Research Article |
Corresponding author: Jun-Qing Yan ( yanjunqing1990@jxau.edu.cn ) Academic editor: Olivier Raspé
© 2025 Lin-Gen Chen, Hong Chen, Ling Ding, Yu-Qin Xu, Hui Zeng, Sheng-Nan Wang, Jun-Qing Yan.
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:
Chen L-G, Chen H, Ding L, Xu Y-Q, Zeng H, Wang S-N, Yan J-Q (2025) Four new species of Entoloma subgenus Cyanula (Entolomataceae, Agaricales) from subtropical regions of China. MycoKeys 116: 303-325. https://doi.org/10.3897/mycokeys.116.145568
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In this study, four species of Entoloma subgenus Cyanula (E. orientosinense, E. subgriseosquamulosum, E. subpraegracile, and E. wuyishanense) from subtropical regions of China, are described as new to science based on morphological and phylogenetic analyses. Morphologically, E. orientosinense is characterized by the white basidiomata, relatively large basidiospores, and carneogriseum-type lamellae edge; E. subgriseosquamulosum is recognized by the fuscous pileus, crowded and adnate lamellae, and medium-sized basidiospores; E. subpraegracile is identified by the yellow pileus and intervenose lamellae with sterile or heterogeneous edge; E. wuyishanense is distinct by the blue basidiomata and fertile lamellae edge with slightly bluish pigmentation near the stipe. Entoloma orientosinense belongs to sect. Caesiocincta, subsect. Queletia, and E. wuyishanense belongs to sect. Poliopodes. The remaining two species each form independent branches and do not belong to any known sections. Detailed descriptions, color photos, and a key to related species are presented.
Basidiomycetes, new taxa, phylogeny, taxonomy
Entoloma (Fr.) P. Kumm. is one of the most diverse genera within Agaricales, well-characterized by pink to brownish spore prints and angular basidiospores viewed in all views (
In the past, based on morphological taxonomy, Cyanuli was introduced by
The species of Entoloma subg. Cyanula are mainly characterized by their collybioid habit, vividly colorful (often blue, violaceous to brown) and squamulose pileus, absence of clamp connections, and presence of brilliant granules and intracellular pigments in hyphae. So far, at least 500–600 species of E. subg. Cyanula have been discovered worldwide.
According to previous studies, there are 13 species belonging to Entoloma subg. Cyanula in China, 7 of which were newly described (
The collection sites of the specimens in this study were all located in the subtropical region of East China, and these specimens were deposited in the Herbarium of Fungi, Jiangxi Agricultural University (
Genomic DNA was extracted from dried specimens with the NuClean Plant Genomic DNA kit (CWBIO, China) (
PCR amplification was conducted with a 25 μL reaction system as follows: 1 µL DNA, 1 µL each for forward and reverse primers, 9.5 µL ddH2O, and 12.5 µL 2 × Taq Master Mix (Dye Plus, Vazyme Biotechnology Co. Ltd., Nanjing City, China). PCR was carried out using a touchdown amplification procedure following
In total, 173 sequences (126 ITS sequences and 47 LSU sequences) of 126 samples were used for phylogenetic analyses based on Bayesian inference (BI) and Maximum likelihood (ML). The selection of sequences for the phylogenetic analyses was based on the results of ITS BLAST and of
Details of sequences used in the phylogenetic analyses. Newly generated sequences were in bold.
