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Research Article
Species diversity of Tricholoma (Agaricales, Tricholomataceae) from Shanxi Province of northern China with the description of four new species
expand article infoNing Mao‡, Jia-Jia Yang‡, Yu-Xin Zhang‡, Ting Li§, Li Fan‡
‡ Capital Normal University, Beijing, China
§ Natural History Museum of China, Beijing, China

Corresponding author: Li Fan ( fanli@mail.cnu.edu.cn )

Academic editor: Zai-Wei Ge
© 2025 Ning Mao, Jia-Jia Yang, Yu-Xin Zhang, Ting Li, Li Fan.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Mao N, Yang J-J, Zhang Y-X, Li T, Fan L (2025) Species diversity of Tricholoma (Agaricales, Tricholomataceae) from Shanxi Province of northern China with the description of four new species. MycoKeys 112: 59-80. https://doi.org/10.3897/mycokeys.112.132652
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Abstract

Species of Tricholoma are of great economic and ecological value. There are many studies on Tricholoma worldwide, but the areas investigated are generally North America and Europe. There is limited knowledge about Tricholoma in China. In this study, 21 species of Tricholoma were confirmed in Shanxi Province based on morphological and molecular phylogenetic analyses. These species are located in eight sections, viz. sect. Atrosquamosa, sect. Genuina, sect. Lasciva, sect. Matsutake, sect. Pardinicuti, sect. Rigida, sect. Terrea and sect. Tricholoma. Of these, four species are described as new to science: T. flavoviride, T. fumeobrunneum, T. parafulvum, and T. viscidum.

Key words

Basidiomycota, ectomycorrhiza, phylogenetic analyses, taxonomy

Introduction

Tricholoma (Fr.) Staude was erected as a genus in the year 1857 (Staude 1857). This genus is mainly characterized by fleshy basidiomata, adnexed to emarginate lamellae, a central stipe, white spore prints, subglobose to oblong basidiospores, simple pileipellis structures and often the absence of well-differentiated cystidia (Kost 1981; Singer 1986; Christensen and Noordeloos 1999; Bessette et al. 2013; Christensen and Heilmann-Clausen 2013; Heilmann-Clausen et al. 2017; Reschke et al. 2018; Ding et al. 2023). Species of Tricholoma are ecologically and economically vital, not only for forming ectomycorrhizal symbioses with trees of the families Pinaceae, Betulaceae, Fagaceae, Salicaceae, Myrtaceae and Nothofagaceae, but also as famous delicacy mushrooms, such as T. matsutake (S. Ito & S. Imai) Singer and its allies (Bougher 1996; Chapela and Garbelotto 2004; Matsushita et al. 2005; Suzuki 2005; Tedersoo et al. 2010; Ota et al. 2012; Bessette et al. 2013; Christensen and Heilmann-Clausen 2013; Sánchez-García and Matheny 2016; Heilmann-Clausen et al. 2017; Reschke et al. 2018; Ding et al. 2023). Based on morphological and molecular approaches, a number of infrageneric classifications of Tricholoma have been proposed (Bon 1984, 1991; Singer 1986; Riva 1988, 2003; Christensen and Noordeloos 1999; Noordeloos and Christensen 1999; Christensen and Heilmann-Clausen 2008; Reschke et al. 2018; Landry et al. 2022; Trudell et al. 2022; Ding et al. 2023). For example, Ding et al. (2023) assessed subgenera and section boundaries within Tricholoma by using morphology and phylogenetic evidence from 50-locus, in which a total of four subgenera and 11 sections were identified.

Recently, some important taxonomic works have been focused on Tricholoma in China (Cui et al. 2022; Ding et al. 2023; Yang et al. 2023). However, our knowledge of Tricholoma is significantly insufficient in Shanxi Province. During the last seven years, we collected a lot of specimens of Tricholoma in this region. Our morphological examinations and molecular analyses based on these collections showed they represented 21 species, including four un­described species. The aim of this paper is to describe these new species and clarify the species diversity and geographic distribution of Tricholoma in Shanxi Province, northern China.

Materials and methods

Morphological studies

Collections were obtained and photographed in the field from Shanxi Province, North China, dried in a fruit drier at 40–45 °C, and deposited at BJTC (Herbarium Biology Department, Capital Normal University) and HSA (Herbarium of Shanxi Institute for Functional Foods, Shanxi Agricultural University). Macroscopic characters were recorded from fresh specimens. Standardized color values matching the color of the description were taken from ColorHexa (http://www.colorhexa.com/). Microscopic characteristics were observed in sections obtained from dry specimens mounted in 5% KOH, Congo Red, or Melzer’s reagent (Dring 1971). Dimensions of basidiospores are given using the following format ‘(a–)b–c(–d)’, where the range ‘b–c’ represents at least 90% of the measured values, and ‘a’ and ‘d’ are the most extreme values. ‘Q’ refers to the length/width ratio of basidiospores in the side view.

DNA extraction, PCR amplification, and DNA sequencing

A small amount of basidiomata material (20–30 mg) was crushed by shaking for 45 seconds at 30 Hz 2–4 times (Mixer Mill MM301, Haan, Germany) in a 1.5 mL tube, together with a 3 mm diam. tungsten carbide ball. Total genomic DNA was extracted from the powdered basidiomata using NuClean Plant Genomic DNA Kit (CWBIO, Beijing, China), following the manufacturer’s instructions. The ITS region was amplified using the primers ITS1-F/ITS4 (White et al. 1990; Gardes and Bruns 1993). Polymerase Chain Reactions (PCR) for ITS region was performed in 25 µl reaction containing 2 μl DNA template, 1 µl primer (10 µM) each, 12.5 µl of 2 × Master Mix [Tiangen Biotech (Beijing) Co.], 8.5 μl ddH2O. PCR reactions were implemented as follows: an initial denaturation at 94 °C for 5 min; then 35 cycles of the following denaturation at 94 °C for 30 s, annealing at 55 °C for 45 s, 72 °C for 1 min; and a final extension at 72 °C for 10 min. The PCR products were sent to Beijing Zhongkexilin Biotechnology Co. Ltd. (Beijing, China) for purifying, sequencing, and editing. Accession numbers of new and downloaded sequences stored in the NCBI database are provided in Table 1.

