Research Article |
Corresponding author: Dong-Mei Wu ( wdm0999123@sina.com ) Corresponding author: Bao-Kai Cui ( cuibaokai@bjfu.edu.cn ) Academic editor: María P. Martín
© 2022 Shun Liu, Tai-Min Xu, Chang-Ge Song, Chang-Lin Zhao, Dong-Mei Wu, Bao-Kai Cui.
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:
Liu S, Xu T-M, Song C-G, Zhao C-L, Wu D-M, Cui B-K (2022) Species diversity, molecular phylogeny and ecological habits of Cyanosporus (Polyporales, Basidiomycota) with an emphasis on Chinese collections. MycoKeys 86: 19-46. https://doi.org/10.3897/mycokeys.86.78305
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Cyanosporus is a genus widely distributed in Asia, Europe, North America, South America and Oceania. It grows on different angiosperm and gymnosperm trees and can cause brown rot of wood. Blue-tinted basidiomata of Cyanosporus makes it easy to distinguish from other genera, but the similar morphological characters make it difficult to identify species within the genus. Phylogeny and taxonomy of Cyanosporus were carried out based on worldwide samples with an emphasis on Chinese collections, and the species diversity of the genus is updated. Four new species, C. flavus, C. rigidus, C. subungulatus and C. tenuicontextus, are described based on the evidence of morphological characters, distribution areas, host trees and molecular phylogenetic analyses inferred from the internal transcribed spacer (ITS) regions, the large subunit of nuclear ribosomal RNA gene (nLSU), the small subunit of nuclear ribosomal RNA gene (nSSU), the small subunit of mitochondrial rRNA gene (mtSSU), the largest subunit of RNA polymerase II (RPB1), the second largest subunit of RNA polymerase II (RPB2), and the translation elongation factor 1-α gene (TEF). Our study expanded the number of Cyanosporus species to 35 around the world including 23 species from China. Detailed descriptions of the four new species and the geographical locations of the Cyanosporus species in China are provided.
brown-rot fungi, distribution areas, host trees, multi-gene phylogeny, new species
Cyanosporus was proposed as a monotypic genus for Polyporus caesius (Schrad.) Fr. based on its cyanophilous basidiospores (
Previously, species identification of the P. caesia complex was only based on morphological characters and host trees in China, and only two species were recorded from China before
Cyanosporus species usually have blue-tinted basidiomata, which makes it easy to recognize. Some specimens with blue-tinted basidiomata were collected during investigations into the diversity of polypores in China, and four undescribed species of Cyanosporus were discovered. To confirm the affinity of the undescribed species to Cyanosporus, phylogenetic analyses were carried out based on the combined datasets of ITS+TEF and ITS+nLSU+nSSU+mtSSU+RPB1+RPB2+TEF sequences. During the investigation and study of Cyanosporus, the information of host trees and distribution areas of species in the genus from China were also obtained (Table
The main ecological habits of Cyanosporus with an emphasis on distribution areas and host trees. New species are shown in bold.
Species | Distribution in the world | Distribution in China | Climate zone | Host | Reference |
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C. alni (Niemelä & Vampola) B.K. Cui, L.L. Shen & Y.C. Dai | Europe (Czech Republic, Denmark, Finland, Germany, Norway, Poland, Russia, Slovakia), East Asia (China) | Guizhou, Hebei | Temperate | Angiosperm (Alnus, Betula, Corylus, Fagus, Populus, Quercus) |
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C. arbuti (Spirin) B.K. Cui & Shun Liu | North America (USA) | Temperate | Angiosperm (Arbutus) |
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C. auricomus (Spirin & Niemelä) B.K. Cui & Shun Liu | Europe (Finland, Poland, Russia), East Asia (China) | Inner Mongolia | Temperate to boreal | Gymnosperm (Pinus, Picea) |
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C. bifarius (Spirin) B.K. Cui & Shun Liu | Europe (Russia), East Asia (China, Japan) | Jilin, Sichuan, Yunnan | Cold temperate | Gymnosperm (Picea, Pinus, Larix) |
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C. bubalinus B.K. Cui & Shun Liu | East Asia (China) | Yunnan | Temperate | Gymnosperm (Pinus) |
