Corresponding author: Zhu L. Yang (
Academic editor: M.P. Martín
Phylogenetically, the genera
He Z-M, Yang ZL (2021) A new clitocyboid genus
The widespread genus
Based on phylogenetic analyses over the past 20 years, (i) many new genera within the Tricholomatoid clade were proposed to accommodate previous
The molecular phylogenetic relationships among members of the
Recently, some collections were shown to be closely related to
Twenty-three specimens of species similar to
Specimens used in phylogenetic analysis and their GenBank accession numbers. The newly generated sequences are shown in bold.
Species | Voucher | Locality | GenBank accession number | |||||
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nrLSU |
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MDF352 | Costa Rica |
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DIC595b | Brazil |
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MDF058 | Ecuador |
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China: Jilin |
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AFTOL-ID 542 |
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DJL06TN40 | USA |
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TUB 011588 |
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Lueck2 | Germany |
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CBS 398.79 | France |
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AFTOL-ID 1017 | USA |
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AH24539 | Spain |
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DJL-BZ-1883 | Belize |
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TENN63483 | India |
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Iqbal-568 | Bangladesh |
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AFTOL-ID 1684 | USA |
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GE18008-1 | France |
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GE18035 | Canada |
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GE18033 | Canada |
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AFTOL-ID 1523 | USA |
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OSC 113683 |
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DJL06TN80 | USA |
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G0171 | Hungary |
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14562 | Italy |
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4804 | Russia |
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G0460 | Hungary |
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NAMA 2015-206 | USA |
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NAMA 2015-318 | USA |
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Mushroom Observer 302917 | USA |
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DIC215 | Ecuador |
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DIC110 | Bolivia |
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DIC119 | Ecuador |
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DIC146 | Bolivia |
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MDF017 | Bolivia |
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MDF200 | Brazil |
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Lueck7 | Germany |
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DJL-Scot-8 | UK |
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CFMR PR-6601 | Puerto Rico |
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AFTOL-ID 1682 | USA |
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China: Tibet |
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CDS 41163 | Ecuador |
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DIC115 | Guatemala |
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Ertz 10475 | Portugal |
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DIC113 | Brazil |
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G1063 | Poland |
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CFMR DEN-25 | Denmark |
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TJB-10048 (CORT) | Australia |
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CFMR PR-5408 | Puerto Rico |
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GDGM41710 | China: Jilin |
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Herb. Roux n. 4044 | France |
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Herb. Roux n. 3666 | Moldova |
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A. Gminder 971488 | Italy |
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E. C. 08191 | Italy |
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JM96/33 |
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AFTOL-ID 1727 | USA |
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QCAM6000 | Ecuador |
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CFMR BZ-3923 | Belize |
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AFTOL-ID 2020 | USA |
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AFTOL-ID 1997 | Puerto Rico |
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FO 46714 |
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CFMR NC-256 | USA |
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AFTOL-ID 1715 | USA |
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AFTOL-ID 729 |
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US97/138 | Germany |
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GDGM70059 | USA |
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China: Tibet |
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China: Tibet |
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China: Yunnan |
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China: Yunnan |
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China: Tibet |
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Germany |
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AFTOL-ID 1723 | USA |
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China: Yunnan |
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China: Jilin |
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China: Tibet |
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China: Tibet |
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GAL18249 | USA |
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S-270-FB1 | Japan |
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GWG H. |
UK |
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GDGM 44492 | China |
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DJL05TN62 (TENN) | USA |
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GRSM 77065 | USA |
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China: Yunnan |
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China: Yunnan |
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China: Yunnan |
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China: Yunnan |
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China: Yunnan |
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China: Yunnan |
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China: Yunnan |
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China: Yunnan |
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China: Yunnan |
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China: Jilin |
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China: Yunnan |
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CFMR ENG-3 | UK |
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DJL05TN80 (TENN) | USA |
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TJB-10034 (CORT) | Thailand |
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AFTOL-ID 1557 | USA |
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China |
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GDGM 50075 | China: Hunan |
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GDGM 43351 | China: Sichuan |
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CFMR HHB-2808 | USA |
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DAOM 226047 | Canada |
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IO. 14. 214 | Spain |
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IO. 14. 177 | Sweden |
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IO. 14. 167 | Sweden |
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IO.14. 200 | France |
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IO. 14. 196 | Spain |
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AFTOL-ID 773 |
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UPS F-611822 | Sweden |
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AFTOL-ID 1924 | USA |
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ZRL20151311 |
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IO. 14. 142 | Sweden |
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AFTOL-ID 536 | USA |
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CFMR L-8014 | USA |
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AFTOL-ID 537 |
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ZRL20151760 |
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IO. 15. 122 | Spain |
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IO. 14. 92 | Sweden |
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IO. 14. 165 | Sweden |
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Macroscopic characters of species were described based on the raw field record data and photographs. Colors used in description referred to
Total genomic DNA was extracted using the Ezup Column
The PCR mixtures contained 1× PCR buffer, 1.5mM MgCl2, 0.2mM dNTPs, each primer at 0.4 μM, 1.25U of
Sequences used for phylogenetic analysis (presented in Table
As shown in Table
The combined dataset
In order to accurately determine the position of
In addition, an
Phylogram showing the phylogenetic relationships among
Characterized generally by clitocyboid basidiomes, convex to funnel-shaped pileus, decurrent lamellae, absence of veils, inamyloid basidiospores and presence of clamps.
From the type genus
Basidiomes small, medium-sized to large, mostly clitocyboid, rarely omphalinoid or mycenoid; veils absent. Pileus convex, applanate to funnel-shaped; surface usually dry, smooth, lubricous or rarely viscid. Lamellae decurrent to deeply decurrent. Basidiospores ellipsoid, oblong or subglobose, thin-walled and inamyloid. Pileipellis usually a cutis, sometimes ixocutis or trichoderm. Lamellar trama regular, subregular, interwoven or bidirectional. Clamp connections present.