Table 1.
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Specimens used in ITS phylogenetic analysis and their GenBank accession numbers. Newly generated sequences are in bold.

Taxon name Specimen voucher Origin ITS
Leucopaxillus laterarius KUN-HKAS 106319 China MW724394
Leucopaxillus tricolor MB 000946 Germany MW724429
Tricholoma albobrunneum MC99-060 France LT000077
Tricholoma albobrunneum KUN-HKAS 68189 China MW724479
Tricholoma album MC95-159 Denmark LT000008
Tricholoma album MB 006366 Germany MW724416
Tricholoma album MB 006323 Germany MW724421
Tricholoma ammophilum WTU-F-073083 USA MW597140
Tricholoma ammophilum WTU-F-073015 USA MW597199
Tricholoma ammophilum HSA 371 China PQ499533
Tricholoma argyraceum MEN9491 Netherlands LT000198
Tricholoma argyraceum BJTC FM2197 China PQ499519
Tricholoma argyraceum BJTC FM2372 China PQ499520
Tricholoma atrosquamosum BJTC FM4346 China PQ499525
Tricholoma aurantium MC97-227 Denmark LT000012
Tricholoma aurantium BJTC FM2193 China PQ499534
Tricholoma aurantium BJTC FM2195 China PQ499535
“Tricholoma auratum” Tk1 Japan AB289663
“Tricholoma auratum” ISK1 Japan AB289662
Tricholoma badicephalum UBC-F-16235r Canada MW597207
Tricholoma badicephalum WTU-F-073095 USA MW597309
Tricholoma bakamatsutake TNS:F-12866 Japan AB699654
Tricholoma bakamatsutake KUN-HKAS 106313 China MW724402
Tricholoma bakamatsutake KUN-HKAS 107570 China MW724468
Tricholoma bakamatsutake BJTC FM3440 China PQ499506
Tricholoma bakamatsutake BJTC FM3441 China PQ499507
Tricholoma basirubens MC01-209 Croatia LT000001
Tricholoma basirubens TL5303 Sweden LT000158
Tricholoma batschii KMS436 USA AF377238
Tricholoma batschii MB-003027 Germany MF034298
Tricholoma bonii LUG-F8450 Italy LT000101
Tricholoma bonii HMJAU35946 China MW724393
Tricholoma bonii BJTC FM1280 China PQ499508
Tricholoma bonii BJTC FM1388 China PQ499509
Tricholoma bonii BJTC FM920 China PQ499510
Tricholoma boudieri HKAS97070 China MW724437
Tricholoma boudieri HKAS97163 China MW724373
Tricholoma boudieri BJTC FM1528 China PQ499521
Tricholoma boudieri BJTC FM2752 China PQ499522
Tricholoma cingulatum MC96-134 Denmark LT000015
Tricholoma cingulatum BJTC FM1172 China PQ499516
Tricholoma cingulatum BJTC FM820 China PQ499517
Tricholoma cingulatum BJTC FM1191 China PQ499518
Tricholoma citrinum MB-305716 China MF034262
Tricholoma citrinum KUN-HKAS 71086 China MW724356
Tricholoma colossus MC97-047 Sweden LT000164
Tricholoma colossus MB-002363 Germany MF034285
Tricholoma columbetta MC95-181 Denmark LT000017
Tricholoma columbetta MQ20-HRL3139-QFB32663 Canada MW628118
Tricholoma dulciolens H:7002022 Sweden AB738883
Tricholoma dulciolens - USA AF309523
Tricholoma elegans OTA:61947 New Zealand JX178630
Tricholoma elegans TENN:063711 New Zealand KJ417316
Tricholoma equestre MC94-027 Denmark LT000018
Tricholoma equestre MC96-155 Denmark LT000020
Tricholoma equestre HMJAU22249 Belarus MW724392
“Tricholoma equestre” MB305549 China MF034257
“Tricholoma equestre” MB305676 China MF034261
“Tricholoma equestre” MB-301506 China MF034239
“Tricholoma equestre” EqFrW France HM590874
Tricholoma filamentosum C-F-35924 Sweden LT000165
Tricholoma filamentosum MB 000950 (KR9404) Germany MW724422
“Tricholoma flavovirens” trh545 USA AF458449
“Tricholoma flavovirens” trh546 USA AF458452
“Tricholoma flavovirens” 613 Japan AB036895
“Tricholoma flavovirens” KGP52 USA DQ822834
Tricholoma flavoviride BJTC FM3966 China PQ499559
Tricholoma flavoviride BJTC FM4164 China PQ499560
Tricholoma flavoviride BJTC FM3512 China PQ499561
Tricholoma focale JV97-239 Sweden LT000166
Tricholoma focale KUN-HKAS 106309 China MW724460
Tricholoma forteflavescens HKAS93511 China MF034207
Tricholoma forteflavescens MB-301985 China MF034246
Tricholoma frondosae type I MC95-130 Sweden LT000167
Tricholoma frondosae type I KUN-HKAS 98072 China MW724365
Tricholoma frondosae type I KUN-HKAS 87149 China MW724346
Tricholoma frondosae type II MC96-235 Denmark LT000023
Tricholoma frondosae type II MC00-225 Slovenia LT000140
Tricholoma frondosae type II BJTC FM2026 China PQ499556
Tricholoma frondosae type II BJTC FM3895 China PQ499557
Tricholoma frondosae type II BJTC FM4157 China PQ499558
Tricholoma fulvomaculatum HKAS107572 China MW724472
Tricholoma fulvomaculatum HKAS107576 China MW724473
Tricholoma fulvum JHC04-251 Sweden LT000171
Tricholoma fulvum MQ20-YL-CMMF001495 Canada MW627880
Tricholoma fumeobrunneum BJTC FM3445 China PQ499552
Tricholoma fumeobrunneum BJTC FM3436 China PQ499553
Tricholoma fumeobrunneum BJTC FM3571 China PQ499554
Tricholoma fumeobrunneum BJTC FM3574 China PQ499555
Tricholoma highlandense HKAS70192 China KY488549
Tricholoma highlandense KUN-HKAS 107590 China MW724452
Tricholoma ilkkae S-F173364 Sweden LT222028
Tricholoma ilkkae S-F513823 Sweden LT222029