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C. caesiosimulans (G.F. Atk.) B.K. Cui & Shun Liu | Europe (Finland, Russia), North America (USA) | Temperate | Angiosperm (Corylus, Fagus, Populus) and gymnosperm (Abies, Picea) |
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C. caesius (Schrad.) McGinty | Europe (Czech Republic, Denmark, Finland, France, Germany, Russia, Slovakia, Spain, UK) | Common in temperate, rare in south boreal zone | Angiosperm (Betula, Fagus, Salix) and gymnosperm (Abies, Picea) |
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C. coeruleivirens (Corner) B.K. Cui, Shun Liu & Y.C. Dai | Asia (China, Indonesia), Europe (Russia) | Hunan, Jilin, Zhejiang | Warm temperate | Angiosperm (Tilia, Ulmus) |
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C. comatus (Miettinen) B.K. Cui & Shun Liu | North America (USA), East Asia (China) | Sichuan, Xizang | Temperate | Angiosperm (Acer) and gymnosperm (Abies, Picea, Tsuga) |
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C. cyanescens (Miettinen) B.K. Cui & Shun Liu | Europe (Estonia, Finland, France, Poland, Russia, Spain, Sweden) | Temperate to Mediterranean mountains | Gymnosperm (Abies, Picea, Pinus) |
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C. flavus B.K. Cui & Shun Liu | East Asia (China) | Sichuan | Plateau humid climate | Gymnosperm (Abies, Picea) | Present study |
C. fusiformis B.K. Cui, L.L. Shen & Y.C. Dai | East Asia (China) | Guizhou, Sichuan | North temperate to subtropical | Angiosperm (Rhododendron) |
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C. glauca (Spirin & Miettinen) B.K. Cui & Shun Liu | East Asia (China), Europe (Russia) | Jilin | Cold temperate mountains | Gymnosperm (Abies, Picea) |
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C. gossypinus (Moug. & Lév.) B.K. Cui & Shun Liu | Europe (France) | Temperate | Gymnosperm (Cedrus) |
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C. hirsutus B.K. Cui & Shun Liu | East Asia (China) | Qinghai, Sichuan, Yunnan | Temperate to plateau continental climate | Gymnosperm (Abies, Picea) |
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C. livens (Miettinen & Vlasák) B.K. Cui & Shun Liu | North America (Canada, USA) | Temperate | Angiosperm (Acer, Betula, Fagus) and gymnosperm (Abies, Larix, Picea, Tsuga) |
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C. luteocaesius (A. David) B.K. Cui, L.L. Shen & Y.C. Dai | Europe (France) | Mediterranean | Gymnosperm (Pinus) |
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C. magnus (Miettinen) B.K. Cui & Shun Liu | East Asia (China) | Chongqin, Jilin, Hainan, Yunnan | Temperate | Angiosperm (Populus) and gymnosperm (Cunninghamia) |
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C. mediterraneocaesius (M. Pieri & B. Rivoire) B.K. Cui, L.L. Shen & Y.C. Dai | Europe (France, Spain) | Warm temperate to Mediterranean | Angiosperm (Buxus, Erica, Populus, Quercus) and gymnosperm (Cedrus, Juniperus, Pinus) |
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C. microporus B.K. Cui, L.L. Shen & Y.C. Dai | East Asia (China) | Yunnan | subtropical | Angiosperm (undetermined) |
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C. nothofagicola B.K. Cui, Shun Liu & Y.C. Dai | Oceania (Australia), South America (Argentina) | Temperate marine climate | Angiosperm (Nothofagus) |
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C. piceicola B.K. Cui, L.L. Shen & Y.C. Dai | East Asia (China) | Sichuan, Xizang, Yunnan | Warm temperate to subtropical | Gymnosperm (Picea) |
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C. populi (Miettinen) B.K. Cui & Shun Liu | East Asia (China), Europe (Finland, Norway, Poland, Russia), North America (USA) | Qinghai, Jilin, Sichuan, Yunnan | Boreal to temperate | Angiosperm (Acer, Alnus, Betula, Populus, Salix) and gymnosperm (Picea) |
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C. rigidus B.K. Cui & Shun | East Asia (China) | Yunnan | Warm temperate | Gymnosperm (Picea) | Present study |
C. simulans (P. Karst.) B.K. Cui & Shun Liu | East Asia (China), Europe (Estonia, Finland, France, Germany, Norway, Russia), North America (Canada, USA) | Jilin | Warm temperate to boreal | Angiosperm (Corylus, Fagus, Populus, Sorbus, Ulmus) and gymnosperm (Abies, Cedrus, Juniperus, Picea, Pinus, Thuja, Tsuga) |