Species | Location | Voucher Number | GenBank No. (ITS) | GenBank No. (LSU) | Sequence origin |
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Entoloma albidosimulans | Australia | MEN 2004-065, isotype | — | MK277956 |
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E. albinellum | USA | TENN:070403 | KY777375 | — | Unpublished in GenBank |
E. argus | Vietnam | LE F-312694, holotype | OM987263 | OM996175 |
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E. argus | Vietnam | LE F-315916 | OM987264 | — |
|
E. arion | Vietnam | LE F-312691, holotype | OM987259 | OM996176 |
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E. arion | Vietnam | LE F-312692 | OM987260 | — |
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E. arion | Vietnam | LE F-315917 | OM987261 | — |
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E. asprellum | Estonia | TUF106064 | UDB011486 | — | UNITE |
E. atropapillatum | Brazil | FK0898, holotype | KF679354 | KF738940 |
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E. azureosquamulosum | China | HKAS53408 | JQ410334 | JQ410326 |
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E. azureosquamulosum | China | GDGM29254 | JQ410335 | — |
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E. azureosquamulosum | China | GDGM27355, holotype | NR_137086 | NG_059214 |
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E. caespitosum | China | GDGM27564 | JQ281477 | JQ320130 |
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E. caespitosum | China | GDGM24025 | JQ281490 | JQ410327 |
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E. caespitosum | China | GDGM24026 | JQ281491 | JQ320133 |
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E. calceus | Norway | O-F-259457, holotype | NR_182489 | — |
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E. calceus | France | LIP0402265 | ON008492 | — |
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E. calceus | Norway | JL12-19 | ON008493 | — |
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E. callipygmaeum | Russia | LE312488 | MZ145205 | — |
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E. callipygmaeum | Russia | LE312487 | MZ145206 | — |
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E. callipygmaeum | Russia | LE253784, holotype | MZ145207 | — |
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E. carneogriseum | Norway | O-F-256479 | UDB07673714 | — | UNITE |
E. cetratum | Sweden | LE311888, neotype | OL338280 | — |
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E. chalybeum | Russia | LE254353 | KC898445 | KC898500 |
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E. chalybeum | Denmark | TUF105760 | UDB034191 | — | UNITE |
E. consanguineum | New Zealand | PDD80751 | MW775252 | — | Unpublished in GenBank |
E. consanguineum | New Zealand | PDD80751 | MW775268 | — | Unpublished in GenBank |
E. coracis | Norway | O-F-256850, holotype | MW934571 | MW934251 |
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E. coracis | Norway | O-F-67255 | MW934572 | — |
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E. coracis | Norway | O-F-251952 | MW934573 | — |
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E. corvinum | France | FA4261 | OR419868 | — |
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E. cyanostipitum | China | GDGM31318, holotype | KY711237 | KY972694 |
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E. cyanostipitum | China | SAAS2239 | KY711238 | KY972695 |
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E. cyanostipitum | China | GDGM31294 | KY972700 | KY972693 |
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E. dislocatum | Spain | L0607565, holotype | ON008483 | — |
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E. dislocatum | Spain | SFC-080612-01 | ON008484 | — |
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E. dislocatum | Italy | TUF105920, paratype | UDB0799300 | — | UNITE |
E. exile | Germany | Lueck8 | KP965773 | KP965791 |
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E. exile | – | KM187354 | MF977976 | — | Unpublished in GenBank |
E. griseocyaneum | Russia | LE254351 | KC898444 | KC898498 |
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E. griseocyaneum | Germany | KaiR997 | MZ611684 | — |
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E. icarus | Vietnam | LE F-312696, holotype | OM987257 | OM996174 |
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E. icarus | Vietnam | LE F-312697 | OM987258 | — |
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E. incanum | Sweden | LE312503, neotype | OK161247 | OK161275 |
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E. incanum | Russia | LE311794 | OK161249 | OK161276 |
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E. incanum | Russia | LE315858 | OK161250 | — |
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E. isborscanum | Russia | LE312486 | MW934564 | — |
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E. isborscanum | Russia | LE302088, holotype | MW934566 | MW934253 |
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E. linkii | Norway | O-F-256353 | UDB07673651 | — | UNITE |
E. mastoideum | China | GDGM28820 | JQ281476 | JQ410328 |
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E. mastoideum | China | GDGM26597 | JQ291564 | JQ320126 |
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E. meridionale | Greece | ACAM2014-0127 | OL679698 | — |
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E. meridionale | Greece | ACAM2018-0152 | OL679699 | — |
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E. meridionale | Greece | ACAM2018-0153, holotype | OL679700 | — |
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E. minutigranulosum | Russia | LE312484 | MZ145210 | — |
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E. minutigranulosum | Russia | LE312483 | MZ145212 | — |
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E. minutigranulosum | Russia | LE302096, holotype | MZ145214 | — |
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E. mougeotii | Estonia | TUF106917 | UDB015645 | — | UNITE |
E. mougeotii | Estonia | TUF101633 | UDB016265 | — | UNITE |