“Tricholoma joachimii” O-F167194 Norway LT222022
Tricholoma imbricatum KUN-HKAS 112559 China MW724476
Tricholoma imbricatum KUN-HKAS 87886 China MW724327
Tricholoma imbricatum BJTC FM1170 China PQ499549
Tricholoma imbricatum BJTC FM1445 China PQ499550
Tricholoma imbricatum BJTC FM1446 China PQ499551
Tricholoma inocybeoides MC03-229 Denmark LT000025
Tricholoma inocybeoides MC97-060 Sweden LT000176
Tricholoma lascivum MC00-519 Denmark LT000028
Tricholoma lascivum MB-303096 Ukraine MF034316
Tricholoma lishanense BJTC FM1023 China PQ499526
Tricholoma lishanense BJTC FM1137 China PQ499527
Tricholoma lishanense BJTC FM1735 China PQ499528
Tricholoma matsutake KUN-HKAS 98323 China MW724385
Tricholoma matsutake MC03-600 Sweden LT000178
Tricholoma matsutake TNS:F-12850 Japan AB699630
Tricholoma murrillianum SAT-16-319-01 USA KY660032
Tricholoma murrillianum NY586560 USA LT220179
Tricholoma olivaceotinctum MC97103 Sweden FJ544861
Tricholoma olivaceotinctum KUN-HKAS 107586 China MW724405
Tricholoma olivaceum HKAS93513 China MF034209
Tricholoma olivaceum KUN-HKAS 68600 China MW724351
Tricholoma orienticolossus HAKS99341 China MT124443
Tricholoma orienticolossus HAKS98045 China MT124444
Tricholoma orientifulvum HAKS107157 China MT114682
Tricholoma orientifulvum HAKS107156 China MT124445
Tricholoma parafulvum BJTC FM2500 China PQ499536
Tricholoma parafulvum BJTC FM4065 China PQ499537
Tricholoma parafulvum BJTC FM4210 China PQ499538
Tricholoma pardinum C-F-96190 USA LT000142
Tricholoma pardinum MB 006381 Germany MW724424
Tricholoma pessundatum JV04-482 Denmark LT000032
Tricholoma pessundatum MQ20-JLAB931-CMMF009347 Canada MW628012
Tricholoma populinum O-F63960 Norway JN019594
Tricholoma populinum KUN-HKAS 106657 China MW724411
Tricholoma populinum BJTC FM1144 China PQ499529
Tricholoma populinum BJTC FM1145 China PQ499530
Tricholoma populinum BJTC FM1165 China PQ499531
Tricholoma populinum BJTC FM1979 China PQ499532
Tricholoma psammopus MC04-600 Slovenia LT000145
Tricholoma psammopus KUN-HKAS 106302 China MW724436
Tricholoma psammopus BJTC FM2673 China PQ499546
Tricholoma psammopus BJTC FM1405 China PQ499547
Tricholoma psammopus BJTC FM814 China PQ499548
Tricholoma qiaomianjun KUN-HKAS 101303 China OK036719
Tricholoma qiaomianjun KUN-HKAS 115901 China OK036720
Tricholoma roseoacerbum MQ20-HRL1010a-QFB32619 Canada MW628060
Tricholoma roseoacerbum KUN-HKAS 88046 China MW724332
Tricholoma rufobrunneum KUN-HKAS49069 China OL331894
Tricholoma rufobrunneum KUN-HKAS90808 China OL331895
Tricholoma saponaceum C-F23337 Denmark LT000038
Tricholoma saponaceum JHC00-049 Norway LT000123
Tricholoma saponaceum MB-002941 Germany MF034221
Tricholoma saponaceum BJTC FM2772 China PQ499523
Tricholoma saponaceum BJTC FM2773 China PQ499524
Tricholoma sinoacerbum GDGM:44680 China KT160219
Tricholoma sinoacerbum KUN-HKAS 105349 China MW724434
Tricholoma sinopardinum KUN-HKAS 91129 China MW724361
Tricholoma sinopardinum HKAS82533 China KY488552
Tricholoma smithii DBG:CLO4513 USA MG719957
Tricholoma sp. HKAS106303 China MW724450
Tricholoma sp. HKAS101281 China MW724443
Tricholoma squarrulosum JHC93-224 Denmark LT000047
Tricholoma squarrulosum JHC93-262 Denmark LT000048
Tricholoma stans MC95-145 Sweden LT000189
Tricholoma stans KUN-HKAS 87940 China MW724329
Tricholoma stiparophyllum MC95-117 Sweden LT000190
Tricholoma stiparophyllum MQ20-GUE1522-CMMF014811 Canada MW628089
Tricholoma sudum JV96-306 Denmark LT000050
Tricholoma sudum MC98-601 Denmark LT000051
Tricholoma terreum MEN95192 Germany LT000098
Tricholoma terreum KUN-HKAS 69914 China MW724459
Tricholoma terreum BJTC FM2414 China PQ499511
Tricholoma terreum BJTC FM3657 China PQ499512
Tricholoma terreum BJTC FM3677 China PQ499513
Tricholoma triste E3754 Germany LT000099
Tricholoma triste BJTC FM1269 China PQ499514
Tricholoma triste BJTC FM3817 China PQ499515
“Tricholoma ulvinenii” JuV13229F Finland LT000068
“Tricholoma ulvinenii” JuV26740F Finland LT000069
“Tricholoma ulvinenii” IK931613 Finland LT000067
Tricholoma umbonatum type I MC00A01 Denmark LT000063
Tricholoma umbonatum type II TRgmb00651 Italy LT000114
Tricholoma ustale JHC92-299 Denmark LT000064
Tricholoma ustale MB-002924 Germany MF034288
Tricholoma ustaloides MC99-067 France LT000094
Tricholoma ustaloides MB-002929 Germany MF034291
Tricholoma vaccinum MC95-109 Sweden LT000195
Tricholoma vaccinum DBG:23466 USA MF034272
Tricholoma vaccinum KUN-HKAS 98065 China MW724364
Tricholoma vaccinum BJTC FM937 China PQ499543
Tricholoma vaccinum BJTC FM943 China PQ499544
Tricholoma vaccinum BJTC FM1425 China PQ499545
Tricholoma venenatoides WTU-F-073089 USA MW597303
Tricholoma viscidum BJTC FM4198 China PQ499539
Tricholoma viscidum BJTC FM3388 China PQ499540
Tricholoma viscidum BJTC FM4038 China PQ499541
Tricholoma viscidum BJTC FM4199 China PQ499542