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C. subcaesius (A. David) B.K. Cui, L.L. Shen & Y.C. Dai | Europe (Czech Republic, Finland, France, Russia, UK) | Temperate | Angiosperm (Alnus, Carpinus, Crataegus, Corylus, Fagus, Fraxinus, Malus, Populus, Prunus, Quercus, Salix, Ulmus) |
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C. subhirsutus B.K. Cui, L.L. Shen & Y.C. Dai | East Asia (China) | Guizhou, Fujian, Yunnan | Warm temperate to subtropical | Angiosperm (Pterocarya) |
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C. submicroporus B.K. Cui & Shun Liu | East Asia (China) | Sichuan, Yunnan, Zhejiang | Alpine plateau to subtropical | Angiosperm (Alnus, Cyclobalanopsis) |
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C. subungulatus B.K. Cui & Shun Liu | East Asia (China) | Yunnan | Subtropical | Angiosperm (undetermined) and gymnosperm (Pinus) | Present study |
C. subviridis (Ryvarden & Guzmán) B.K. Cui & Shun Liu | Europe (Finland), North America (Mexico, USA) | Temperate to boreal | Gymnosperm (Abies, Picea, Pinus) |
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C. tenuicontextus B.K. Cui & Shun Liu | East Asia (China) | Yunnan | Subtropical | Angiosperm (undetermined) and gymnosperm (Pinus) | Present study |
C. tenuis B.K. Cui, Shun Liu & Y.C. Dai | East Asia (China) | Sichuan | Subtropical monsoon to Alpine plateau | Gymnosperm (Picea) |
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C. tricolor B.K. Cui, L.L. Shen & Y.C. Dai | East Asia (China) | Sichuan, Xizang | Alpine plateau | Gymnosperm (Abies, Picea) |
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C. ungulatus B.K. Cui, L.L. Shen & Y.C. Dai | East Asia (China) | Sichuan | Subtropical monsoon to Alpine plateau | Angiosperm (Castanopsis) and gymnosperm (Abies) |
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C. yanae (Miettinen & Kotir.) B.K. Cui & Shun Liu | Europe (Russia) | Temperate continental climate | Gymnosperm (Larix, Pinus) |
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The examined specimens were deposited in the herbarium of the Institute of microbiology, Beijing Forestry University (
In the text the following abbreviations were used: IKI = Melzer’s reagent, IKI– = neither amyloid nor dextrinoid, KOH = 5% potassium hydroxide, CB = Cotton Blue, CB + = cyanophilous, CB – = acyanophilous, L = mean spore length (arithmetic average of all spores), W = mean spore width (arithmetic average of all spores), Q = variation in the L/W ratios between the specimens studied, n (a/b) = number of spores (a) measured from given number (b) of specimens.
A cetyl trimethylammonium bromide (CTAB) rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd, Beijing, China) was used to extract total genomic DNA from dried specimens, and performed the polymerase chain reaction (PCR) according to the manufacturer’s instructions with some modifications as described by
The PCR cycling schedule for ITS, mtSSU and TEF included an initial denaturation at 95 °C for 3 min, followed by 35 cycles at 94 °C for 40 s, 54 °C for ITS and mtSSU, 54–55 °C for TEF for 45 s, 72 °C for 1 min, and a final extension at 72 °C for 10 min. The PCR cycling schedule for nLSU and nSSU included an initial denaturation at 94 °C for 1 min, followed by 35 cycles at 94 °C for 30 s, 50 °C for nLSU and 52 °C for nSSU for 1 min, 72 °C for 1.5 min, and a final extension at 72 °C for 10 min. The PCR procedure for RPB1 and RPB2 follow
Additional sequences were downloaded from GenBank (Table
Most parsimonious phylogenies were inferred from the combined 2-gene dataset (ITS+TEF) and 7-gene dataset (ITS+nLSU+nSSU+mtSSU+RPB1+RPB2+TEF), and their congruences were evaluated with the incongruence length difference (ILD) test (
MrModeltest 2.3 (
The combined 2-gene (ITS+TEF) sequences dataset had an aligned length of 1015 characters, of which 502 characters were constant, 62 were variable and parsimony-uninformative, and 451 were parsimony-informative. MP analysis yielded 10 equally parsimonious trees (TL = 2396, CI = 0.379, RI = 0.735, RC = 0.279, HI = 0.621). The best model for the concatenate sequence dataset estimated and applied in the Bayesian inference was GTR+I+G with equal frequency of nucleotides. ML analysis resulted in a similar topology as MP and Bayesian analyses, and only the ML topology is shown in Fig.
