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Research Article
Two new species of Craterellus (Cantharellales, Hydnaceae) with veined hymenophore from north-eastern China
expand article infoGui-Ping Zhao, Jia-Jun Hu, Yong-Lan Tuo, Yu Li, Bo Zhang
‡ Jilin Agricultural University, Changchun, China
Open Access

Abstract

In this contribution to the genus Craterellus in northern China, two new species are introduced: Craterellus connatus and C. striatus. These species and C. atrobrunneolus, initially described in south-western China, are highly similar and closely related. The species delimitation is molecularly supported by multigene phylogenetic analysis of the nr LSU and tef-1α region. Craterellus connatus is characterised by its medium-sized basidiomata, greyish-brown and smooth pileus with an off-white margin, the hymenophore with a strongly anastomosing vein, turning khaki upon drying, connate stipe, broad ellipsoid to ellipsoid basidiospores (6.1–7.8 × 4.8–5.9 μm), slender basidia with (2)4–6 sterigmata and the absence of clamp connection. Craterellus striatus is characterised by its small-sized basidiomata, fibrillose, greyish-brown to yellowish-brown, fully perforated pileus with a brown fringe, the hymenophore with a forking vein, the stipe inflated at the base, broad ellipsoid to ellipsoid basidiospores (6.8–8.0 × 5.1–6.0 μm), 2–6 spored basidia, encrusted hyphae and the absence of clamp connection. Detailed macroscopic and microscopic descriptions, accompanied by illustrations and a taxonomic discussion, are presented. A key to the Chinese Craterellus species is also provided.

Keywords

Chinese species, molecular phylogeny, morphology

Introduction

Craterellus Pers., typified with C. cornucopioides (L.) Pers. (Persoon 1825), belongs to Hydnaceae of Cantharellales (Hibbett et al. 2014). It is well represented in northern temperate zones and occurs in the tropics (Dahlman et al. 2000). Species in this genus are characterised by funnel-shaped fruiting bodies, hollow stipes, usually dark grey to black or yellow pileus (Corner 1966). Craterellus is an important genus of wild edible mushrooms and is renowned for its high economic, medicinal and ecological values (Pilz 2003). Approximately 162 records of Craterellus (including infraspecific taxa) have been found in Index Fungorum (http://www.indexfungorum.org). However, most of these species have been transferred to other genera, based on a phylogenetic analysis of the large nuclear subunit (nr LSU) and the internal transcribed spacer region (ITS), including Cantharellus Adans. ex Fr., Gomphus Pers. and Polyozellus Murrill etc. (Dahlman et al. 2000; Contu et al. 2009; Olariaga et al. 2009; Wilson et al. 2012; Montoya et al. 2021). Nearly 100 names have been considered legitimate in MycoBank (https://www.mycobank.org/) to date.

In China, species diversity, taxonomy and phylogeny of macrofungi have been investigated in recent years, many new species having been discovered (Zhong et al. 2018; Cui et al. 2019; Sun et al. 2020; Zhang et al. 2020a; Cao et al. 2021a, b; Liu et al. 2021, 2022a, b; Ji et al. 2022; Sun et al. 2022). Craterellus is one of the important fungal genera of macrofungi, 14 species of Craterellus having been recorded in China: C. albidus Chun Y. Deng, M. Zhang & J. Zhang, C. atratus (Corner) Yomyart, Watling, Phosri, Piap. & Sihan., C. atrobrunneolus T. Cao & H.S. Yuan, C. aureus Berk. & M.A. Curtis, C. badiogriseus T. Cao & H.S. Yuan, C. cornucopioides, C. croceialbus T. Cao & H.S. Yuan, C. lutescens (Fr.) Fr., C. luteus T.H. Li & X.R. Zhong, C. macrosporus T. Cao & H.S. Yuan, C. odoratus (Schwein.) Fr., C. sinuosus (Fr.) Fr., C. squamatus T. Cao & H.S. Yuan, C. tubaeformis (Fr.) Quél (Bi 1994; Mao 1998; Dai et al. 2010; Gu et al. 2012; Li et al. 2015; Zhong et al. 2018; Zhang et al. 2020a; Cao et al. 2021a, b). Nevertheless, due to their less attractive appearance, which can be confused with the environment, it is challenging to find Craterellus species in the field, causing them to be overlooked and thus remain undescribed. In addition, some species of Craterellus, especially those with the non-perforated pileus, are easily confused with other morphologically similar taxa, for example,, some species of Clitocybe (Fr.) Staude, Gomphus or Polyozellus, making it never easily distinguished from others in fieldwork.

