24h. Colonies grow on PDA attaining 38–48 mm
diameter in 40d at 20–25°C in the condition of 12h-dark and 12h-light, with smooth, floccose, pale brown mycelium
on the surface, reverse white, with filamentous, undulate margin.
Material examined: China, Yunnan Province, small river of Puzhehei, on dead submerged decaying wood of
unidentified plants, 23 June 2018, Hao Yang, P37 (IFRD500–012), living culture = KUMCC 20–0190 (new record
from Yunnan, new habitat record).
Known hosts and distribution: Soil (Germany, Papua New Guinea, Zaire), Homo sapiens (Netherlands), Wheat
straw (South Africa), Triticum aestivum (Netherlands), Malus sylvestris (Switzerland), Cannomois virgate (South
Africa)
GenBank Numbers: ITS MT559103
Notes:—Myrmecridium was introduced by Arzanlou et al. (2007) with M. schulzeri as type species, which
was described as Chloridium schulzerii (Sacc.) Sacc. and Rhinocladiella schulzeri (Sacc.) Matsush. Our isolate fits
the characters of Myrmecridium well in having macronematous, unbranched, septate conidiophores, polyblastic
conidiogenous cells with denticles, and hyaline, thin-walled, smooth, fusoid or ellipsoidal to obovoidal conidia
(Arzanlou et al. 2007, Jie et al. 2013, Peintner et al. 2016, Réblová et al. 2016). The sequence data in ITS gene region
of our isolate are identical to that of M. schulzeri. Thus, we identified our isolate as M. schulzeri. Our isolate is a new
geographic record in China and a new habitat record from freshwater.
Mushrooms
Panus similis Berk. & Br. In Journ. Linn. Soc., Bot. 14:43 (1873) (FIGURES 10,11)
Pileus (4.9B) 4–16 cm diameter, thin, deeply infundibuliform; surface brown to dark chestnut brown, finely velutinate
at the centre, radially plicate-sulcate with the striae extending almost to the centre, margin curved downwards, ciliate.
Lamellae decurrent, ochraceous buff, darkening at maturity, 1.5–3 mm broad, moderately spaced with lamellulae of
five lengths; entire edge. Stipe central, 4–17 cm × 1.5–2 mm, solid, cylindric, slightly expanded at the base; surface
concolorous with the pileus, uniformly velutinate and felt-like. Context 1–2 mm thick at the centre, coriaceous,
white. Generative hyphae (4.7E) 2–4 μm diameter, very thin-walled, frequently branching with clamp connections.
Skeletal hyphae (4.7E) 2–5 μm diameter, cylindric, sinuous with a thickened hyaline wall, unbranched. Basidiospores
(4.7A) (5.5–6.5 × 2.5–3.5 (5.5 ± 0.3 × 3 ± 0.2) μm, Q =1.83, hyaline, ellipsoid to oblong cylindric, thin-walled, with
few contents. Basidia 17–29 × 4–5 μm, clavate, cylindric, bearing 4 sterigmata. Lamella-edge sterile, with small
Cheilocystidia, soon collapsing. Cheilocystidia crowded, 17–26 × 3–6 μm, nodulose-clavate, hyaline, irregular, thin-
walled. Sclerocystidia (4.7D) very abundant, very crowded, 19–41 × 4–9 μm, irregularly fusoid, elongate, with a thick,
hyaline wall. Hymenophoral trama irregular of radiate construction, hyaline. Subhymenial layer slightly developed.
Pileipellis on epicutis, up to 115 μm thick, of more or less repent hyaline, up to 160 μm long, 115 μm diameter, with a
thickened wall of 1.5–3.5 μm. Stipitipellis similar to Pileipellis. Smell mushroomy, edible when it is young.
Material examined:—CHINA, Yunnan Province, Xishuangbanna, elevation 400 m, rainforest dominated by
Castanopsis sp. and Dipterocarpus sp.; 4 June 2018, Samantha C. Karunarathna (HKAS 121668) (new country
record).