Phylogenetic analyses

For this study, the ITS dataset was compiled to identify the new species and to investigate their phylogenetic position in the Tricholoma. Leucopaxillus tricolor (Peck) Kühner and Leucopaxillus laterarius (Peck) Singer & A.H. Sm. were selected as outgroups based on previous studies (Ding et al. 2023). The sequences of the ITS were aligned in the online version of MAFFT 7.110 using default parameters (Katoh and Standley 2013) and manually edited in BioEdit v.7.0.9 (Hall 1999). The final alignments were submitted to TreeBASE 31776.

Phylogenetic analyses were conducted using maximum likelihood (ML) and Bayesian inference (BI). ML analysis was carried out in RAxML 8.0.14 (Stamatakis 2014) with parameters at default settings using a GTRGAMMAI model. 1000 bootstrap replicates were computed in RAxML using a rapid bootstrap analysis and search for the best-scoring ML tree. BI analysis was performed in MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003). The best-fitted substitution model for ITS was determined through MrModeltest v2.3 (Nylander 2004) by using the Akaike Information Criterion (AIC). GTR+I+G was chosen as the best model for ITS. We used two independent runs with four Markov chains Monte Carlo (MCMC) for 3 775 000 generations under the default settings. The average standard deviations of split frequency (ASDSF) values were far lower than 0.01 at the end of the runs. Trees were sampled every 100 generations after burn-in (25% of trees were discarded as the burn-in phase of the analyses, set up well after convergence), and 50% of majority-rule consensus trees were constructed.

Clades with bootstrap support (MLBS) ≥ 70% and Bayesian posterior probability (BPP) ≥ 0.95 were considered significantly supported (Hillis and Bull 1993; Alfaro et al. 2003). All phylogenetic trees were viewed with TreeView (Page 2001).

Results

Phylogenetic analyses

The ITS dataset consisted of 191 sequences, including 56 sequences newly generated from our collections. The length of the dataset was 578 bp after the exclusion of poorly aligned sites. Our present analyses (Fig. 1) revealed that members of Tricholoma in this dataset formed a monophyletic lineage with high likelihood bootstrap support (MLB = 100%) and strong posterior probability support (BPP = 1.00). The sequences of our 56 collections formed 21 strongly support clades, indicating they were 21 distinct species, and these phylogenetic species were respectively placed in eight sections, i.e. sect. Atrosquamosa, sect. Genuina, sect. Lasciva, sect. Matsutake, sect. Pardinicuti, sect. Rigida, sect. Terrea and sect. Tricholoma. Of them, four species are described as new species (see Taxonomy in this paper), and the remaining 17 clades correspond well to the previously described species (also see the photos of their basidiocarps in Figs 4, 5).

Figure 1. 

Phylogenetic tree generated from a maximum likelihood analysis based on ITS sequences, showing the phylogenetic relationships of Tricholoma. Numbers representing likelihood bootstrap support (MLB ≥ 70%, left) and significant Bayesian posterior probability (BPP ≥ 0.95, right) are indicated above the nodes. Novel sequences are printed in bold.

Taxonomy

Tricholoma flavoviride L. Fan, N. Mao & J.J Yang, sp. nov.

MycoBank No: MB 856305
Figs 2A, B, 3A, C

Diagnosis

It is distinguished by the combination of the following features: greenish-yellow to yellowish-brown pileus, stipe surface with brownish fibrils, or reflexed squamules. It is most similar to T. citrinum Y.Y. Cui & Zhu L. Yang but differs by habitat associated with broadleaf forest.

Holotype

China • Shanxi Province, Pu County, Wulu Mountain, 36°33'34"N, 111°12'40"E, elev. 1510 m, on the ground in broadleaf forest dominated by Quercus sp., 28 September 2023, J.Z Cao, MS808 (BJTC FM4164), GenBank Acc. No.: ITS = PQ499560, mtSSU = PQ499564, rpb2 = PQ509907, mcm7 = PQ509917.

Etymology

flavoviride (Lat.): referring to the color of the basidiomata.

Description

Basidiomata small to medium-sized. Pileus 20–55 mm diam., at first hemispherical to convex, later plano-convex to applanate with age, with an umbo at center; surface dry, slightly viscid when wet, greenish-yellow (#dbd252) to yellowish-brown (#c2a677), center darker to dark brown (#7a614d) when mature, covered with brown (#b49976) squamules; margin incurved or not. Lamellae sinuate, crowded, greenish-yellow (#b4ae6c) to yellowish-brown (#d9c45b); lamellulae in 2–4 tiers, concolorous with lamellae. Stipe 32–70 mm long, 12–17 mm diam., cylindrical, equal or enlarge downwards, dry, white (#f9f9f8) at apex, pale yellow (#e7e1a5) to yellowish brown (#d7cb67) below, covered with brownish (#9c8c58) fibrils or reflexed squamules. Context white (#ffffff). Odor unrecorded. Taste not recorded.

