Corresponding authors: Tai-Hui Li (
Academic editor: R. H. Nilsson
Based on phylogenetic analyses, some newly studied Chinese mushroom specimens were found to represent two distinct species within the genus
Li J-P, Antonín V, Gates G, Jiang L, Li T-H, Li Y, Song B, Deng C-Y (2022) Emending Gymnopus sect. Gymnopus (Agaricales, Omphalotaceae) by including two new species from southern China. MycoKeys 87: 183–204.
Morphologically, the current circumscription of
Phylogenetically,
In this study, two new species of
For Latin names:
For phylogenetic analysis:
For collection locality:
For climate:
For soil type: B = brown;
Nine collections from China were examined in this study: one came from the Guizhou Province (Tongren City), three collections from the Yunnan Province (one from Pu’er City and two from Maguan County) and five collections from the Guangdong Province (one from Guangzhou City, one from Shenzhen City and three from Xinyi City). The exact localities and their environmental characteristics are shown in Table
The environmental characteristics of localities for each collection.
Locality | Climate | Average annual temperature | Average annual rainfall | Major soil type | References |
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16.9 °C | 1351 mm |
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16.9 °C | 1345 mm |
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17.8 °C | 1514.6 mm | ||
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22 °C | 1725 mm |
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22.4 °C | 1948.4 mm | ||
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18 °C | 2300–2600 mm |
Fresh basidiomata were photographed and used for macromorphological descriptions. The colours are coded from
Micromorphological structures were observed via a ZEISS Axio Lab. A1 microscope based on the hand-made sections of dried basidiomata mounted in 5 % KOH on a glass slide. When necessary, Congo Red solution was used as a stain and Melzer’s reagent was used to test amyloid or dextrinoid reactions. For the various microscopic structures, ‘n’ refers to the number of measured elements. For basidiospores, ‘E’ represents the quotient of length and width in any one spore, and ‘Q’ represents the mean of E values. Basidiospore measurements do not include apiculus and are presented as ‘(a)b–c(d)’, where ‘b–c’ represents the minimum of 90 % of the measured values and ‘a’ and ‘d’ represent the extreme values. The main body (sterigmata or excrescences not included) of basidia, basidioles, pleurocystidia and cheilocystidia were measured (if present).
Genomic DNA was extracted from dried tissue via a Magen HiPure Fungal DNA Kit (Magen Biotech
Information on DNA sequences used in the phylogenetic analyses. Newly generated sequences are highlighted in bold and type specimen is marked with an asterisk (*).
Taxon name | ITS | LSU | Collection No. | Locality | Reference |
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Sw2-1 | Japan | GenBank | |
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SFSU:DED8813 | Not given |
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China |
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CULTENN5609 | USA |
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CBS 240.53 | France |
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CBS 239.53 | France |
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CULTENN5021h2 | Canada |
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TENN:F-59594 | Russia |
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URM 87728 | Brazil |
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SFSU:AWW118 | Indonesia |
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XAL: Cesar50 | Mexico |
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BRNM 714974 | South Korea |
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BRNM 747547 | South Korea |
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BRNM 714927 | South Korea |
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BRNM 781447 | South Korea |
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TENN:F-57012 | Not given |
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TENN:F-61125 | USA | Hughes and Petersen (2016) |
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TENN:F-69323 | USA | Hughes and Petersen (2016) |
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TENN:F-55679 | USA | Hughes and Petersen (2016) | |
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TENN:F-59300 | Austria |
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TENN:F-69254 | Slovakia | Hughes and Petersen (2016) |
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TENN:F-59217 | France |
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BRNM 714849 | Czech Republic |
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TENN:F-48143 | USA | Hughes and Petersen (2016) | |
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TFB 4529 | USA | Hughes and Petersen (2016) | |
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TENN:F-53490 | USA | Hughes and Petersen (2016) | |
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TENN:F-51233 | USA | Hughes and Petersen (2016) | |
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BCN:SCM B-4058 | Spain |
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TENN:F-52970 | USA |
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Unavailable | SFSU:DED 8209 | São Tomé |
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JMCR 143 | Not given |
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TENN:F-14505 | USA |
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CULTENN4975 | USA | Hughes and Petersen (2016) | |
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SFSU-DED5097 | USA | Hughes and Petersen (2016) | |
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TENN:F-65135 | Belgium | Hughes and Petersen (2016) |
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* |
China | This study |
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China | This study | |
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China | This study | |
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China | This study | |
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Unavailable | China | This study | ||
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China | ||
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TENN:F-50999 | Puerto Rico | Hughes and Petersen (2016) |
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China | This study | |
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China | This study | |
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* |
China | This study |
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China | This study | |
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BRNM 714981 | South Korea |
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TENN:F-65912 | USA | Hughes and Petersen (2016) |
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PDD:96595 | New Zealand | GenBank |
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URM 87730 | Brazil |
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Sara Landvik:NN008037 | Sweden |
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MUCL35155 | Not given | Klonowska et al. (2013) |
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PDD:106823 | New Zealand | GenBank |
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PDD:113265 | New Zealand | GenBank |
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Unavailable | NEHU.MBSRJ.48 | India |
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TENN:F-55630 | Russia |
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TENN:F-53474 | USA |
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TENN:F-50319 | Sweden |
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Representative species and their sequences were selected to cover all sections of
A BLAST search of
The combined dataset comprised 113 sequences including 58
Phylogram generated by
In the generated phylogenetic tree (Fig.