Maximum likelihood tree illustrating the phylogeny of Cyanosporus and its related genera in the antrodia clade based on the combined sequences dataset of ITS+TEF. Branches are labelled with maximum likelihood bootstrap higher than 50%, parsimony bootstrap proportions higher than 50% and Bayesian posterior probabilities more than 0.90 respectively. Bold names = New species.
The combined 7-gene (ITS+nLSU+nSSU+mtSSU+RPB1+RPB2+TEF) sequences dataset had an aligned length of 5634 characters, of which 3843 characters were constant, 247 were variable and parsimony-uninformative, and 1544 were parsimony-informative. MP analysis yielded 23 equally parsimonious trees (TL = 5756, CI = 0.468, RI = 0.752, RC = 0.352, HI = 0.532). The best model for the concatenate sequence dataset estimated and applied in the Bayesian inference was GTR+I+G with equal frequency of nucleotides. ML analysis resulted in a similar topology as MP and Bayesian analyses, and only the ML topology is shown in Fig.
Maximum likelihood tree illustrating the phylogeny of Cyanosporus and its related genera in the antrodia clade based on the combined sequences dataset of ITS+nLSU+nSSU+mtSSU+RPB1+RPB2+TEF. Branches are labelled with maximum likelihood bootstrap higher than 50%, parsimony bootstrap propwortions higher than 50% and Bayesian posterior probabilities more than 0.90 respectively. Bold names = New species.
The phylogenetic trees inferred from ITS+TEF and ITS+nLSU+nSSU+mtSSU+RPB1+RPB2+TEF gene sequences were all obtained from 106 fungal samples representing 65 taxa of Cyanosporus and its related genera within the antrodia clade. 74 samples representing 35 taxa of Cyanosporus clustered together and separated from species of Postia and other related genera. As for Cyanosporus, the sequences used in phylogenetic analyses include 28 holotype specimen sequences, one isotype specimen sequence and one neotype specimen sequence (Table
Cyanosporus flavus is characterised by flabelliform to semicircular and hirsute pileus with ash grey to light vinaceous grey pileal surface when fresh, buff to lemon-chrome pore surface when dry, and allantoid and slightly curved basidiospores (4.6–5.2 × 0.8–1.3 μm).
China. Sichuan Province, Jiuzhaigou County, on stump of Picea sp., 19.IX.2020, Cui 18547 (
Flavus (Lat.): referring to its lemon-chrome pore surface when dry.
Basidiomata annual, pileate, soft and watery, without odour or taste when fresh, becoming corky to fragile and light in weight upon drying. Pileus flabelliform to semicircular, projecting up to 3.2 cm, 5.7 cm wide and 0.9 cm thick at base. Pileal surface ash-grey to light vinaceous grey when fresh, becoming pale mouse-grey to mouse-grey when dry, hirsute; margin acute to slightly obtuse, white with a little blue tint when fresh, olivaceous buff to greyish brown when dry. Pore surface white to cream when fresh, becoming buff to lemon-chrome when dry; sterile margin narrow to almost lacking; pores angular, 5–7 per mm; dissepiments thin, entire to lacerate. Context white to cream, soft corky, up to 6 mm thick. Tubes pale mouse-grey to ash-grey, fragile, up to 4 mm long.
Hyphal system monomitic; generative hyphae with clamp connections, IKI–, CB–; hyphae unchanged in KOH.
Generative hyphae hyaline, thin- to slightly thick-walled with a wide lumen, occasionally branched, loosely interwoven, 2.7–6.5 μm in diam.