Figure 2. 

Photographs of basidiomata in their natural habitat A, B Tricholoma flavoviride (A BJTC FM3996, B BJTC FM4164) C, D T. fumeobrunneum (C BJTC FM3571, D BJTC FM3574) E, F T. parafulvum (E BJTC FM2500, F BJTC FM4065) G, H T. viscidum (G BJTC FM4038, H BJTC FM4198). Scale bars: 1 cm.

Basidiospores [90/3/3] 5–6.5 × 3.5–5 μm, Q = 1.2–1.6, ellipsoid, smooth, inamyloid, usually containing one large oil droplet. Basidia 24–37 × 5.5–7.5 μm, clavate, (2–)4-spored, sterigmata up to 4.5 μm long. Cystidioid cells in hymenium absent. Hymenophoral trama regular, composed of cylindrical hyphae, 4–10 μm wide, colorless in water and 5% KOH. Pileipellis a cutis, composed of cylindrical hyphae, 4–10 µm wide, colorless or yellowish in water and 5% KOH. Stipitipellis a cutis, composed of parallel hyphae, 2–6 µm wide, colorless in water and 5% KOH. Clamps absent in all tissues.

Figure 3. 

Microscopic features A, C Tricholoma flavoviride B, D T. fumeobrunneum E, G T. parafulvum F, H T. viscidum A, B, E, F basidiospores C, D, G, H basidia. Scale bars: 5 μm.

Ecology and habitat

Scattered or gregarious on the ground in broadleaf forest dominated by Quercus sp., currently only known from Shanxi Province, northern China.

Figure 4. 

Photographs of basidiomata in their natural habita A, B Tricholoma ammophilum (HSA 371) C T. argyraceum (BJTC FM3571) D T. aurantium (BJTC FM2195) E T. bakamatsutake (BJTC FM3441) F T. bonii (BJTC FM905) G T. boudieri (BJTC FM2752) H T. cingulatum (BJTC FM1172) I T. frondosae (BJTC FM3895) J T. imbricatum (BJTC FM1170) K T. lishanense (BJTC FM1735) L T. populinum (BJTC FM1144). Scale bars: 1 cm.

Additional specimens examined

China • Shanxi Province, Pu County, Wulu Mountain, on the ground in broadleaf forest dominated by Quercus sp., 28 September 2023, J.Z Cao, CF2196 (BJTC FM3966), GenBank Acc. No.: ITS = PQ499559, mtSSU = PQ499563, rpb2 = PQ509906, mcm7 = PQ509916; ibid., Qinshui County, Lishan Mountain, on the ground in broadleaf forest dominated by Quercus sp., 7 September 2023, N. Mao, MNM829 (BJTC FM3512), GenBank Acc. No.: ITS = PQ499561, mtSSU = PQ499562, rpb2 = PQ509905, mcm7 = PQ509915.

Figure 5. 

Photographs of basidiomata in their natural habitat A Tricholoma psammopus (BJTC FM814) B T. saponaceum (BJTC FM2773) C T. atrosquamosum (BJTC FM4346) D T. terreum (HSA 419) E T. triste (BJTC FM3817) F T. vaccinum (BJTC FM943). Scale bars: 1 cm.

Notes

Tricholoma flavoviride belongs to the sect. Tricholoma (Fig. 1). Tricholoma citrinum, a species recently described from Yunnan Province of southwestern China (Cui et al. 2022), is morphologically and phylogenetically closely related to the new species. Both species have brownish-yellow pileus and ellipsoid basidiospores. However, T. citrinum is distinguished from T. flavoviride by its relatively paler pileus, stipe sometimes smooth, and habitat association with pine forests. Morphologically, T. flavoviride is also easily confused with T. equestre (L.) P. Kumm., and T. frondosae Kalamees & Shchukin. Tricholoma equestre is distinguished from T. flavoviride by its larger basidiospores (6.5–9 × 4–5.5 μm) and habitat associated with pine forests; T. frondosae by its paler pileus and larger basidiospores (7–8.5 × 4–5 μm) (Christensen and Heilmann-Clausen 2013; Reschke et al. 2018; Cui et al. 2022).

Tricholoma fumeobrunneum L. Fan, N. Mao & J.J Yang, sp. nov.

MycoBank No: MB 856306
Figs 2C, D, 3B, D

Diagnosis

It is distinguished by the combination of the following features: pale gray to grayish-brown pileus, smooth stipe, and broadly ellipsoid to ellipsoid basidiospores. It is most similar to T. sinopardinum Zhu L. Yang, X.X. Ding, G. Kost & Rexer but differs by the smaller basidiospores (6.5–8.5 × 5–6.5 μm).

Holotype

China • Shanxi Province, Lingchuan County, Lishan Mountain, 35°36'6"N, 113°20'32"E, elev. 1185 m, on the ground in broadleaf forest dominated by Quercus sp., 28 August 2023, Y. Li, MS492 (BJTC FM3436), GenBank Acc. No.: ITS = PQ499553, mtSSU = PQ499565, tef1-α = PQ509897, rpb2 = PQ509908, mcm7 = PQ509918.

Etymology

fumeobrunneum (Lat.): referring to the cap color of the basidiomata.

Description

Basidiomata small to medium-sized. Pileus 25–77 mm diam., at first convex, later broadly convex, plano-convex to applanate with age, with an umbo at center; surface dry, pale gray (#cac7c0) to grayish-brown (#9d8166), center darker to dark gray (#aaa9a3) to grayish black (#6b6157) when mature, covered with dark gray (#8e8c95) reflexed fibrillose squamules; margin at first involute, later incurved, cracking with age. Lamellae sinuate, moderately crowded, dirty white (#f9f8e9) to pale yellow (#f1eeca), turning yellowish brown (#e7dbb2) to brown (#d6bb9c) with age; lamellulae in 2–4 tiers, concolorous with lamellae. Stipe 50–76 mm long, 7–15 mm diam., cylindrical to clavate, enlarged downwards, dry, white (#f6f7f7), pale yellow (#f1ffe3) to yellowish brown (#ab9065), smooth. Context white (#f9f9f9), up to 3.3 mm. Odor unrecorded. Taste not recorded.