Usually gregarious or caespitose on ground, rarely on wood; widespread in temperate and tropical regions.
The genera
Differs from
Basidiomes small, clitocyboid. Pileus convex, applanate to infundibuliform; surface dry, greyish (2B1), grey-brown (5C4) to dark grey-brown (5E4); center depressed with age. Lamellae decurrent to deeply decurrent, white (1A1) to cream (1A2), thin, moderately crowded, sometimes furcate and interveined. Stipe central, subcylindrical, concolorous with pileus. Basidiospores ellipsoid, oblong to cylindrical, colourless, hyaline, smooth, thin-walled, inamyloid; ratio of basidia to basidiospore length less than 5. Pileipellis and stipitipellis a cutis. Lamellar trama subregular. Clamp connections abundant, present in all parts of basidiome.
Saprophytic, usually gregarious or caespitose on the ground of coniferous or coniferous and broad-leaved mixed forest; distributed in the temperate and subtropical zones from June to November.
Differs from
China. Yunnan Province: Kunming City, near Yeya Lake, at
Basidiomes small, clitocyboid. Pileus 0.5–2 cm in diam, at first nearly applanate, then concave; surface dry and rugose, gray-brown (5E2-4) to gray-black (4F2-4) in the center and gray-brown (5C2-4) or gray (5B1-2) towards margin; center often slightly umbonate; margin straight and undulating; context thin and white (1A1) to cream (1A2). Lamellae deeply decurrent, white (1A1) to cream (1A2), thin (up to 2 mm high), crowded, sometimes forked and intervenose. Stipe 2.5–6 × 0.2–0.4 cm, central, narrowly cylindrical to subcylindrical, sometimes flexuous, hollow; surface dry and nearly smooth, concolorous with pileus; context white (1A1).
Basidiospores [60/3/3] 5–6 (6.5) × (2.5)3–3.5(4) μm, Q = (1.38)1.55–1.95(2), Qm = 1.73 ± 0.14, elongate, colorless, hyaline, smooth, thin-walled, inamyloid. Basidia 20–24 × 5–6 μm, clavate, 4-spored, colorless, hyaline, thin-walled; sterigmata up to 4 μm long; ratio of basidia to basidiospore length values about 3–5. Cystidia absent. Lamellar trama subregular; hyphae colorless, hyaline, cylindrical, thin-walled, 3–10 µm wide. Pileipellis a cutis, but in places upright or trichodermial in appearance, made up with thin-walled cylindrical hyphae 3–9 µm wide. Stipitipellis a cutis, composed of thin-walled cylindrical hyphae 3–10 μm wide. Clamp connections present in all parts of basidiome.
Basidiomes of described
Gregarious or caespitose, growing saprotrophically in forest litter, often under conifers, on the ground, known from subtropical zone of Yunnan, China; from July to October.
China. Yunnan Province: Dali Bai Autonomous Prefecture, Yunlong Country, Tianchi National Nature Reserve, at
Differs from
China. Yunnan Province: Baoshan City, Longyang District, Shuizhai Village, at
Basidiomes small, clitocyboid. Pileus 1.5–3 cm in diam, plano-convex to funnel-shaped; surface dry and nearly smooth, greyish-brown (4B2-3) to grey-brown (4E3-5); center depressed, usually with a low umbo, somewhat darker; margin generally straight and undulating, incurved when old; context thin and white (1A1). Lamellae deeply decurrent, white (1A1) to cream (1A2), thin, 1–2 mm high, relatively crowded, sometimes forked and intervenose. Stipe 1–3 × 0.2–0.4 cm, central, subcylindrical, hollow; surface dry and nearly smooth, concolorous with pileus; context white (1A1).
Microscopic features of
Basidiospores [60/3/3] (7)7.5–10.5(11.5) × 3–4 μm, Q = (2.05)2.11–3(3.33), Qm = 2.56 ± 0.3, cylindrical, colorless, hyaline, smooth, thin-walled, inamyloid. Basidia 20–30 × 4–5.5 μm, clavate, 2-spored, colorless, hyaline, thin-walled; sterigmata up to 10 μm long; ratio of basidia to basidiospore length less than 5 (about 2–4). Cystidia absent. Lamellar trama subregular, colorless, hyaline, made up of thin-walled cylindrical hyphae with 3–10 µm wide. Pileipellis a cutis, composed of thin-walled cylindrical hyphae 3–11 µm wide. Stipitipellis a cutis, composed of thin-walled cylindrical hyphae 3–10 μm wide. Clamp connections in all parts of basidiomes.
Saprophytic, usually gregarious on the ground of coniferous or coniferous and broad-leaved mixed forest, known from Yunnan, China; July to September.
China. Yunnan Province: Kunming City, Qipan Mountain, at
Microscopic features of
In our current study, the new clitocyboid species were clustered into a monophyletic lineage (
Besides the molecular analyses, morphological data also support its separation from the relative genera.
For a long time,
In previous studies,
The authors are very grateful to their colleagues at Kunming Institute of Botany, Chinese Academy of Sciences, including Drs. Xiang-Hua Wang, Jiao Qin, Bang Feng, Qi Zhao and Master students Hua Qu and Si-Peng Jian for collecting and providing specimens; and Drs. Gang Wu, Yang-Yang Cui, Qing Cai for providing help on morphological observation and phylogenetic analysis. This study was financed by Yunnan Ten-Thousand-Talents Plan – Yunling Scholar Project and Postdoctoral Directional Training Foundation of Yunnan Province.
Alignment of
fasta file
Alignment of
fasta file
Alignment of
fasta file