E. mougeotii | Estonia | TUF106505 | UDB019720 | — | UNITE |
E. mutabilipes | Finland | TUR610/12 | LN850550 | — |
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E. mutabilipes | Estonia | TUR8788 | LN850551 | — |
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E. notabile | Cyprus | L-0607514, holotype | OL343537 | — |
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E. olivaceomarginatum | USA | PUL00036174 | ON561593 | — | Unpublished in GenBank |
E. orientosinense | China | HFJAU1414, holotype | PQ584686 | — | This work |
E. orientosinense | China | HFJAU1907 | PQ584690 | PQ584707 | This work |
E. orientosinense | China | HFJAU2616 | PQ584687 | — | This work |
E. orientosinense | China | HFJAU2920 | PQ584688 | PQ584708 | This work |
E. orientosinense | China | HFJAU4048 | PQ584689 | PQ584709 | This work |
E. pallidostriatum | Spain | L-0607566, holotype | NR_177630 | — |
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E. perasprellum | France | GC01100310, holotype | MZ145177 | — |
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E. perasprellum | Sweden | GB-0204547 / JBJ 19-107 | MZ145179 | — |
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E. perasprellum | Sweden | GB-0204548 / JBJ 19-122 | MZ145180 | — |
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E. perchalybeum | Sweden | GB-0209474, holotype | NR_182490 | — |
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E. perchalybeum | Finland | TUR190180 | ON008495 | — |
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E. poliopus | Estonia | TUF120264 | UDB024655 | — | UNITE |
E. praegracile | China | GDGM29251 | JQ281482 | JQ320129 |
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E. praegracile | China | GDGM29256 | JQ320107 | — |
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E. pseudocoelestinum | Germany | Lueck10 | KP965774 | KP965792 |
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E. pseudocoelestinum | – | KM132400 | MF977966 | — | Unpublished in GenBank |
E. pseudosubcorvinum | Thailand | SDBR-CMUNK0985, holotype | MZ215769 | MZ203540 |
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E. pseudosubcorvinum | Thailand | SDBR-CMUNK1367 | MZ215770 | MZ203541 |
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E. pulchripes | Russia | LE312485 | MZ145187 | — |
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E. pulchripes | Russia | LE311808, holotype | MZ145188 | — |
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E. pulchripes | Russia | LE311809 | MZ145189 | — |
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E. queletii | Turkey | OKA-TR1002 | MT741747 | — | Unpublished in GenBank |
E. queletii | Estonia | TUF141044 | UDB07674927 | — | UNITE |
E. riparium | Italy | L-0607563, holotype | NR_177632 | — |
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E. riparium | Estonia | TUF120259 | UDB024650 | — | UNITE |
E. septentrionale | Norway | O-F-254295, holotype | NR_174647 | — |
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E. sericeum | Germany | KaiR237 | OL338118 | OL338542 |
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E. sericeum | – | VHAs03 2 | DQ367430 | DQ367423 | Unpublished in GenBank |
E. serrulatum | Norway | O-F-158208/DMS-730296 | MZ869016 | — |
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E. serrulatum | Russia | LE254361 | KC898447 | KC898501 |
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E. serrulatum | Iran | EnSe-1 | KT833862 | — | Unpublished in GenBank |
E. sicoense | Portugal | PO F2244, holotype | OR026624 | — |
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E. sicoense | Portugal | PO F2245 | OR026625 | — |
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E. subcaesiocinctum | China | SAAS103 | KY711235 | KY972698 |
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E. subcaesiocinctum | China | SAAS133, holotype | KY711236 | KY972697 |
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E. subcorvinum | USA | MGW1494 | KY744168 | — | Unpublished in GenBank |
E. subcorvinum | USA | SAT1518905 | KY744169 | — | Unpublished in GenBank |
E. subgriseosquamulosum | China | HFJAU3967 | PQ584696 | — | This work |
E. subgriseosquamulosum | China | HFJAU3969, holotype | PQ584697 | PQ584721 | This work |
E. subpraegracile | China | HFJAU1822, holotype | PQ584698 | PQ584710 | This work |
E. subpraegracile | China | HFJAU3094 | PQ584706 | PQ584711 | This work |
E. subpraegracile | China | HFJAU3164 | PQ584700 | PQ584712 | This work |
E. subpraegracile | China | HFJAU3168 | PQ584705 | — | This work |
E. subpraegracile | China | HFJAU5110 | PQ584701 | — | This work |
E. subpraegracile | China | HFJAU5115 | PQ584699 | PQ584713 | This work |
E. subpraegracile | China | HFJAU5140 | PQ584702 | PQ584714 | This work |
E. subpraegracile | China | HFJAU5175 | PQ584703 | PQ584715 | This work |
E. subpraegracile | China | HFJAU5177 | PQ584704 | PQ584716 | This work |
E. subserrulatum | USA | TENN:068464 | KY744143 | — | Unpublished in GenBank |
E. subserrulatum | USA | TENN:070407 | KY744177 | — | Unpublished in GenBank |
E. subtenuicystidiatum | China | GDGM 28459, holotype | JQ320109 | JQ320116 |
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E. subtenuicystidiatum | China | GDGM 29246 | JQ320114 | JQ320132 |
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E. turci | Austria | WU25055 | UDB0802163 | — | UNITE |
E. viridomarginatum | The Netherlands | JAC15761 | MW775255 | — | Unpublished in GenBank |
E. viridomarginatum | The Netherlands | JAC12344 | MW775264 | — | Unpublished in GenBank |
E. wuyishanense | China | HFJAU3571, holotype | PQ584691 | — | This work |
E. wuyishanense | China | HFJAU3871 | PQ584692 | PQ584717 | This work |
E. wuyishanense | China | HFJAU3874 | PQ584694 | PQ584718 | This work |
E. wuyishanense | China | HFJAU3878 | PQ584693 | PQ584719 | This work |
E. wuyishanense | China | HFJAU3881 | PQ584695 | PQ584720 | This work |
A total of 2136 characters were used in subsequent analyses (ITS, 841 bp; LSU, 1,295 bp), of which 1382 were constant, 670 were parsimony-informative, and 84 were singleton. For Bayesian analysis, the average standard deviation of split frequencies was less than 0.01 after 20 million generations.