Generative hyphae hyaline, thin- to slightly thick-walled with a wide lumen, rarely branched, interwoven, 2.2–4.7 μm in diam. Cystidia absent; cystidioles present, fusoid, thin-walled, 12.3–17.8 × 2.2–3.5 μm. Basidia clavate, bearing four sterigmata and a basal clamp connection, 13.2–16.5 × 3.2–5.5 μm; basidioles dominant, in shape similar to basidia, but smaller, 12.6–15.7 × 2.9–5.2 μm.
Basidiospores slim allantoid, slightly curved, hyaline, thin- to slightly thick-walled, smooth, IKI–, CB–, 4.6–5.2 × 0.8–1.3 μm, L = 5 μm, W = 0.99 μm, Q = 4.96–5.25 (n = 60/2).
Brown rot.
China. Sichuan Province, Jiuzhaigou County, Jiuzhaigou Nature Reserve, on fallen trunk of Abies sp., 20.IX.2020, Cui 18562 (
Cyanosporus rigidus is characterised by corky, hard corky to rigid basidiomata with a buff yellow to clay-buff and tomentose pileal surface when fresh, becoming olivaceous buff to greyish brown when dry, smaller and cylindrical to allantoid basidiospores (3.7–4.2 × 0.9–1.3 μm).
China. Yunnan Province, Yulong County, Laojun Mountain, Jiushijiu Longtan, on fallen trunk of Abies sp., 15.IX.2018, Cui 17032 (
Rigidus (Lat.): referring to the rigid basidiomata.
Basidiomata annual, pileate, corky, without odour or taste when fresh, becoming hard corky to rigid upon drying. Pileus flabelliform, projecting up to 1.6 cm, 3.8 cm wide and 0.6 cm thick at base. Pileal surface tomentose, buff yellow to clay-buff, when fresh, becoming smooth, rugose, olivaceous buff to greyish brown when dry; margin obtuse. Pore surface white to cream when fresh, becoming buff-yellow to pinkish buff when dry; sterile margin narrow to almost lacking; pores angular, 5–8 per mm; dissepiments thin, entire to lacerate. Context cream to buff, hard corky, up to 4 mm thick. Tubes cream to pinkish buff, brittle, up to 5 mm long.
Hyphal system monomitic; generative hyphae with clamp connections, IKI–, CB–; hyphae unchanged in KOH.
Generative hyphae hyaline, thin- to slightly thick-walled with a wide lumen, rarely branched, loosely interwoven, 2.2–5 μm in diam.
Generative hyphae hyaline, thin- to slightly thick-walled with a wide lumen, occasionally branched, interwoven, 2–4 μm in diam. Cystidia and cystidioles absent. Basidia clavate, bearing four sterigmata and a basal clamp connection, 12.4–14.8 × 3–4.2 μm; basidioles dominant, in shape similar to basidia, but smaller, 11.8–13.9 × 2.6–4 μm.
Basidiospores allantoid to cylindrical, slightly curved, hyaline, thin- to slightly thick-walled, smooth, IKI–, CB–, (3.5–)3.7–4.2 × (0.8–)0.9–1.3(–1.4) μm, L = 3.94 μm, W = 1.09 μm, Q = 3.66 (n = 60/1).
Brown rot.
Cyanosporus subungulatus is characterised by shell-shaped pileus with a pale mouse-grey to ash-grey pileal surface when fresh, dark-grey to mouse-grey when dry, allantoid to cylindrical and slightly curved basidiospores (4.5–5.2 × 1.1–1.4 μm).
China. Yunnan Province, Yangbi County, Shimenguan Nature Reserve, on fallen trunk of Pinus sp., 6.IX.2019, Cui 18046 (
Subungulatus (Lat.): referring to the species resembling Cyanosporus ungulatus in morphology.
Basidiomata annual, pileate, soft corky, without odour or taste when fresh, becoming corky to fragile and light in weight upon drying. Pileus shell-shaped, projecting up to 1.7 cm, 2.8 cm wide and 1.2 cm thick at base. Pileal surface velutinate, pale mouse-grey to ash-grey when fresh, becoming smooth, rugose, dark-grey to mouse-grey when dry; margin obtuse. Pore surface white to cream when fresh, becoming cream to pinkish buff when dry; sterile margin narrow to almost lacking; pores round, 4–6 per mm; dissepiments thin, entire to lacerate. Context white to cream, soft corky, up to 5 mm thick. Tubes pale mouse-grey to ash-grey, fragile, up to 6 mm long.