Notes: Panus similis has a palaeotropical Distribution and is most commonly found in south-east Asia and
Australasia, but also extends westwards across equatorial Africa. It is recognized by the excellently velutinate to
glabrescent pileus with noticeable radially sulcate striate, combined with the subdistant lamellae. Large basidiocarps
are frequently encountered almost always associated with a prominent pseudosclerotium. This study reports P. similis
for the first time from China, based on both morphological characteristics (Figs. 10, 11) and phylogenetic analysis (Fig.
12).
Discussion
Why Yungui Plateau is important in Chinese mycology
Hawksworth & Lucking (2017), and Hyde et al. (2020) identified biodiversity-rich areas for revealing undiscovered
or missing fungal species. Hence, Yungui Plateau is an important region to conduct intensive research to discover new

WIJAYAWARDENE ET AL.
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fungal taxa. Figure 13 shows that Yunnan and Guizhou Provinces are the leading provinces in introducing new species
in China.
FIGURE 10. Panus similis a. Basidiospores, b. Basidia, c. Cheilocystidia, d. Sclerocystidia, e. Generative hyphae and Skeletal Hyphae,
f. Hairs on pileus. Scale bars: b, c, d = 20 µm; a, e, f = 10 µm

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FIGURE 11. Basidiocarps of Panus similis (HKAS 121668) in the field.
Species prediction: host-fungi ratio, insect-fungi ratio?
Predicting species number in the Kingdom Fungi is a topic of considerable controversy. Some studies predicted
global species number (e.g., Blackwell 2011, Tedersoo et al. 2014, Hawksworth & Lücking 2017), while some studies
predicted the species number in particular geographical regions (e.g., Crous et al. 2006 in South Africa; Dai & Zhuang
2010 in China). However, different studies have used different techniques for calculation, leading to divergent species
numbers. The most recent study carried out using high-throughput sequencing revealed 6.28 million fungal species
globally (Baldrian et al. 2021).
Hawksworth (1991, 2001) assumed the plant:fungi ratio as 1:6; thus, based on this ratio, Feng & Yang (2018)
predicted 104,000 fungal species should be present in Yunnan (number of vascular plants: 17,427 species *6 =
ca. 104,000). However, Hawksworth & Lücking (2017) regarded 1:8 as a more accurate plant:fungi ratio, raising
the estimation of fungal species to 139,416. However, it is also very important to consider the insect:fungi ratio.
Environmental sequencing enhances species number as expected in Tedersoo et al. (2014). Nevertheless, Lücking &
Hawksworth (2018) mentioned that these ratios might underestimate when tropical regions are taken in to account.
Hyde et al. (2018) has also suggested ‘that a large proportion of new species awaits discovery and possibly lie in
tropical regions such as Thailand’. Hence, it is extremely likely that many more fungal species exist in Yungui Plateau
awaiting discovery and description.

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FIGURE 12. Phylogram of Panus generated from Maximum likelihood analysis of ITS sequence data. Lentinus crinitus (MK408650)
was selected as the outgroup taxon. Maximum likelihood bootstrap values greater than 60% are indicated above the nodes. The new record
Panus similis (HKAS 121668) is in black bold.
FIGURE 13. Number of species introduced from different provinces in China over the last decade (2010–2020).

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Future work
It is necessary to recognize important recent changes in mycology, since outdated techniques, methodologies and
literature have resulted in inaccurate calculations and conclusions. Changes in nomenclature (Hawksworth 2012; May
et al. 2019; Wijayawardene et al. 2021), fungal barcoding (Schoch et al. 2012), genes for precise pathogenic species
identification and environmental sequencing to identify unculturable taxa (Wu et al. 2019) are some important landmark
developments in the last decade and comprise the foundational bedrock for future mycological work. Active mycology
research groups in Yungui Plateau are recognized as 1) those who work on only macro fungi (i.e., mushrooms, other
macrofungi and mushroom domestication); 2) those who work only on micro fungi; and 3) those who work on both
micro and macro fungi. In the future, collaboration between these groups and overseas institutions will be essential to
overcome limitations in data and funding.