The results of the phylogenetic analysis were shown in Fig.
Refers to its type specimen originating from the eastern regions of China.
China • Anhui Province, Chizhou City, Shitan County, Guniujiang Nature Reserve, 30.0303°N, 117.5290°E, alt. 783 m, 9 October 2019, collected by Yu-Peng Ge, HFJAU1414.
Entoloma orientosinense is mainly characterized by the white, collybioid basidiomata, fibrillous and not striate pileus, narrow, adnate to decurrent lamellae, glabrous stipe, 5–6 angled basidiospores, sterile lamellae edge of carneogriseum-type, cylindrical to subclavate cheilocystidia, absence of cell pigments and clamp connections in hyphae. It differs from E. albinellum by its non-striate pileus, adnate to decurrent lamellae, and smaller basidiospores.
Basidiomata rather small, collybioid. Pileus 8–20 mm wide, convex then flattened with depressed center, with entire margin, slightly hygrophanous, fibrillous when young, then repent or raised scaly, not translucently striate, white (3A1–2). Lamellae moderately distant, 1.5–2.0 mm wide, with three types of lamellulae, adnate to decurrent, subventricose, initially white, then pink (11B4–6), with serrulate and concolourous edge. Stipe 20–25 × 2.0–3.0 mm, central, terete, tapered upwards, hollow, concolorous or paler with the pileus, minutely tomentose in the upper part elsewhere smooth and glabrous, base with white tomentum. Context thin, concolorous to the surface. Odor indistinct, taste not tested.
Basidiospores (8.5)9.3–11.0(12.5) × (6.0)6.5–8.0(9.0) μm, (av = 10.1 ×7.3 μm), Q = 1.2–1.6(1.7) (Qm = 1.4 ± 0.07, n = 200), heterodiametrical, 5–6 angles in profile view, thick-walled, inamyloid. Basidia 40–52 × 11–13 μm, clavate, 4-spored, sterigmata 5.0–10 μm long, clampless. Pleurocystidia absent. Lamellae edge sterile of carneogriseum-type. Cheilocystidia regularly dispersed in the lamellae edge, 17–47 × 4.0–7.0 μm, narrowly cylindrical to subclavate, septate, with slightly inflated apex. Lamellar trama regular, made up of cylindrical hyphae 7.0–13 µm wide. Pileipellis a cutis made up of cylindrical hyphae 8.0–11 μm broad, with transitions to a trichoderm towards the margin with clavate terminal elements 10–18 μm wide, not pigmented. Stipitipellis a cutis composed of densely arranged, cylindrical hyphae, up to 11 μm wide, slightly constricted at the septa, with acute or tapered end. Clamp connections absent.
Solitary or scattered on soil in mixed coniferous-broad-leaved forest, or on rotten wood, soil, and moss in broadleaved forest.
So far known from eastern China.
China • Fujian Province, Wuyishan City, 27.7139°N, 117.6533°E, alt. 1113 m, 27 June 2022, collected by Jun-Qing Yan and Bing-Ring Ke, HFJAU4048 • Zhejiang Province, Lishui City, Suichang County, Huangtakou Village, 28.2679°N, 118.9435°E, alt. 346 m, 12 July 2020, collected by Jun-Qing Yan and Yan-Liu Chen, HFJAU1907 • Qingtian County, Shigu Lake, 28.2063°N, 120.0415°E, alt. 1130 m, 31 July 2021, collected by Jun-Qing Yan, Bing-Ring Ke, and Zhi-Heng Zeng, HFJAU2616 • Nanyang Village, 27.9603°N, 120.0020°E, alt. 522 m, 6 August 2021, collected by Yu-Peng Ge and Lan-Yu Sun, HFJAU2920.