Hyphal system monomitic; generative hyphae with clamp connections, IKI–, CB–; hyphae unchanged in KOH.
Generative hyphae hyaline, slightly thick-walled with a wide lumen, rarely branched, loosely interwoven, 2.5–6.4 μm in diam.
Generative hyphae hyaline, slightly thick-walled with a wide lumen, occasionally branched, interwoven, 2–4.2 μm in diam. Cystidia and cystidioles absent. Basidia clavate, bearing four sterigmata and a basal clamp connection, 13.6–17.8 × 3–5.5 μm; basidioles dominant, in shape similar to basidia, but smaller, 12.8–17.2 × 2.4–5.2 μm.
Basidiospores allantoid to cylindrical, slightly curved, hyaline, thin- to slightly thick-walled, smooth, IKI–, CB–, (4.3–)4.5–5.2 × 1.1–1.4 μm, L = 4.73 μm, W = 1.22 μm, Q = 3.48–3.66 (n = 60/2).
Brown rot.
China, Yunnan Province, Xichou County, Xiaoqiaogou Nature Reserve, on fallen angiosperm trunk, 14.I.2019, Zhao 10833 (
Cyanosporus tenuicontextus is characterised by flabelliform pileus with a velutinate, cream to pinkish buff with a little blue tint pileal surface when fresh, becoming glabrous, light vinaceous grey to pale mouse-grey when dry, small and round pores (6–8 per mm), thin context (up to 0.8 mm) and allantoid basidiospores (3.8–4.3 × 0.8–1.2 μm).
China. Yunnan Province, Lanping County, Tongdian Town, Luoguqing, on fallen trunk of Pinus sp., 19.IX.2017, Cui 16280 (
Tenuicontextus (Lat.): referring to the species having thin context.
Basidiomata annual, pileate, soft corky, without odour or taste when fresh, becoming corky to fragile and light in weight upon drying. Pileus flabelliform, projecting up to 1.3 cm, 3.2 cm wide and 0.5 cm thick at base. Pileal surface velutinate, cream to pinkish buff with a little blue tint when fresh, becoming glabrous, light vinaceous grey to pale mouse-grey when dry; margin acute. Pore surface white to cream when fresh, becoming pinkish buff to buff when dry; sterile margin narrow to almost lacking; pores round, 6–8 per mm; dissepiments thin, entire to lacerate. Context cream to buff, soft corky, up to 0.8 mm thick. Tubes pale mouse-grey to buff, fragile, up to 4.3 mm long.
Hyphal system monomitic; generative hyphae with clamp connections, IKI–, CB–; hyphae unchanged in KOH.
Generative hyphae hyaline, thin- to slightly thick-walled with a wide lumen, occasionally branched, loosely interwoven, 2.3–5.5 μm in diam.
Generative hyphae hyaline, thin- to slightly thick-walled with a wide lumen, occasionally branched, interwoven, 2–4 μm in diam. Cystidia absent; cystidioles present, fusoid, thin-walled, 9.5–14.6 × 2.8–3.4 μm. Basidia clavate, bearing four sterigmata and a basal clamp connection, 11.7–16.8 × 3.4–4.3 μm; basidioles dominant, in shape similar to basidia, but smaller, 10.6–14.7 × 2.9–3.6 μm.
Basidiospores allantoid, slightly curved, hyaline, thin- to slightly thick-walled, smooth, IKI–, CB–, (3.7–)3.8–4.3 × 0.8–1.2 μm, L = 3.97 μm, W = 1.02 μm, Q = 3.78–4.26 (n = 60/2).
Brown rot.
China. Yunnan Province, Yuxi, Xinping County, Mopanshan National Forest Park, on angiosperm stump, 16.I.2017, Zhao 813 (
In the current phylogenetic analyses based on the combined datasets of ITS+TEF and ITS+nLSU+mtSSU+nSSU+RPB1+RPB2+TEF sequences, species of Cyanosporus formed a highly supported lineage, distant from Postia and other brown-rot fungal genera (Figs
A list of species, specimens, and GenBank accession number of sequences used for phylogenetic analyses in this study.