We list potential research areas below that could be explored in the future.
Micro fungi
1. Looking for species in less-studied life modes, e.g., lichenicolous, species on rocks and karst regions (including artefacts).
2. Epitypification of old species on the Yungui Plateau.
3. Resolving species complexes of important phytopathogens using barcoding.
4. Screening for secondary metabolites of endophytic species.
5. Genomic studies of agriculturally and industrially important species.
Mushrooms
1. Domestication of new wild edible and medicinal mushrooms.
2. Checklists and guidebooks of edible, medicinal and poisonous mushrooms.
Acknowledgments
This work was supported by the National Natural Science Foundation of China (No. NSFC 31950410558, NSFC
31760013, NSFC 31960005, NSFC 32000009), Department of Science and Technology of Yunnan Province (No.
2018FB050), the State Key Laboratory of Functions and Applications of Medicinal Plants, Guizhou Medicial
University (No. FAMP201906K); Science and Technology Department of Guizhou Province (QKHRCPT[2017]5101)
and High-Level Talent Recruitment Plan of Yunnan Provinces ("Young Talents" Program and "High-End Foreign
Experts" Program). Nalin N. Wijayawardene would like to thank Dr. Josiane S. Monteiro for providing statistics.
Ting-Chi Wen would like to thank National Natural Science Foundation of China (No. 31760014), and the Science
and Technology Foundations of Guizhou Province (No. KY[2018]039). Chun Y. Deng thanks the Guizhou Province
Science and Technology Support Program, China ([2019]2451). Yingqian Kang would like to thank Guizhou Scientific
Plan Project [qiankehe Support (2020) 4Y220] and National Natural Science Foundation of China ［32060034; 111
Plan of China [(2019) 475]. Austin Smith in World Agroforestry Center, Kunming Institute of Botany is thanked
for English correction in the manuscript. Saowaluck Tibpromma would like to thank the International Postdoctoral
Exchange Fellowship Program (number Y9180822S1), CAS President’s International Fellowship Initiative (PIFI)
(number 2020PC0009), China Postdoctoral Science Foundation, and the Yunnan Human Resources and Social Security
Department Foundation for funding her postdoctoral research. Samantha C. Karunarathna thanks CAS President’s
International Fellowship Initiative (PIFI) for funding his postdoctoral research (number 2018PC0006) and the National
Science Foundation of China (NSFC) for funding this research work under project code 31750110478.
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Supplementary Table 1. Sources of isolates and GenBank accession numbers for the genus Tolypocladium and
Mucispora.
Name Voucher GenBank Accession no. References
SSU ITS LSU
Adelosphaeria catenata CBS 138679 KT278692 KT278707 KT278707 Réblová et al. (2016)
Ascotaiwania fusiformis MFLUCC 15-0621 KX550898 KX550893 KX550893 Yang et al. (2016)
A. fusiformis MFLUCC 15-0625 KX550898 KX550894 KX550894 Yang et al. (2016)
A. mitriformis HKUCC 3706 NA AF132324 AF132324 Ranghoo et al. (1999)
A. sawadae SS 00051 HQ446283 HQ446363 HQ446363 Boonyuen et al. (2011)
Bactrodesmiastrum monilioides FMR 10756 NA KF771879 KF771879 Hernández-Restrepo et al.