Basidiospores [120/3/4] 6.5–8.5 × 5–6.5 μm, Q = 1.3–1.6, broadly ellipsoid to ellipsoid, smooth, inamyloid, usually containing one large oil droplet. Basidia 34–46 × 6–9 μm, clavate, (2–)4-spored, sterigmata up to 6 μm long. Cystidioid cells in hymenium absent. Hymenophoral trama regular, composed of cylindrical hyphae, 3–12 μm wide, colorless in water and 5% KOH. Pileipellis a cutis, composed of cylindrical hyphae, 3–10 µm wide, colorless or yellowish-brown in water and 5% KOH. Stipitipellis a cutis, composed of parallel hyphae, 1.5–6 µm wide, colorless in water and 5% KOH. Clamps common in all tissues.

Ecology and habitat

Scattered or gregarious on the ground in broadleaf forest dominated by Quercus sp., currently only known from Shanxi Province, northern China.

Other specimens examined

China • Shanxi Province, Lingchuan County, Lishan Mountain, on the ground in broadleaf forest dominated by Quercus sp., 5 September 2023, N. Mao, MNM891 (BJTC FM3571), GenBank Acc. No.: ITS = PQ499554; ibid., Lingchuan County, Lishan Mountain, 35°36'6"N, 113°20'32"E, elev. 1190 m, on the ground in a broadleaf forest dominated by Quercus sp., 28 August 2023 H.M. Ji, MS492 (BJTC FM3445), GenBank Acc. No.: ITS = PQ499552, mtSSU = PQ499566, tef1-α = PQ509898, rpb2 = PQ509909, mcm7 = PQ509919; ibid., 5 September 2023, N. Mao, MNM894 (BJTC FM3574), GenBank Acc. No.: ITS = PQ499555, mtSSU = PQ499567, tef1-α = PQ509899, mcm7 = PQ509920.

Notes

Tricholoma fumeobrunneum belongs to the sect. Pardinicutis (Fig. 1). Tricholoma fumeobrunneum was phylogenetically sister to T. sinopardinum, a species found in Xizang Autonomous, southwestern China. However, T. sinopardinum differs from the new species by its stipe covered with brownish, brown to dark brown fibrillose to reflexed squamules and larger basidiospores (8.5–10.5 × 6.5–7.5 μm) (Yang et al. 2017). Tricholoma highlandense Zhu L. Yang, X.X. Ding, G. Kost & Rexer, another species from China in this section, is easily distinguished from T. fumeobrunneum by its pileus without a tinge of gray and occurrence in red acid soil (Yang et al. 2017).

Tricholoma parafulvum L. Fan, N. Mao & J.J Yang, sp. nov.

MycoBank No: MB 856307
Figs 2E, F, 3E, G

Diagnosis

It is distinguished by the combination of the following features: yellowish brown, brown to dark brown pileus, stipe surface covered with brown fibrils, and broadly ellipsoid to ellipsoid basidiospores. It is most similar to Tricholoma fulvum (DC.) Bigeard & H. Guill. but differs in the pileus surface without being radially streaky.

Holotype

China • Shanxi Province, Qinshui County, Lishan Mountain, 35°36'5"N, 113°20'29"E, elev. 1600 m, on the ground in broadleaf forest dominated by Quercus sp., 5 October 2023, J.Z. Cao, CF2300 (BJTC FM4065), GenBank Acc. No.: ITS = PQ499537, mtSSU = PQ499568, tef1-α = PQ509900, rpb2 = PQ509910, mcm7 = PQ509921.

Etymology

Parafulvum (Lat.): referring to the fact that the new species is very similar to Tricholoma fulvum in basidiomatal appearance.

Description

Basidiomata small to medium-sized. Pileus 30–66 mm diam., at first hemispherical to convex, later plano-convex to applanate with age, with depression or an umbo at center; surface dry, slightly viscid when wet, yellowish brown (#edd6b8), brown (#d49d76) to dark brown (#b78877), darker in the center and paler towards the margin, covered with brown (#a1735b) to dark brown (#845546) reflexed fibrillose squamules; margin at first involute, later incurved, cracking with age. Lamellae sinuate, moderately crowded, yellow (#f3edd7) to yellowish brown (#ddb68b) when young, turning dark brown (#c1986e) with age; lamellulae in 2–3 tiers, concolorous with lamellae. Stipe 42–83 mm long, 5–22 mm diam., cylindrical to clavate, enlarge downwards, dry, white (#f5fae3), yellowish brown (#c1a36f) to brown (#9d7963), covered with brown (#7e5c43) fibrils. Context white (#f7f8f7), up to 11 mm. Odor unrecorded. Taste not recorded.

Basidiospores [90/3/3] 4.5–6 × 3.5–4.5 μm, Q = 1.3–1.4, broadly ellipsoid to ellipsoid, smooth, inamyloid, usually containing one large oil droplet. Basidia 23–37 × 4.5–7 μm, clavate, (2–)4-spored, sterigmata up to 5.5 μm long. Cystidioid cells in hymenium absent. Hymenophoral trama regular, composed of cylindrical hyphae, 3–7 μm wide, colorless in water and 5% KOH. Pileipellis a cutis, composed of cylindrical hyphae, 3–8 µm wide, colorless or yellowish-brown in water and 5% KOH. Stipitipellis a cutis, composed of parallel hyphae, 2–7 µm wide, colorless in water and 5% KOH. Clamps absent in all parts of basidioma.

Ecology and habitat

Scattered or gregarious on the ground in broadleaf forest dominated by Quercus sp. or Betula sp., currently only known from Shanxi Province, northern China.