During a recent survey of macrofungi in northern China, we discovered some exciting and novel species of Craterellus. In this contribution, the subsequent morphological and molecular analyses of the transcription elongation factor 1-alpha (tef-1α) and nr LSU sequences represented two new species, C. connatus G. P. Zhao, J. J. Hu, B. Zhang & Y. Li and C. striatus G. P. Zhao, J. J. Hu, B. Zhang & Y. Li which are described and illustrated herein. A key of Craterellus in China is provided as well.

Materials and methods

Vouchers and morphological analyses

This paper is principally based on materials collected by the senior author and collaborators over the past four years in northern China and specimens have been deposited at the Mycological Herbarium of Jilin Agricultural University (HMJAU).

Photographs and descriptions of macroscopic morphological characteristics were made from fresh materials in the field. The colours correspond to the “Flora of British fungi: colour identification chart” (Royal Botanic Garden 1969). Collections were dried in an oven at 45 °C and rehydrated with 95% alcohol. All microscopic observations and measurements were made in ammoniacal Congo red with a 5% aqueous potassium hydroxide (KOH) solution to improve tissue dissociation and matrix dissolution. Melzer’s Reagent was also used in the study. Measurements of basidiospores cite length and length/width ratio (Q) in this format: (minimum–) mean minus standard deviation–mean value–mean plus standard deviation (–maximum measured), Qm = average Q of all basidiospores measured ± sample standard deviation; spore measurements are based on 20 spores. Microscopic features were examined with the aid of the ZEISS Axio Lab A1.

DNA extraction, amplification and sequencing

Genomic DNA was extracted from dried or fresh material stored in desiccant silica gel. The extraction method followed the nuclear Plant Genomic DNA Kit (Kangwei Century Biotechnology Company Limited, Beijing, China). The amplified segments were the nr LSU and the tef-1α regions. The nr LSU region was amplified using LR0R and LR5 (Vilgalys and Hester 1990). The tef-1α region was amplified using tefF and tefR (Morehouse et al. 2003). Polymerase chain reaction (PCR) amplifications were performed in a total volume of 25 μl containing 1 μl template DNA, 1 μl of each primer, 10.5 μl distilled water and 12.5 μl PCR mix (2× Es Taq MasterMix, CWBIO, China). PCR amplification conditions followed Vilgalys and Hester (1990) for the nr LSU region and Morehouse et al. (2003) for the tef-1α region. The PCR products were subjected to electrophoresis on 1% agarose gel. Sequencing was performed by Sangon Biotech (Shanghai, China) using the same primer pairs used for the PCR.

Phylogenetic analyses

This study is based on around 25 specimens, including two outgroups (Hydnum ellipsosporum Ostrow & Beenken and Cantharellus cibarius Fr.) (Bijeesh et al. 2018; Zhong et al. 2018; Zhang et al. 2020a; Cao et al. 2021b). Sequences of the nr LSU and tef-1α regions were newly produced in this study and downloaded from GenBank (http://www.ncbi.nlm.nih.gov/). Detailed sample information is provided in Table 1.

Table 1.

Information on the specimens that were used in the phylogenetic analyses. Sequences that were newly generated in this study are indicated in black bold. T: Type.