(2015)
B. obovatum FMR 6482 NA FR870266 FR870266 Hernández-Restrepo et al.
(2013)
B. pyriforme FMR 10747 NA FR870265 FR870265 Hernández-Restrepo et al.
(2013)
B. pyriforme FMR 11931 NA HE646637 HE646637 Hernández-Restrepo et al.
(2013)
Brachysporiella setosa HKUCC 3713 NA AF132334 AF132334 Ranghoo et al. (1999)
Canalisporium exiguum SS 00809 GQ390266 GQ390281 GQ390281 Sri-indrasutdhi et al. (2010)
C. grenadoideum BCC 20507 GQ390252 GQ390267 GQ390267 Sri-indrasutdhi et al. (2010)
C. pulchrum SS 03982 GQ390262 GQ390277 GQ390277 Sri-indrasutdhi et al. (2010)
Conioscypha lignicola CBS 335.93 JQ437439 AY484513 AY484513 Réblová & Seifert (2004)
C. minutispora FMR 11245 HF937347 KF924559 KF924559 Crous et al. (2014)
C. peruviana ILL 41202 NA KF781539 KF781539 Zelski et al. (2015)
C. varia CBS 113653 AY484511 AY484512 AY484512 Réblová & Seifert (2004)
Fuscosporella aquatica MFLUCC 16-0859 NG_062433 MG388209 MG388209 This study
F. pyriformis MFLUCC 16-0570 KX550900 KX550896 KX550896 Yang et al. (2016)
Helicoön farinosum DAOM 241947 NA JQ429230 JQ429230 Réblová et al. (2012)
Leotia lubrica AFTOL-ID 1 AY544746 AY544644 AY544644 Lutzoni et al. (2004)
Melanotrigonum ovale CBS 138742 KT278695 KT278708 KT278708 Réblová et al. (2016)
M. ovale CBS 138743 KT278696 KT278709 KT278709 Réblová et al. (2016)
Microglossum rufum AFTOL-ID 1292 DQ471033 DQ470981 DQ470981 Spatafora et al. (2006)
Mucispora hydei GMB0028 MW800164 MW797039 MW797122 This study
M. infundibulata MFLU 18-0142 NG_073505 MH457139 MH457139 Hyde et al. (2020)
M. obscuriseptata MFLUCC 15-0618 KX550897 KX550892 KX550892 Yang et al. (2016)
M. phangngaensis MFLUCC 16-0865 MG388207 MG388210 MG388210 This study
Parafuscosporella aquatica MFLU 19-0550 NA MN512343 MN512343 Yang et al. (2020)
P. garethii FF00725.01 KX958428 KX958430 KX958430 Boonyuen et al. (2016)
P. pyriformis MFLU 19-0525 NA MN512339 MN512339 Yang et al. (2020)
P. moniliformis MFLUCC 15-0626 KX550899 KX550895 KX550895 Yang et al. (2016)
P. mucosa MFLUCC 16-0571 MG388208 MG388211 MG388211 This study
Phaeoisaria fasciculata CBS 127885 KT278693 KT278705 KT278705 Réblová et al. (2016)
P. fasciculata DAOM 230055 KT278694 KT278706 KT278706 Réblová et al. (2016)
P. microspora MFLUCC 16-0033 NA MF167351 MF167351 Hyde et al. (2017)
P. sedimenticola CGMCC 3.14949 NA JQ031561 JQ031561 Cheng et al. (2014)
Phragmocephala stemphylioides DAOM 673211 NA KT278717 KT278717 Réblová et al. (2016)
Plagiascoma frondosum CBS 139031 KT278701 KT278713 KT278713 Réblová et al. (2016)
Pleurotheciella centenaria DAOM 229631 JQ429246 JQ429234 JQ429234 Réblová et al. (2012)
P. rivularia CBS 125238 JQ429244 JQ429232 JQ429232 Réblová et al. (2012)
P. rivularia CBS 125237 JQ429245 JQ429233 JQ429233 Réblová et al. (2012)