Morphologically, Entoloma orientosinense has much in common with E. albidosimulans G.M. Gates & Noordel. and E. albinellum with regard to the white and collybioid basidiomata. However, E. albidosimulans is distinct by its broader (up to 6 mm), adnate-emarginate lamellae, and belonging to E. subg. Alboleptonia species (
In the molecular data, E. orientosinense fits well within subg. Cyanula, sect. Caesiocincta, subsect. Queletia including E. albinellum and E. queletii. E. queletii is distinguished from new species by the vinaceous-pink pileus and larger basidiospores (10–13 × 6.5–9.0 μm) (
Refers to its morphology similar to “Entoloma griseosquamulosum”.
China • Fujian Province, Wuyishan City, Yangzhuang Town, Xiyuan Village, 27.7632°N, 117.8139°E, alt. 533 m, 26 June 2022, collected by Jun-Qing Yan, Cheng-Feng Nie, and Lin-Gen Chen, HFJAU3969.
Entoloma subgriseosquamulosum is mainly characterized by the rather small, collybioid basidiomata, fuscous pileus, crowded and adnate lamellae, glabrous stipe, medium-sized basidiospores with 5–6 angles, mostly 5 angles, and absence of clamp connections. It differs from E. griseosquamulosum G.M. Gates & Noordel. by its gray stipe, smaller basidiospores, and absence of brilliant granules in hyphae.
Basidiomata rather small, collybioid. Pileus 11–20 mm wide, campanulate to convex with slight depressed center, with entire margin, not hygrophanous, gray hairy scaly with denser center, translucently striate almost up to 1/2 of the radius, fuscous (4D2–4F2) to dark gray (1F1–4F1), darker at center. Lamellae relatively crowded, 2.0–4.0 mm wide, with two types of lamellulae, adnate to emarginate, ventricose, initially white, then brownish-rose, with entire and concolorous edge. Stipe 20–42 × 1.5–3.0 mm, central, terete, equal, hollow, gray (1C1–1E1), darker downwards, sparsely white fibrillous in the upper part elsewhere smooth and glabrous, base with white mycelium. Context thin, white. Odor indistinct, taste not tested.
E. subgriseosquamulosum A–D basidiomata A, B HFJAU3969, holotype C, D HFJAU3967 E basidiospores F cheilocystidia G basidia G, H pileipellis. G was observed in H2O, remaining microstructures all were observed in 5% KOH, and used 1% Congo red as the stain except E. Scale bars: 10 mm (A–D); 20 μm (E); 30 μm (F–H).
Basidiospores (8.1)8.4–10.5(11) × (6.0)6.5–8.0(8.5) μm, (av = 9.3 ×7.4 μm), Q = 1.1–1.4(1.5) (Qm = 1.3 ± 0.07, n = 100), subisodiameterical or heterodiametrical, 5–6 angles, mostly 5 angles in profile view, thick-walled, inamyloid. Basidia 27–36 × 10–13 μm, clavate, 4-spored, sterigmata 6.0–9.0 μm long, clampless. Pleurocystidia absent. Lamellae edge sterile of poliopus-type. Cheilocystidia 27–64 × 9.0–14 μm, clavate. Lamellar trama regular, made up of cylindrical hyphae 7.0–12 µm wide. Pileipellis a trichoderm made up of cylindrical hyphae 6.0–12 μm broad, with clavate terminal elements and yellow-brown intracellular pigment. Stipitipellis a cutis composed of densely arranged, cylindrical hyphae, 7.0–17 μm wide, slightly constricted at the septa, with acute or attenuated end. Clamp connections absent.
Solitary on soil or moss in broad-leaved forest.
So far known from Fujian Province in China.
China • Fujian Province, Wuyishan City, Yangzhuang Town, Xiyuan Village, 27.7652°N, 117.8164°E, alt. 512 m, 26 June 2022, collected by Jun-Qing Yan, Cheng-Feng Nie, and Lin-Gen Chen, HFJAU3967.
Morphologically, several similar species within Entoloma subg. Cyanula that share brown to brown-gray pileus can be distinguished from the new species as follows: E. anatinum (Lasch) Donk is characterized by its larger basidiospores (9.0–13.5 × 7.5–9.0 μm) with 6–9 angles, and fertile lamellae edge (
Phylogenetically, E. cyanostipitum Xiao L. He & W.H. Peng is closest to the new species. However, E. cyanostipitum is distinct by the deep blue pileus margin, lamellae edge and stipe, and the ITS region, with an 84% similarity (
Refers to its macroscopic morphology similar to “Entoloma praegracile”
China • Zhejiang Province, Lishui City, Qingyuan County, Bandaihoushang Village, 27.6748°N, 119.0780°E, alt. 1084 m, 7 July 2020, collected by Jun-Qing Yan and Yan-Liu Chen, HFJAU1822.