Taxon Voucher number Country GenBank accession number References
nr LSU tef-1α
Craterellus albidus HGASMF013581 (T) China MT921161 Zhang et al. (2020a)
C. albidus HGASMF0110046 China MT921162 Zhang et al. (2020a)
C. albostrigosus CAL1624 (T) India MG593194 Bijeesh et al. (2018)
C. atratoides TH9232 (T) Guyana NG042660 Wilson et al. (2012)
C. atratoides MCA1313 Guyana JQ915119 Wilson et al. (2012)
C. atratus MCA1070 Guyana JQ915118 Wilson et al. (2012)
C. atratus TH9203 Guyana JQ915133 Wilson et al. (2012)
C. atrobrunneolus Yuan13878 (T) China MN894058 Cao et al. (2021b)
C. badiogriseus Yuan 14776 (T) China MW979532 MW999432 Cao et al. (2021a)
C. badiogriseus Yuan 14779 China MW979533 MW999433 Cao et al. (2021a)
C. caeruleofuscus MH17001 USA MT237468 Genbank
C. cinereofimbriatus TH9075 (T) Guyana JQ915131 Wilson et al. (2012)
C. cinereofimbriatus TH8999 Guyana JQ915130 Wilson et al. (2012)
C. cinereofimbriatus TH9264 Guyana JQ915138 Wilson et al. (2012)
C. connatus HMJAU 61462 (T) China OM509448 ON125915 This study
C. connatus HMJAU 61462 China ON125916 This study
C. cornucopioides HbO53302 Norway AF105301 Dahlman et al. (2000)
C. cornucopioides UPSF11801 USA AF105299 Dahlman et al. (2000)
C. excelsus MCA3107 Guyana JQ915121 Wilson et al. (2012)
C. excelsus TH7515 Guyana JQ915127 Wilson et al. (2012)
C. fallax AFTOL286 USA AY700188 Genbank
C. ignicolor UPSF11794 USA AF105314 Dahlman et al. (2000)
C. indicus PUN3884 (T) India HM113529 Kumari et al. (2012)
C. indicus MSR7 India HQ450770 Kumari et al. (2012)
C. indicus MSR8 India HQ450771 Kumari et al. (2012)
C. inusitatus CAL 1625 (T) India MG593195 Bijeesh et al. (2018)
C. lutescens UPSF11789 Sweden AF105302 Dahlman et al. (2000)
C. lutescens UPSF11790 Sweden AF105303 Dahlman et al. (2000)
C. lutescens UPSF11791 Spain AF105304 Dahlman et al. (2000)
C. luteus GDGM46432 China MG727898 Zhong et al. (2018)
C. luteus GDGM48105 (T) China MG701171 Zhong et al. (2018)
C. luteus GDGM49495 China MG806926 Zhong et al. (2018)
C. striatus HMJAU 61463 (T) China OM509446 ON125913 This study
C. striatus HMJAU 61463 China OM509447 ON125914 This study
C. odoratus UPSF11794 USA AF105306 Dahlman et al. (2000)
C. olivaceoluteus TH9205 (T) Guyana JQ915135 Wilson et al. (2012)
C. olivaceoluteus MCA3186 Guyana JQ915124 Wilson et al. (2012)
C. parvogriseus CAL1533 (T) India MF421098 Das et al. (2017)
C. parvogriseus CAL1534 India NG059049 Das et al. (2017)
C. pleurotoides TH9220 (T) Guyana JQ915136 Wilson et al. (2012)
C. pleurotoides MCA3124 Guyana JQ915123 Wilson et al. (2012)
C. strigosus MAC1750 Guyana JQ915120 Wilson et al. (2012)
C. strigosus TH9204 (T) Guyana JQ915134 Wilson et al. (2012)
C. tubaeformis UPSF11793 Sweden AF105307 Dahlman et al. (2000)
C. tubaeformis BB 07.293 Slovakia KF294640 GQ914989 Buyck and Hofstetter (2011)
Cantharellus cibarius BIO 10986 (T) Sweden KR677539 KX828823 Olariaga et al. (2017)
Hydnum ellipsosoporxum FD3281 Switzerland KX086217 Genbank

Sequences were assembled and edited using the software package Sequencher 5.4.6 (Gene Codes Corp., USA). Alignment of sequence data was performed with MUSCLE in MEGA 7.0.21 (Kumar et al. 2016). For Bayesian Inference (BI) analyses, the most suitable substitution model for each gene partition was calculated with ModelFinder (Kalyaanamoorthy et al. 2017) in PhyloSuite 1.2.2. (Zhang et al. 2020b). BI analyses were performed using MrBayes in PhyloSuite 1.2.2 (Zhang et al. 2020b), implementing the Markov Chain Monte Carlo (MCMC) technique. Four simultaneous Markov chains were run from random trees, keeping one tree every 200th generation until the average standard deviation of split frequencies was below 0.01. The Maximum Likelihood analysis was performed using RAxML-HPC2 on XSEDE 8.2.12 (Stamatakis 2014) in the CIPRES Science Gateway portal (https://www.phylo.org/portal2/tools.action) with the GTRGAMMA model and searching for the most likely tree with 1000 heuristic replicates. The bootstrap support (BS) of ≥ 50% in the ML tree and BPP of ≥ 0.75 indicated statistical significance. The phylogenetic trees were visualised using FigTree 1.4.23 (Andrew 2016).