Species | Sample no. | Locality | GenBank accessions | References | ||||||
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ITS | nLSU | mtSSU | nSSU | RPB1 | RPB2 | TEF | ||||
Amaropostia hainanensis | Cui 13739 (holotype) | China | KX900909 | KX900979 | KX901053 | KX901123 | KX901171 | KX901223 |
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A. stiptica | Cui 10043 | China | KX900906 | KX900976 | KX901046 | KX901119 | KX901167 | KX901219 |
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Amylocystis lapponica | HHB-13400 | United States | KC585237 | KC585059 |
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A. lapponica | OKM-4418 | United States | KC585238 | KC585060 |
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Antrodia serpens | Dai 7465 | Luxembourg | KR605813 | KR605752 | KR606013 | KR605913 | KR610832 | KR610742 |
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A. tanakae | Cui 9743 | China | KR605814 | KR605753 | KR606014 | KR605914 | KR610833 | KR610743 |
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Calcipostia guttulata | Cui 10018 | China | KF727432 | KJ684978 | KX901065 | KX901138 | KX901181 | KX901236 | KX901276 |
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C. guttulata | Cui 10028 | China | KF727433 | KJ684979 | KX901066 | KX901139 | KX901182 | KX901237 | KX901277 |
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Cyanosporus alni | Petr Vampola 12.10.1995 (holotype) | Slovakia | MG137026 |
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C. alni | Cui 7185 | China | KX900879 | KX900949 | KX901017 | KX901092 | KX901155 | KX901202 | KX901254 |
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C. alni | Dai 14845 | Poland | KX900880 | KX900950 | KX901018 | KX901093 | KX901156 | KX901203 | KX901255 |
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C. arbuti | Viacheslav Spirin 8327 (holotype) | United States | MG137039 | MG137132 |
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C. auricomus | Cui 13518 | China | KX900887 | KX900957 | KX901025 | KX901100 | KX901209 |
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C. auricomus | Cui 13519 | China | KX900888 | KX900958 | KX901026 | KX901101 |
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C. auricomus | Tuomo Niemelä 8310 (holotype) | Finland | MG137040 |
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C. bifarius | Viacheslav Spirin 6402 (holotype) | Russia | MG137043 | MG137133 |
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C. bifarius | Cui 17534 | China | OL423598* | OL423608* | OL437195* | OL423620* | OL444985* | OL446999* | OL444994* | Present study |
C. bifarius | Cui 16277 | China | OL423599* | OL423609* | OL437196* | OL423621* | OL444986* | OL447000* | OL444995* | Present study |
C. bubalinus | Cui 16976 | China | MW182172 | MW182225 | MW182208 | MW182189 | MW191547 | MW191563 | MW191530 |
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C. bubalinus | Cui 16985 (holotype) | China | MW182173 | MW182226 | MW182209 | MW182190 | MW191548 | MW191564 | MW191531 |
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C. caesiosimulans | Viacheslav Spirin 4199 | Russia | MG137061 | MG137140 |
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C. caesiosimulans | Otto Miettinen 16976 (holotype) | United States | MG137054 | MG137137 |
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C. caesius | Gerhard Schuster 51 (neotype) | Germany | MG137045 |
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C. caesius | Otto Miettinen 14156 | Finland | MG137048 | MG137134 |
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C. caesius | Cui 18630 | France | OL423600* | OL423610* | OL437197* | OL423622* | OL444996* | Present study | ||
C. caesius aff GB | K 32713 | United Kingdom | AY599576 |
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C. caesius aff GB | K 32425 | United Kingdom | AY599575 |
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C. coeruleivirens | Otto Miettinen 12214 | Indonesia | MG137063 |
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C. coeruleivirens | Dai 19220 | China | MW182174 | MW182227 | MW182210 | MW182191 | MW191549 | MW191532 |
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C. comatus | Otto Miettinen 14755,1 (holotype) | United States | MG137066 |
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C. cyanescens | Otto Miettinen 13602 (holotype) | Finland | MG137067 | MG137142 |
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C. cyanescens | Otto Miettinen 15919,2 | Spain | MG137071 | MG137144 |
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C. flavus | Cui 18547 | China | MW448564 * | MW448561 * | MW448557 * | MW452596 * | MW452599 * | MW452601 | Present study | |