Additional specimens examined

China • Shanxi Province, Qinshui County, Lishan Mountain, on the ground in broadleaf forest dominated by Quercus sp., 5 October 2023, J.Z. Cao, MS855 (BJTC FM4210), GenBank Acc. No.: ITS = PQ499538, mtSSU = PQ499569, tef1-α = PQ509901, rpb2 = PQ509911, mcm7 = PQ509922; ibid., Loufan County, Yundingshan Mountain, on the ground in broadleaf forest dominated by Betula sp., 23 August 2022, N. Mao, MNM738 (BJTC FM2500), GenBank Acc. No.: ITS = PQ499536.

Notes

Tricholoma parafulvum belongs to sect. Genuina (Fig. 1). Tricholoma fulvum is morphologically similar and phylogenetically closely related to the new species. They all have brownish caps and ellipsoid basidiospores. However, T. fulvum differs from T. parafulvum by its pileus with radially streaky and relatively larger spores (5.8–7.2 × 4.6–5.1 μm) (Christensen and Heilmann-Clausen 2013). Four species from this section are also found in Shanxi Province, i.e., Tricholoma aurantium, T. ammophilum, T. populinum and T. viscidum. Of them, Tricholoma viscidum is similar to T. parafulvum, both of which are associated with Quercus spp. However, T. viscidum differs from T. parafulvum by its pileus being extremely viscid when wet and relatively larger spores (5–6.5 × 4.5–5.5 μm). The remaining three species are easily distinguished from T. parafulvum, T. ammophilum and T. populinum by their habitats associated with Populus spp., T. aurantium by its stipe surface covered with dense, orange to brownish orange scaly (Heilmann-Clausen et al. 2017).

Tricholoma viscidum L. Fan, N. Mao & J.J Yang, sp. nov.

MycoBank No: MB 856308
Figs 2G, H, 3F, H

Diagnosis

It is distinguished by the combination of the following features: pileus surface viscid when wet, stipe surface covered with brown to dark brown fibrils, and broadly ellipsoid basidiospores. It is most similar to T. ustaloides Romagn. but differs in the absence of a sharply defined zone of white color on the upper part of the stipe surface.

Holotype

China • Shanxi Province, Qinshui County, Lishan Mountain, 35°29'8"N, 113°1'19"E, elev. 1658 m, on the ground in broadleaf forest dominated by Quercus sp., 29 September 2023, J.Z. Cao, MS843 (BJTC FM4198), GenBank Acc. No.: ITS = PQ499539, mtSSU = PQ499571, tef1-α = PQ509903, rpb2 = PQ509913, mcm7 = PQ509924.

Etymology

viscidum (Lat.): referring to the viscid cap of basidiomata when wet.

Description

Basidiomata small, medium to large-sized. Pileus 33–86 mm diam., at first hemispherical to convex, later plano-convex with age, with an umbo at center; surface viscid when wet, yellowish brown (#e6d3b8), brown (#d0b79d) to reddish brown (#9d6463), darker in the center and paler towards the margin, covered with brown (#ab7d6b) fibrils; margin at first involute, later incurved, cracking with age. Lamellae sinuate, crowded, dirty white (#f3fbd0) to pale yellow (#f1ffb8), turning brown (#c09a7e) with age; lamellulae in 2–3 tiers, concolorous with lamellae. Stipe 49–113 mm long, 10–18 mm diam., cylindrical to clavate, enlarged downwards, dry, yellowish brown (#c79f5c) to brown (#9f6327), covered with brown (#b99976) to dark brown (#8b6c68) fibrils. Context white (#f3eeea), up to 13 mm. Odor unrecorded. Taste not recorded.

Basidiospores [120/3/4] 5–6.5 × 4.5–5.5 μm, Q = 1.1–1.25, broadly ellipsoid to ellipsoid, smooth, inamyloid, usually containing one large oil droplet. Basidia 28–37 × 5–8 μm, clavate, (2–)4-spored, sterigmata up to 5.5 μm long. Cystidioid cells in hymenium absent. Hymenophoral trama regular, composed of cylindrical hyphae, 5–10.5 μm wide, colorless in water and 5% KOH. Pileipellis a cutis, composed of cylindrical hyphae, 4–9.5 µm wide, yellowish-brown in water and 5% KOH. Stipitipellis a cutis, composed of parallel hyphae, 2.5–6 µm wide, colorless in water and 5% KOH. Clamps absent in tissues.

Ecology and habitat

Scattered or gregarious on the ground in broadleaf forest dominated by Quercus sp., currently only known from Shanxi Province, northern China.

Additional specimens examined

China • Shanxi Province, Qinshui County, Lishan Mountain, on the ground in broadleaf forest dominated by Quercus sp., 5 October 2023, J.Z. Cao, CF2268 (BJTC FM4038), GenBank Acc. No.: ITS = PQ499541; ibid., 29 September 2023, J.Z. Cao, MS844 (BJTC FM4199), GenBank Acc. No.: ITS = PQ499542, mtSSU = PQ499572, tef1-α = PQ509904, rpb2 = PQ509914, mcm7 = PQ509925; ibid., 25 August 2023, H.M. Ji MS443 (BJTC FM3388), GenBank Acc. No.: ITS = PQ499540, mtSSU = PQ499570, tef1-α = PQ509902, rpb2 = PQ509912, mcm7 = PQ509923.

Notes

Tricholoma viscidum belongs to the sect. Genuina (Fig. 1). Tricholoma ustaloides is closely related to the new species in morphology and phylogeny. They all have ellipsoid basidiospores and are associated with the Quercus spp. However, T. ustaloides differs from T. viscidum by its upper part of the stipe decorated with a distinctly and sharply delimited zone (Christensen and Heilmann-Clausen 2013; Halama et al. 2016). Tricholoma ustale (Fr.) P. Kumm., a European species often associated with Fagus spp. and Carpinus spp., is also easily confused with the new species as there are some reports that it is related to Quercus forests. However, T. ustale differs by its stipe flesh turning reddish brown when cut or bruised and its phylogenetic position (Christensen and Heilmann-Clausen 2013).