Results

Phylogeny

Seven new DNA sequences (3 nr LSU, 4 tef-1α) were produced for this study. After removing introns and low-homology regions, the final combined alignment of these two genes totalled 2,027 characters (nr LSU: 1,024 characters; tef-1α: 1,003 characters). The best models for the BI analysis of the concatenated dataset were SYM+I+G4 for nr LSU and SYM+G4 for tef-1α. The most likely tree inferred by ML analysis of the combined dataset exhibited a quite similarly supported topology as the Bayesian majority-rule consensus tree with an average standard deviation of split frequencies = 0.005143. The most likely tree, based on 1000 searches, is depicted in Fig. 1 with associated bootstrap values. Maximum Likelihood Bootstrap (MLBS) and Bayesian Posterior Probability (BPP) were established along the branches. Phylogenetic analyses show (Fig. 1) that our two new species, C. connatus and C. striatus were clustered together with C. atrobrunneolus in a monophyletic clade (MLBS = 99%, BPP = 1.00). C. connatus (MLBS = 88%, BPP = 0.93) formed sister relationships (MLBS = 88%, BPP = 0.87) with C. striatus (MLBS = 60%, BPP = 0.77).

Figure 1. 

BI best tree inferred from the nr LSU and tef-1α region analysis for 25 specimens. Branches that received both bootstrap support (MLBS) ≥ 50% and Bayesian posterior probabilities (PP) ≥ 0.75 are in bold; branches supported by either MLBS or BPP are in grey. Both values (MLBS/BPP) are reported along the branches. Taxon names shown in bold indicate the specimens examined in this study.

Taxonomy

Craterellus connatus G.P. Zhao, J.J. Hu, B. Zhang & Y. Li, sp. nov.

MycoBank No: 842527
Figs 2, 3

Holotype

China. Jilin Province, Jilin City, Jiaohe County, Lafashan National Forest Part, Red Leaves Canyon, alt. 802.5 m, 43.75°N, 127.10°E, 5 September 2018, Bo Zhang HMJAU 61462, GenBank Acc. nos.: nr LSU = OM509448, tef-1α = ON125915, ON125916).

Figure 2. 

Craterellus connatus (HMJAU 60411, holotype) A fresh basidiocarps B connate stipes C margin of pileus D hymenophore. Scale bars: 1 cm (A, B)

Etymology

Connatus: referring to several stipes grown together from the base upwards.

Diagnosis

Differs from other Craterellus species by its greyish-brown and smooth pileus with an off-white margin, hymenophore with a strongly anastomosing vein and the colouration turning khaki upon drying.

Figure 3. 

Microscopic characteristics of Craterellus connatus (HMJAU 60411) A basidiospores B basidia C pileipellis. Scale bars: 5 μm (A); 10 μm (B); 20 μm (C).

Description

Pileus 18–30 mm in diam., infundibuliform, deeply depressed at the centre, perforated continuously to the base of the stipe; margin inrolled when young, then expand upwards, becoming upturned finally, broadly wavy; surface greyish-brown in the centre, the marginal edge dirty white, smooth. Hymenophore consists of longitudinal, anastomosing veined, off-white, turning khaki when drying and decurrent. Stipe 25–30 × 3 mm, equal to subcylindrical, hollow, central, smooth, greyish-brown. Text fleshy, greyish-brown. Odour light. Taste unknown. Spore print not obtained.

Basidiospores 6.1–6.9–7.8(8.1) × 4.8–5.3–5.9 μm, Q = (1.16)1.20–1.32–1.45(1.56), Qm = 1.32 ± 0.09, broad ellipsoid to ellipsoid, smooth, thin-walled, pale yellow in 5% aqueous KOH, inamyloid. Basidia (40)50–72.5(75) × (6)7.5–8(10) μm, clavate, sterigmata (2)4-6. Cystidia absent. Hymenium in transverse section 125–175 μm thick, yellowish-brown in 5% aqueous KOH. Pileipellis scarcely differentiated from the trama, a cutis of vastly inflated hyphae, yellowish-brown in 5% aqueous KOH; individual hyphae (22)27–63(64) × 6–15 μm, thin-walled, branched occasionally, secondary septation absent, hyaline in 5% aqueous KOH. Pileus trama up to 500 μm thick, subparallel, yellowish-brown in 5% aqueous KOH; individual hyphae (29)33–60(125) × (4)5–12(13) μm, cylindrical, thin-walled, branching frequently, secondary septation absent. Subhymenium up to 10 μm thick. Stipitipellis composed of a tightly packed mass of subparallel, narrow, cylindrical hyphae, yellowish-brown in 5% aqueous KOH; individual hyphae (33)37–79(102) × 4–13(16) μm, thin-walled, hyaline to pale yellow in 5% aqueous KOH, branched frequently, secondary septation absent. Clamp connections absent.