China, Guangdong Province, Shenzhen City, Wutongshan Scenic Area, 16 September 2019, H. Huang, L.Q. Wu & N. Zhan (
The epithet ‘
Differs from
Basidiomata omphalinoid, collybioid or gymnopoid. Pileus 10–40 mm broad, membranous, hemispheric when young, becoming convex, plano-convex to applanate, generally umbilicate to sometimes slightly depressed at the centre, inflexed then straight or reflexed at margin, with a marginal zone often undulating with age, glabrous, radially striate or grooved towards the margin, orange (6B7) or reddish orange (7B7) to brown (7D8) overall when young, somewhat reddish orange (7B7) or dark brown (7F8), then paler towards the margin, white or pale orange (6A3) to light brown (6D4), often greyish orange (6B4) to dark brown (6F8) at the disc. Lamellae adnate, broad, ventricose to broadly ventricose, white when fresh, sometimes with greyish red (7B4) to brown (7E7) tint somewhere, margin entire to split and sometimes grooved, L = 12–17, l = 3–5. Stipe 10–30 mm long, 2–4 mm thick in the middle, central, cylindrical, or compressed, with dense basal mycelium when young that disappears when old, hollow, fibrous, glabrous, slightly longitudinally striate when old, rooting deep in the substrate, but eventually attaches to the stump, dull white to greyish red (7B4) when young, soon darker towards the base, white to reddish orange (7A7) at apex, finally entirely dark brown (7F8). Odour not distinctive.
Basidiomata of
Microscopic features of
Basidiospores [n=80] (3.5–) 4.0–5.5 (–6.0) × 2.5–3 (–3.5) μm (average= 4.63 × 2.93 μm, E = 1.33–1.83 (–2), Q=1.58), obovoid, ellipsoid to subellipsoid, sometimes amygdaliform. Basidia [n=20] 17–31 × 3–5 μm, clavate, 4-spored. Basidioles [n=20] 17–32 × 4–5.5 μm, clavate, cylindrical. Lamellar edge sterile. Cheilocystidia [n=20] 17–32 × 4–10 μm, irregularly clavate, sphaeropedunculate or almost so, with tendency to be inflated, with or without finger-like apical projection(s) or more or less diverticulate elements. Pileipellis a cutis composed of cylindrical, thin-walled hyphae, up to 12.5 μm wide, smooth or with scattered diverticula, hyaline to slightly brownish;
Saprotrophic, gregarious or in small clusters, usually rooting around the roots and stumps in broadleaf forests.
China, Guangdong Province, Guangzhou City, Tianluhu Forest Park, longitude and latitude not recorded, alt. not recorded, 4 April 2019, T.H. Li, W.Q. Deng, J.Y. Xu & J.P. Li (
Among the known species of
China, Guangdong Province, Xinyi City, Yunkaishan National Nature Reserve,
The epithet “
Differs from
Basidiomata gymnopoid or collybioid. Pileus 10–20 mm broad, membranous, hemispherical when young, then convex, with slightly inflexed margin, expanding to plano -convex , with a depressed disc, undulating at the margin, glabrous, radially striate or grooved towards the margin, often pale orange (6A3) to light brown (6D8), darker at the centre, sometimes to dark brown (6F8), white to light brown (6D8) towards the margin. Lamellae adnate, linear to arcuate, sometimes furcate to branched or venose, generally split at the edge, dull white to brownish orange (7C7), pale at the edge, sometimes with brown (7E8) to dark brown (7F8) tints somewhere, L = 10–20, l = 3–4. Stipe 11–21 mm long, 0.8–1 mm thick in middle, central, cylindrical, straight or sometimes curved, insititious, hollow, fibrous, glabrous, rooting deep in the substrate, but eventually attaches to the stump, white to orange-white (6A2) at first, slightly darker at base, then darker towards the apex, finally entirely light brown (7D8) to brown (7E8). Odour not distinctive.