Entoloma subpraegracile is mainly characterized by the yellow, glabrous, and striate pileus, white, adnexed to adnate lamellae with tiny lateral veins, 5–7 angled and medium-sized basidiospores, sterile or heterogeneous lamellae edge of serrulatum-type, cylindrical or clavate cheilocystidia, and absence of clamp connections. It differs from E. praegracile by the larger basidiomata, and sterile or heterogeneous lamellae edge.
Basidiomata rather small. Pileus 10–20 mm wide, conical when young, then convex to flattened with depressed, rarely cuspidate center, with entire margin, not hygrophanous, smooth and glabrous, translucently striate almost up to the center, ochre (7B4–6), grayish yellow (1A4–5) to tawny (2C4–6), darker at center. Lamellae relatively dense, 1.5–2.0 mm wide, with tiny lateral veins and two or three types of lamellulae, adnate to adnexed, subventricose, white, with entire and concolorous edge. Stipe 25–35 × 2.0–2.5 mm, central, terete. equal, hollow, concolorous or paler with the pileus, smooth and glabrous, sometimes grooved, white tomentose at the base. Context thin, concolorous to the surface. Odor indistinct, taste not tested.
E. subpraegracile A, B basidiomata A HFJAU1822, holotype B HFJAU5115 C basidiospores D pileipellis E cheilocystidia F heterogeneous lamellae edge. All microscopic structures were observed in 5% KOH, and used 1% Congo red as the stain except C. Scale bars: 10 mm (A, B); 20 μm (C); 30 μm (D–F).
Basidiospores (7.0)8.5–10.5(12) × (6.0)6.5–7.5(8.5) μm, (av = 9.6 ×7.0 μm), Q = 1.2–1.6(1.7) (Qm = 1.4 ± 0.08, n = 200), heterodiametrical, 5–7(8) angles in profile view, appearing nodulose, thick-walled, inamyloid. Basidia 27–37 × 9–12 μm, clavate, slightly constricted at middle, mainly 2-spored, sterigmata 6.0–12 μm long, clampless. Pleurocystidia absent. Lamellae edge sterile or heterogeneous of poliopus-type. Cheilocystidia dense clusters on lamellae edge, 21–53 × 7.0–14 μm, cylindrical or clavate. Lamellar trama regular, made up of cylindrical hyphae 4.0–8.0 µm wide. Pileipellis a cutis made up of cylindrical hyphae 5.0–12 μm broad, with transitions to a trichoderm towards the center with clavate terminal elements 10–16 μm wide, with tawny intracellular pigment. Stipitipellis a cutis composed of densely arranged, cylindrical hyphae, 7.0–15 μm wide, slightly constricted at the septa, with rounded end. Clamp connections absent.
Solitary or scattered on soil in mixed coniferous-broad-leaved forest.
So far known from eastern China.
China • Fujian Province, Wuyishan City, 27.8594°N, 117.9096°E, alt. 372 m, 12 August 2021, collected by Jun-Qing Yan and Ze-Wei Liu, HFJAU3094 • 27.8563°N, 117.8661°E, alt. 668 m, 13 August 2021, collected by Qin Na, Yu-Peng Ge, and Lan-Yu Sun, HFJAU3164, HFJAU3168 • 27.7221°N, 117.7072°E, alt. 654 m, 16 August 2023, collected by Nian-Kai Zeng, Cheng-Feng Nie, Hua-Zhi Qin, Hui Deng, Tian Jiang, and Run-Xiang Zhao, HFJAU5110, HFJAU5115, HFJAU5140, HFJAU5175, HFJAU5177.
In the phylogenetic tree, E. subpraegracile groups together with E. praegracile. Entoloma praegracile differs from the new species by the smaller pileus (less than 10 mm), fertile lamellae edge, and the ITS sequence with 86% similarity (
Some similar species with a yellow pileus within subg. Cyanula can be distinguished from the new species as follows: E. chloropolium (Fr.) M.M. Moser is recognized by the fertile to heterogeneous lamellae edge, and septate cheilocystidia (
Refers to the collection locality of the holotype specimen – Wuyishan National Natural Park.
China • Fujian Province, Nanping City, Wuyishan National Natural Park, 27.5418°N, 117.4743°E, alt. 422 m, 7 June 2022, collected by Jun-Qing Yan and Lin-Gen Chen, HFJAU3571.