Habitat

Scattered or gregarious in small clusters on the ground in coniferous and angiosperm mixed forests.

Craterellus striatus G.P. Zhao, J.J. Hu, B. Zhang & Y. Li, sp. nov.

MycoBank No: 842531
Figs 4, 5

Holotype

China. Jilin Province, Baishan City, Fusong County, Quanyang Town, alt. 780.4 m, 42.23°N, 121.30°E, 22 August 2021, G. P. Zhao, J. J. Hu, and Bo Zhang (HMJAU 61463, GenBank Acc. nos.: nr LSU = OM509446, OM509447, tef-1α = ON125913, ON125914).

Etymology

Striatus: referring to the fringe of pileus.

Diagnosis

Differs from other Craterellus species by the fibrillose, greyish-brown to yellowish-brown, fully perforated pileus with a brown fringe, the hymenophore with a forking vein, not anastomosing and non-discolouring upon drying.

Figure 4. 

Craterellus striatus (HMJAU 60412, holotype) A Fresh basidiocarps B Stipe C Pileus D Hymenophore. Scale bars: 1 cm (A, B)

Description

Fruiting body 12–45 mm high. Pileus 3–21 mm in diam., 1–10 mm tall, plane-convex firstly, soon infundibuliform and usually perforated continuous to the base of the stipe; margin inrolled when young, then expanding outwards, old becoming incurved, broadly wavy; surface blackish-brown, cream near the margin at first, gradually lighter to the centre with age, yellowish-brown finally, turning pale greyish-brown when drying, covered with brown fringe and fibrillose scales. Hymenophore decurrent, consisting of longitudinal ridges (<1 mm) with prominent forking, cross vein and off-white. Stipe 11–35 × 1–6 mm, cylindrical, inflated at the base, up to 10 mm in diam., hollow, central, smooth, blackish-brown. Text leathery, dirty white. Odour light. Taste unknown. Spore print not obtained.

Figure 5. 

Microscopic characteristics of Craterellus striatus (HMJAU 60412) A spores B basidiobasidia C pileipellis. Scale bars: 10 μm (A); 20 μm (B); 40 μm (C).

Basidiospores (6.5)6.8–7.4–8.0(8.8) × (5.0)5.1–5.5–6.0 μm, Q = (1.18)1.22–1.36–1.53(1.73), Qm = 1.36 ± 0.12, broad ellipsoid to ellipsoid, smooth, thin-walled, hyaline to pale yellow in 5% aqueous KOH, inamyloid. Basidia (34)40–67(68) × 6–9 μm, slender, sterigmata 2–6. Cystidia absent. Hymenium in transverse section 45 μm thick, yellowish-brown in 5% aqueous KOH. Pileipellis scarcely differentiated from trama, a cutis of largely cylindrical hyphae, yellowish-brown in 5% aqueous KOH; individual hyphae (30)40–70(73) × (4)5–14(15) μm, thin-walled, encrusted, branched frequently, secondary septation absent, hyaline in 5% aqueous KOH. Pileus trama up to 500 μm thick, subparallel, yellowish-brown in 5% aqueous KOH; individual hyphae (29)33–60(125) × (4)5–12(13) μm, cylindrical, thin-walled, encrusted, branching frequently, secondary septation absent. Subhymenium up to 10 μm thick. Stipitipellis composed of a tightly-packed mass of subparallel, narrow, cylindrical hyphae, yellowish-brown in 5% aqueous KOH; individual hyphae (33)37–79(102) × 4–13(16) μm, thin-walled, hyaline to pale yellow in 5% aqueous KOH, branched frequently, secondary septation absent. Clamp connections absent.

Habitat

Scattered to gregarious on the ground in a coniferous and angiosperm mixed forest.