P. uniseptata DAOM 673210 NA KT278716 KT278716 Réblová et al. (2012)
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Supplementary Table 1. (Continued)
Name Voucher GenBank Accession no. References
SSU ITS LSU
Pleurothecium floriforme MFLUCC 15-0628 KY697279 KY697277 KY697277 Hyde et al. (2017)
P. recurvatum CBS 131272 JQ429251 JQ429237 JQ429237 Réblová et al. (2012)
P. recurvatum CBS 138747 KT278703 KT278714 KT278714 Réblová et al. (2016)
P. semifecundum CBS 131271 JQ429254 JQ429240 JQ429240 Réblová et al. (2012)
P. semifecundum CBS 131482 JQ429253 JQ429239 JQ429239 Réblová et al. (2012)
Pseudoascotaiwania persoonii A57-14C NA AY094190 AY094190 Campbell & Shearer (2004)
P. persoonii A57-14C NA AY590295 AY590295 Campbell & Shearer (2004)
Purpureocillium lilacinum CBS 284.36 AY489689 NR_111432 NA Luangsa-Ard et al. (2004)
Savoryella longispora SAT 00322 HQ446302 HQ446380 HQ446380 Boonyuen et al. (2011)
S. paucispora SAT 00866 HQ446303 HQ446381 HQ446381 Boonyuen et al. (2011)
S. verrucosa SS 00052 HQ446296 HQ446374 HQ446374 Boonyuen et al. (2011)
Sterigmatobotrys macrocarpa PRM 915682 NA GU017317 GU017317 Réblová and Seifert (2011),
Réblová et al. (2012)
S. macrocarpa DAOM 230059 = CBS 113468 NA GU017316 GU017316 Réblová & Seifert (2011),
Réblová et al. (2012)
Taeniolella rudis DAOM 229838 JQ429256 JQ429241 JQ429241 Réblová et al. (2012)
Tolypocladium album CBS 869.73 KF747309 NR_155018 NA Gazis et al. (2014)
T. album CBS 393.89 NA MH862176 MH873866 Vu et al. (2019)
T. album CBS 968.73B KF747314 MH860832 MH872567 Vu et al. (2019)
T. album CBS 830.73 NG_065021 MH860811 MH872543 Vu et al. (2019)
T. album GB5123 NA AF389191 AF245296 Bills et al. (2002)
T. album GB5502 AY489689 AF389192 AF245297 Bills et al. (2002)
T. amazonense LA100 MW798784 HQ022485 KF747129 Gazis et al. (2014)
T. amazonense LA108 MW798785 HQ022486 KF747130 Gazis et al. (2014)
T. amazonense MS308 NA JQ905653 KF747134 Gazis et al. (2014)
T. amazonense BPI 892889 NA NA NA Gazis et al. (2014)
T. capitatum FLAS-F-60359 NA MF074845 NA Montalva et al. (2019)
T. capitatum OSC 71233 AF049153 NA AY489721 Quandt et al. (2014)
T. cucullae HKAS 55588 NA MW798788 MW798786 This study
T. cucullae GZU A-77 NA MW798789 MW798787 This study
T. cylindrosporum ARSEF 2920 KF747323 MG228381 NA Montalva et al. (2019)
T. cylindrosporum IP 425 KF747321 MG228380 NA Montalva et al. (2019)
T. cylindrosporum IP 419 DQ522545 MG228379 NA Montalva et al. (2019)
T. cylindrosporum NRRL 28025 NA NA AF049173 Quandt et al. (2014)
T. dujiaolongae RCEF6201 NA KF696558 NA Li et al. (2018)
T. dujiaolongae ZBAH632 NA KF696557 NA Li et al. (2018)
T. endophyticum MX575 NA JX155949 KF747155 Gazis et al. (2014)
T. endophyticum MX486 NA KF747245 KF747152 Gazis et al. (2014)
T. fractum OSC 110990 AB027322 NA DQ518759 Sung et al. (2007)