Discussion

Shanxi Province is located in northern China, where the climate ranges from subtropical to cold temperate. A total of 21 species of Tricholoma were confirmed in Shanxi Province in this study (Figs 1–5) based on morphological and molecular data, including four new species described in this paper. They are T. ammophilum A.D. Parker, Grubisha & S.A. Trudell, T. argyraceum (Bull.) Gillet, T. atrosquamosum Sacc., T. aurantium (Schaeff.) Ricken, T. bakamatsutake Hongo, T. bonii Basso & Candusso, T. boudieri Barla, T. cingulatum (Almfelt ex Fr.) Jacobasch, T. flavoviride, T. frondosae, T. fumeobrunneum, T. imbricatum (Fr.) P. Kumm., T. lishanense L. Fan & J.J. Yang, T. parafulvum, T. populinum J.E. Lange, T. psammopus (Kalchbr.) Quél., T. saponaceum (Fr.) P. Kumm., T. terreum (Schaeff.) P. Kumm, T. triste (Scop.) Quél., T. vaccinum (Schaeff.) P. Kumm., and T. viscidum. That means Shanxi Province is rich in species diversity of the genus Tricholoma.

Before the present study, there were nine Tricholoma species reported in Shanxi Province according to the related professional fungal literatures (Cao et al. 1990; Liu 1991; Mao 2000). Of them, five species are confirmed by the present study, including T. argyraceum, T. cingulatum, T. populinum, T. terreum, T. vaccinum. Of the remaining four species, T. gambosum is a species of Calocybe currently as Calocybe gambosa, which is still not yet confirmed from this province; T. matsutake was reported based on the specimens under the tree of Picea sp. from Guancenshan Mountain in late 80th of last century (Cao et al. 1990). We are not able to examine the specimens cited by Cao et al. (1990), and also did not obtain new collections, so its occurrence is not confirmed; T. virgatum (Fr.) P. Kumm. was reported by Mao (2000), but no specimen was cited. According to the picture of basidiomata cited in his book, we suspect this name is probably misapplied for T. bonii; T. zelleri (D.E. Stuntz & A.H. Sm.) Ovrebo & Tylutki was reported by Liu (1991) from Qinshui County in southern Shanxi Province under Pinus armandii and P. tabuliformis, this name has already been treated as a synonym of Tricholoma focale (Fr.) Ricken by Kuo (2018). Tricholoma focale is a distinct species, that is morphologically very similar to the new species T. viscidum described in the present study, but we did not find it again for the time being.

Many species of Tricholoma have been observed having exclusively host association in Shanxi Province in this study; for example, both T. bonii and T. psammopus are associated with Larix gmelinii var. principis-rupprechtii only, T. vaccinum with Picea spp. only, T. populinum with Populus sp. only, T. terreum and T. imbricatum with Pinus tabuliformis only, some species, viz. T. atrosquamosum, T. bakamatsutake, T. fumeobrunneum, T. lishanense and T. viscidum with Quercus spp. only. A few species have a relatively wide host range. Tricholoma cingulatum grows under the trees of Populus spp. or Quercus sp., T. aurantium grows under Quercus sp. or Picea sp., T. argyraceum appears under Pinus tabuliformis, P. sylvestris, Betula sp. and Quercus sp. These host specificities are a useful guide for identifying the Tricholoma in the field.

The distribution of Tricholoma is significantly influenced by the geography of their host and the climate in Shanxi Province. There are two main geographic zones to be recognized in Shanxi Province: i) the subalpine zone with vegetation composed of coniferous Larix-Picea and broadleaf Populus-Betula and with a cold climate, which is mainly located in the mountain area above an altitude of 1600 m and some northern regions; ii) the warm zone with the vegetation composed of conifer’s Pinus spp. and broadleaf Quercus spp. and with a warm climate, which is mapped in most of the central and southern Shanxi Province. Some Tricholoma species are only distributed in the subalpine zone, including T. bonii, T. populinum, T. psammopus, T. triste and T. vaccinum. Only a few species are observed appearing in both subalpine and warm zones, such as T. argyraceum, T. aurantium, and T. cingulatum. In contrast, the warm zone is rich in species of this genus, with at least 13 species found that are only distributed in this zone, including the most popularly encountered T. terreum, which is often confused with the northern T. bonii.

The climate also clearly shaped the geographic map of the Tricholoma species in this province. One of the good samples is the species T. bonii, an exclusive Larix-associated species in this province. It is the most popularly encountered mushroom in the forest of Larix gmelinii var. principis-rupprechtii in the northern region, but in the southern area, it is completely absent although there are many plantations of L. gmelinii var. principis-rupprechtii colonized for more than 50 years. Another example is T. lishanense, one of the most common Tricholoma in Lishan Mountain under Quercus spp.; its distribution is limited to Zhongtiao Mountains, where there are some subtropical climate areas.

Acknowledgements

We thank Dr. J.Z. Cao, who collected specimens and provided valuable suggestions.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

The study was supported by the National Natural Science Foundation of China (No. 32370010, 31750001), the Beijing Natural Science Foundation (No. 5172003), the BJAST Budding Talent Program (Grant No. 24CE-BGS-19).

Author contributions

LF conceived and designed the study; NM wrote the manuscript and conducted phylogenetic analyses; NM and JJY conducted morphological observations; YXZ and LT conducted the experiments.

Author ORCIDs

Ning Mao https://orcid.org/0000-0003-1564-9446

Jia-Jia Yang https://orcid.org/0000-0003-3668-6107

Yu-Xin Zhang https://orcid.org/0000-0002-6757-1903

Ting Li https://orcid.org/0000-0001-7809-3217

Li Fan https://orcid.org/0000-0001-9887-7086

Data availability

All of the data that support the findings of this study are available in the main text.

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