Entoloma wuyishanense is mainly characterized by the rather small and blue basidiomata, squamous and striate pileus, white and adnexed lamellae with fertile edge with slightly bluish pigmentation near the stipe, relatively large basidiospores with 5–6 angles, pileipellis with fuscous intracellular pigment. It differs from E. azureosquamulosum Xiao L. He & T.H. Li by the striate pileus, adnexed lamellae, larger basidiospores, and fertile lamellae edge.
Basidiomata rather small. Pileus 2.0–11 mm wide, conical when young, then convex to flattened with depressed center, with entire, straight or wavy margin, not hygrophanous, squamous with denser center, translucently striate almost up to the center, deep blue (20E4–7) to light gray-blue (20B2–3), darker at center. Lamellae moderately distant, 1.0–3.0 mm wide, with two types of lamellulae, adnexed, ventricose, white, with entire and bluish edge near the stipe. Stipe 9.0–26 × 1.0–2.0 mm, central, terete, equal, hollow, concolorous with pileus, paler downwards, white fibrillose, glabrescent with age, base with white mycelium. Context thin, gray-blue. Odor indistinct, taste not tested.
Basidiospores (9.5)10–13.5(15) × (6.5)7.5–9.5(10) μm, (av = 11.7 ×8.5 μm), Q = 1.2–1.7(2.0) (Qm = 1.4 ± 0.11, n = 200), heterodiametrical, 5–6 angles in profile view, sometimes appearing nodulose, thick-walled, inamyloid. Basidia 25–33 × 10–13 μm, clavate, slightly constricted at middle, 4- or 2-spored, sterigmata 6.0–10 μm long, clampless. Lamellae edge fertile. Cystidia absent. Lamellar trama regular, made up of cylindrical hyphae 5.0–11 µm wide. Pileipellis a trichoderm made up of clavate to pyriform terminal cells, 41–65 × 22–36 μm. Pigment fuscous, intracellular, diffuse in pileipellis. Stipitipellis a cutis composed of densely arranged, cylindrical hyphae, 5.0–12 μm wide, with rounded end. Clamp connections absent.
Solitary or scattered on moss in mixed coniferous-broad-leaved forest.
So far known from eastern China.
China • Zhejiang Province, Lishui City, Songyang County, Zicao Village, 28.4874°N, 119.5783°E, alt. 722 m, 2 July 2022, collected by Jun-Qing Yan, Cheng-Feng Nie, and Meng-Hui Han, HFJAU3871, HFJAU3874, HFJAU3878 • Lishui City, Yunhe County, Chongtou Town, Xiayang Village, 28.0499°N, 119.4732°E, alt. 592 m, 4 July 2022, collected by Jun-Qing Yan and Cheng-Feng Nie, HFJAU3881.
Morphologically, E. azureosquamulosum is the most similar species with the distinction that E. azureosquamulosum exhibits not striate pileus, adnate lamellae, smaller basidiospores (8–10.5 × 6.5–8.0 μm), and sterile lamellae edge (
In the phylogenetic tree, E. wuyishanense belongs to Cyanula sect. Poliopodes, within which several species have blue pileus, including E. argus O.V. Morozova, E.S. Popov, A.V. Alexandrova & Noordel., E. calceus Noordel., Bendiksen, Brandrud, P.-A. Moreau & Vila, E. corvinum (Kühner) Noordel., E. icarus O.V. Morozova, E.S. Popov & Noordel., and E. perchalybeum Noordel., J.B. Jordal & Dima. However, the lamellae edge of the latter in all species is sterile. In addition, E. argus is characterized by the adnate lamellae and smaller basidiospores (≤10 μm) (
Entoloma subg. Cyanula currently is divided into 11 sections (
Notably, based on the results of this phylogenetic analysis, we have realized that the sect. Caesiocincta is divided into three clades. However, since none of the originating branches of these scattered clades is supported, the cause of this result cannot be determined. Additional specimen data are needed for further analysis.
The present study expands our understanding of entolomoid species by providing descriptions and phylogenetic analyses for four new species. The findings enrich our knowledge of the distribution of E. subg. Cyanula species in China and the overall diversity of Entoloma.