Basidiospores [n=80] 4–6 (–6.5) × 2.5–3 (–3.5) μm (average = 4.90 × 2.93 μm, E = (1.29–) 1.33–2.00 (–2.20), Q = 1.68) or [n=20] 6.5–8 × 2.5–3 μm (average = 7.35 × 2.86 μm, E = 2.17–3.2, Q = 2.65), obovoid, ellipsoid to subellipsoid, sometimes amygdaliform. Basidia [n=20] 15–32 × 4–6 μm, clavate, 4-spored, rarely 1–3-spored. Basidioles [n=20] 17–27.5 × 4–6.5 μm, clavate, cylindrical. Lamellar edge sterile. Cheilocystidia [n=20] 20–43 × 4.5–9 μm, irregularly clavate, tending to inflated, with finger-like apical projection(s) or more or less diverticulate elements. Pileipellis a cutis composed of thin-walled, cylindrical hyphae up to 18 μm wide, smooth or with scattered diverticula, hyaline to slightly greyish;
Saprotrophic, gregarious or in small clusters, usually rooting around roots and stumps in broadleaf forests.
China, Guangdong Province, Xinyi City, Yunkaishan National Nature Reserve,
Morphologically, among the known species of
Basidiomata of
Microscopic features of
According to the phylogenetic results, the two new species could be taken to represent a new section within
Morphologically, the taxonomic placement of
As the characteristic of the pileipellis is a significant factor for sectional delimitation in
A very interesting and unusual characteristic is a splitting lamellar edge in both newly described species. What advantage such split lamellar edge could confer is difficult to surmise, but
Pileus membranous or fleshy; stipe smooth or slightly to deeply sulcate-striate, with a well-developed or reduced pseudorrhiza; spore print white to pale ochraceous; cheilocystidia versiform, clavate, fusoid, tending inflated, sometimes with more or less finger-like apical projection(s), or diverticulate elements; pileipellis a cutis, or this transitioning to a trichoderm, with broad terminal elements, mostly inflated, mixed with irregularly branched elements and some resembling
1 | Pileus fleshy; stipe with a distinct pseudorrhiza |
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– | Pileus membranous; stipe without a pseudorrhiza but rooting in the substrate |
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2 | Pileus generally deeply umbilicate; lamellae broad, adnate and ventricose |
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– | Pileus more or less depressed; lamellae adnate, linear to arcuate |
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Grateful thanks are due to Prof. Xiang-Hua Wang (Kunming Institute of Botany, CAS, Kunming, China) for providing specimen(s), sequences, suggestions and photographs, Dr Jerry Adrian Cooper (Landcare Research, New Zealand) for providing sequences and photographs, Dr Rolf Henrik Nilsson (University of Gothenburg, Gothenburg, Sweden) for improving our work, Dr Ji-Ze Xu (Jilin Agricultural Science and Technology University, Jilin, China) for providing specimens and sequence data, Dr Md Iqbal Hosen, Prof. Wang-Qiu Deng, Dr Chao-Qun Wang (Guangdong Institute of Microbiology, Guangzhou, China) and Xiao-Ya An (Shenyang Agricultural University, Shenyang, China) for providing suggestions, Dr Ming Zhang, Mr. Ting Li, Mr. Juan-Yan Xu, Mr. Hao Huang, Mr. Li-Qiang Wu, Mr. Ning Zhan, Mr. Hua-Shu Wen (Guangdong Institute of Microbiology, Guangzhou, China), Prof. Shu-Hong Li (Yunnan Academy of Agricultural Sciences, Kunming, China) for hunting collection(s). This work was supported by the National Natural Science Foundation of China (Nos. 31750001, 31970016), the Science and Technology Planning Project of Guangdong Province, China (2019B121202005, 2018B030320001, 20070617627078), the government procurement project of Shenzhen, China (SZCG2019191412), China Agriculture Research System (CARS-20), the government procurement project of China (ZX2021-FJC083), Projects of Science and Technology Programs of Guizhou Province ([2019]2451, [2019]4007-2), GDAS’ Special Project of Science and Technology Development (Grant No. 2019GDASYL-0104011), and the Project of Comprehensive Scientific Investigation of Dalingshan Forest Park in Dongguan (441901-2021-08594). The studies of V.A. were made possible by the support provided to the Moravian Museum by the Ministry of Culture of the Czech Republic as part of its long-term conceptual development program for research institutions (DKRVO, ref. MK000094862).