T. fumosum WA18945 EF469124 KU925171 KU985053 Crous et al. (2017)
T. geodes ARSEF 2684 NA FJ973059 NA Ghikas et al. (2010)
T. geodes CBS 723.70 NA NR_164431 NA Vu et al. (2019)
T. guangdongensis GDGM 24020 NA EU039881 NA Ke & Ju (2015)
T. inegoense SU-15 NA NA DQ118741 Chaverri et al. (2005)
T. inegoense DQ522547 NA AB027368 Nikoh & Fukatsu (2000)
T. inflatum OSC 71235 NA JN049844 EF469077 Sung et al. (2007)
T. inflatum NBRC 31668 AB027320 AB103381 NA Yokoyama et al. (2004)
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Supplementary Table 1. (Continued)
Name Voucher GenBank Accession no. References
SSU ITS LSU
T. inflatum CBS 714.70 AB027319 MH859916 MH871710 Vu et al. (2019)
T. inflatum CBS 127302 NA MH864514 MH875949 Vu et al. (2019)
T. inflatum CBS 127147 NA MH864439 MH875880 Vu et al. (2019)
T. japonicum OSC 110991 NA JN049824 DQ518761 Kepler et al. (2012)
T. japonicum BCRC FU30561 NA KT873533 NA Ke & Ju (2015)
T. japonicum IFO 9647 NA AB027366 NA Nikoh & Fukatsu (2000)
T. jezoense NA NA AB027365 Nikoh & Fukatsu (2000)
T. longisegmentatum OSC 110992 NA NA EF468816 Sung et al. (2007)
T. longisegmentatum 2731.S AY489691 AJ786568 NA Stensrud et al. (2005)
T. longisegmentatum MHHNU 8699 KJ878910 MK253762 NA Chen & Zhang (2019)
T. longisegmentum HMJAU6903 NA KJ866879 NA Yan & Tolgor (2014)
T. nubicola ARSEF 3434 NA FJ973067 NA Ghikas et al. (2010)
T. nubicola CBS 944.72 NA NA MH878304 Vu et al. (2019)
T. nubicola CBS 568.84 NA NA MH873478 Vu et al. (2019)
T. ophioglossoides OSC 106405 JN941730 NA AY489723 Castlebury et al. (2004)
T. ophioglossoides CBS 100239 AB027323 KU382155 KJ878874 Quandt et al. (2014)
T. ovalisporum CBS 700.92 NA NR_155019 NA Unpublished
T. paradoxum JFL14081002 MF5368LR KX017278 NA Zha et al. (2018)
T. paradoxum HKAS 87772 NA KX017279 NA Zha et al. (2018)
T. paradoxum HMG 20938 NA DQ901630 NA Tian et al. (2010)
T. paradoxum NBRC 106958 NA JN943324 JN941411 Schoch et al. (2012)
T. paradoxum MX338 KF747318 AB027369 AB027369 Nikoh & Fukatsu (2000)
T. pustulatum MRL GB6597 KF747303 NA AF389190 Bills et al. (2002)
T. pustulatum MRL MF5368LR NA MF5368LR AF373282 Bills et al. (2002)
T. pustulatum KaP8.2.2.1 NA KP698195 NA Arhipova et al. (2015)
T. tropicale MX337 NA JQ905660 KF747148 Gazis et al. (2014)
T. tropicale IQ214 NA KF747254 NA Gazis et al. (2014)
T. tropicale MX338 NG_061025 KF747259 KF747149 Gazis et al. (2014)
T. tropicale IQ136 KF747309 NA KF747121 Gazis et al. (2014)
T. tropicale CBS 136897 NA NR_159005 NA Gazis et al. (2014)
T. tundrense ARSEF 3400 KF747314 FJ973069 NA Ghikas et al. (2010)
T. tundrense CBS 569.84 NG_065021 MH861781 MH873479 Vu et al. (2019)
T. varium CBS 429.94 NA MH862472 MH874122 Vu et al. (2019)
NA: Sequences not available.