1 | Pileus white to pink | 2 |
– | Pileus other colored | 3 |
2 | Pileus white, not striate, with depressed center; lamellae adnate to decurrent; basidiospores 5–6 angled; pigment not | E. orientosinense |
– | Pileus pink, striate, with umbonate center; lamellae subfree to adnexed; basidiospores 6–8 angled; pigment yellow encrusting | E. mastoideum |
3 | Pileus yellow-brown to grayish-brown | 4 |
– | Pileus blue to violaceous | 13 |
4 | Pileus glabrous to fibrillose | 5 |
– | Pileus squamulose to velvety | 8 |
5 | Basidiospores Lav ≥ 11 μm; pileus striate, with depressed center; lamellae adnate; lamellae edge sterile or heterogeneous | E. subtenuicystidiatum |
– | Basidiospores Lav < 11 μm | 6 |
6 | Pileus ≥ 20 mm, with margin exceeding lamellae; lamellae adnate-emarginate to adnexed | E. caespitosum |
– | Pileus < 20 mm | 7 |
7 | Lamellae edge sterile or heterogeneous; pileus not hygrophanous | E. subpraegracile |
– | Lamellae edge fertile; pileus hygrophanous | E. praegracile |
8 | Lamellae edge fertile; pileus striate; lamellae adnate or emarginate; basidiospores 8.0–14 × 5.5–10 μm | E. insidiosum |
– | Lamellae edge sterile | 9 |
9 | Basidiospores Lav ≥ 10 μm; pileus striate; lamellae adnexed; pigment yellow-brown intracellular | E. longistriatum |
– | Basidiospores Lav < 10 μm | 10 |
10 | Cheilocystidia cylindrical to clavate | 11 |
– | Cheilocystidia fusiform, lageniform, vesiculose to spheropedunculate | 12 |
11 | Pileus striate; lamellae adnate to emarginate; lamellae edge entire and concolorous with lamellae | E. subgriseosquamulosum |
– | Pileus not striate; lamellae adnexed to short decurrent; lamellae edge serrulate and blue-black | E. subcaesiocinctum |
12 | Cheilocystidia fusiform to lageniform; pileus not striate; lamellae adnexed to free; lamellae edge concolorous with lamellae | E. pseudosubcorvinum |
– | Cheilocystidia vesiculose or spheropedunculate; pileus not striate; lamellae adnate-emarginate; lamellae edge brown | E. pulchripes |
13 | Lamellae edge fertile; pileus squamous, striate; lamellae adnexed; lamellae edge blue; basidiospores 10–13.5 × 7.5–9.5 μm | E. wuyishanense |
– | Lamellae edge sterile | 14 |
14 | Pileus not striate | 15 |
– | Pileus striate | 16 |
15 | Lamellae adnate-emarginate; cheilocystidia fusoid | E. azureosquamulosum |
– | Lamellae short decurrent; cheilocystidia cylindrical to subclavate | E. cyanostipitum |
16 | Cheilocystidia subglobose or sphaeropedunculate; lamellae edge concolorous with lamellae | E. ekaterinae |
– | Cheilocystidia broadly clavate or lageniform; lamellae edge blackish purple | E. callipygmaeum |
The authors are very grateful for assistance of Yu-Peng Ge and Meng-Hui Han in the field specimen collection and the anonymous reviewers of the manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This work was financed by the National Natural Science Foundation of China (32460326, 31960008), Jiangxi Provincial Natural Science Foundation (20224BAB205003), Fujian Provincial Natural Science Foundation (2023J01379), and the Project of FAAS (XTCXGC2021007).
Conceptualization, Jun-Qing Yan; methodology, Jun-Qing Yan and Sheng-Nan Wang; software, Lin-Gen Chen, Hong Chen, Ling Ding and Yu-Qin Xu; formal analysis, Hui Zeng, Jun-Qing Yan, and Sheng-Nan Wang; investigation, Lin-Gen Chen, Hong Chen, Ling Ding, and Jun-Qing Yan; resources, Hui Zeng and Jun-Qing Yan; writing – original draft, Lin-Gen Chen; writing – review and editing, Jun-Qing Yan; visualization, Jun-Qing Yan and Sheng-Nan Wang; supervision, Jun-Qing Yan; project administration, Jun-Qing Yan; funding acquisition, Jun-Qing Yan. All authors have read and agreed to the published version of the manuscript.
Lin-Gen Chen https://orcid.org/0009-0000-2506-181X
Hong Chen https://orcid.org/0009-0004-0107-3962
Ling Ding https://orcid.org/0009-0000-5293-046X
Hui Zeng https://orcid.org/0000-0003-2025-844X
Sheng-Nan Wang https://orcid.org/0000-0003-0648-271X
Jun-Qing Yan https://orcid.org/0000-0003-1128-5171
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Supplementary data
Data type: nex
Explanation note: A nexus file contains alignment sequence and original tree of ML and Bayes.