Key to reported species of Craterellus in China

1 Clamp connections present 2
Clamp connections absent 4
2 Stipe golden yellow to orangish-yellow 3
Stipe grey C. atratus
3 Pileus perforate to the base of the stipe C. tubaeformis
Pileus not perforate C. lutescens
4 Basidiomata not brown, absolutely light colour 5
Basidiomata greyish-brown to blackish-brown 7
5 Pileus white C. albidus
Pileus yellow 6
6 Pileus very small, usually < 10 mm in diam C. aureus
Pileus large, usually > 90 mm in diam C. luteus
7 Basidiospores 10–15 μm long, average length > 11 µm 10
Basidiospores 6–12 μm long, average length < 11 µm 8
8 Hymenophore smooth to slightly folded 9
Hymenophore with well-developed veins or gill-folds C. striatus
9 Basidia long, up to 106 µm long C. badiogriseus
Basidia short 13
10 Basidia with 2 sterigmata 11
Basidia with 2–4 sterigmata 12
11 Basidiospores broad, up to 11.5 μm wide C. macrosporus
Basidiospores 2–4 μm wide C. cornucopioides
12 Pileus surface smooth C. croceialbus
Pileus surface often with darker brown raised scales C. squamatus
13 Margin dirty white C. connatus
Margin not dirty white C. atrobrunneolus

Discussion

In our study, two new species formed a sister relationship with Craterellus atrobrunneolus. These three species show close morphological and phylogenetic similarities with each other. All species share the brown pileus, grey hymenophore, hollow stipe, narrow basidia and absence of clamps. Craterellus atrobrunneolus was initially described in south-western China. It is characterised by a dark brown to brownish-grey colouration, convex to plano-convex pileus with shallow depression, but not perforated centre, smooth to slightly folded hymenophore, absence of clamp connections in all tissues, narrow basidia with 2–6 sterigmata and broad ellipsoid to subglobose basidiospores (Cao et al. 2021b). Craterellus connatus is recognised in the field by the medium-sized, nearly fleshy basidiomata with the greyish-brown, fully perforated pileus, an off-white margin, strongly anastomosed, veined hymenophore and hollow stipe. Microscopically, it possesses broad ellipsoid to ellipsoid basidiospores (Qm = 1.32 ± 0.09), slender basidia with (2)4–6 sterigmata and an absence of clamp connections. Craterellus striatus is characterised by its small-sized basidiomata, blackish-brown pileus that turns yellowish-brown upon drying and is covered with brown fringe and spinous scales and off-white hymenophore consisting of longitudinal ridges (<1 mm) with prominent forking. It has a hollow and brown stipe and broad ellipsoid to ellipsoid basidiospores, 2–6 spored basidia, encrusted hyphae and the absence of the clamp connection.

Craterellus atrobrunneolus differs by its dark brown to almost black throughout, convex but not perforated pileus, while C. connatus possesses greyish-brown pileus with an off-white margin and C. striatus possesses yellowish-brown, perforated pileus with brown fringe. Microscopically, C. atrobrunneolus possesses broad ellipsoid to subglobose basidiospores, while C. connatus and C. striatus possess broad ellipsoid to ellipsoid basidiospores. In addition, C. atrobrunneolus have 2–6 spored basidia, while C. connatus have mostly 4–6 spored basidia. Craterellus striatus have encrusted hyphae, while C. atrobrunneolus did not. Although the two new species have similar microscopic characteristics (spores and basidia), they are separated by their colouration of dried pileus and hymenophore and the configuration of hymenophore. Craterellus striatus is smaller (both pileus and stipe) than C. connatus. Further, C. striatus has a hymenophore composed of the forked longitudinal ridge and non-discolouring upon drying, whereas C. connatus has an anastomosing veined hymenophore, which turns khaki upon drying. Morphology and phylogenetic analyses indicated that the two new species in this study are not conspecific.

However, Craterellus atrobrunneolus was not included in any subgenus in Cao et al. (2021a). According to our study, C. atrobrunneolus, C. conatus and C. striatus have formed an isolated branch. Further research is needed to obtain a more precise infrageneric classification. Additionally, this study suggests that China, especially north-eastern China, has considerable fungal diversity and possibly many endemic species.

Acknowledgements

This study is funded by the National Natural Science Foundation of China (No. 31970020), the Scientific and Technological Tackling Plan for the Key Fields of Xinjiang Production and Construction Corps (No. 2021AB004), the National Key R & D of the Ministry of Science and Technology (2019YFD1001905-33), Research on the creation of excellent edible mushroom resources and high quality & efficient ecological cultivation technology in Jiangxi Province (20212BBF61002) and the National Key R & D of Ministry of Science and Technology (2018YFE0107800).

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