C. flavus | Cui 18562 (holotype) | China | MW448565 * | MW448562 * | MW448558 * | MW452597 * | MW452600 * | MW452602 | Present study | |
C. fusiformis | Cui 10775 | China | KX900868 | KX900938 | KX901006 | KX901081 | KX901191 | KX901245 |
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C. fusiformis | Dai 15036 (holotype) | China | KX900867 | KX900937 | KX901005 | KX901080 | KX901190 | KX901244 |
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C. glaucus | Viacheslav Spirin 5317 | Russia | MG137078 |
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C. glaucus | Viacheslav Spirin 6580 (holotype) | Russia | MG137081 | MG137145 |
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C. gossypinus | Bernard Rivoire 6658 | France | MG137146 |
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C. hirsutus | Cui 17083 (holotype) | China | MW182179 | MW182233 | MW182214 | MW182197 | MW191554 | MW191568 | MW191538 |
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C. hirsutus | Cui 17343 | China | OL423601* | OL423611* | OL437198* | OL423623* | OL444987* | OL447001* | OL444997* | Present study |
C. hirsutus | Cui 17342 | China | OL423602* | OL423612* | OL437199* | OL423624* | OL444988* | OL447002* | OL444998* | Present study |
C. livens | Viacheslav Spirin 8728 | United States | MG137090 | MG137150 |
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C. livens | Otto Miettinen 17177 (holotype) | United States | MG137082 | MG137147 |
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C. luteocaesia | Bernard Rivoire 2605 | France | MG137091 |
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C. magnus | Dai 21105 | China | OL423603* | OL423613* | OL437200* | OL423625* | OL444989* | OL447003* | OL444999* | Present study |
C. magnus | Cui 16983 | China | MW182180 | MW182234 | MW182215 | MW182198 | MW191555 | MW191569 | MW191539 |
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C. magnus | Otto Miettinen 10634 (holotype) | China | KC595944 | KC595944 | MG137151 |
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C. mediterraneocaesius | LY BR 4274 | France | KX900886 | KX901024 | KX901099 |
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C. microporus | Cui 11014 (holotype) | China | KX900878 | KX900948 | KX901016 | KX901091 | KX901201 |
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C. microporus | Dai 11717 | China | KX900877 | KX900947 | KX901015 | KX901090 | KX901200 |
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C. nothofagicola | Cui 16697 (holotype) | Australia | MW182181 | MW182235 | MW182216 | MW182199 | MW191556 | MW191570 | MW191540 |
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C. nothofagicola | Dai 18765 | Australia | MW182182 | MW182236 | MW182217 | MW182200 | MW191557 | MW191541 |
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C. piceicola | Cui 10626 (holotype) | China | KX900862 | KX900932 | KX901001 | KX901075 | KX901185 |
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C. piceicola | Cui 12158 | China | KX900866 | KX900936 | KX901004 | KX901079 | KX901153 | KX901189 | KX901243 |
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C. populi | Otto Miettinen 17043 (holotype) | United States | MG137092 | MG137153 |
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C. populi | Cui 17087a | China | MW182183 | MW182237 | MW182218 | MW182201 | MW191558 | MW191571 | MW191542 |
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C. populi | Dai 18934 | China | OL423604* | OL423614* | OL437201* | OL423626* | OL444990* | OL447004* | OL445000* | Present study |
C. populi | Cui 17557 | China | OL423605* | OL423615* | OL437202* | OL423627* | OL444991* | OL447005* | OL445001* | Present study |
C. rigidus | Cui 17032 (holotype) | China | OL423606 * | OL423617 * | OL437204 * | OL423629 * | OL444993 * | OL445003 * | Present study | |
C. simulans | Otto Miettinen 20422 | Finland | MG137110 | MG137160 |
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C. simulans | Tuomo Niemelä 8846 (holotype) | Finland | MG137103 |
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C. subcaesius | Josef Vlasák 0110/24 | Czech Republic | MG137117 | MG137164 |
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C. subcaesius | Alix David 652 (isotype) | France | MG137116 |
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C. subhirsutus | Cui 11330 | China | KX900873 | KX900943 | KX901011 | KX901086 | KX901196 | KX901250 |
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C. subhirsutus | Dai 14892 (holotype) | China | KX900871 | KX900941 | KX901009 | KX901084 | KX901194 | KX901248 |
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C. submicroporus | Cui 16306 | China | MW182184 | MW182239 | MW182220 | MW182203 | MW191560 | MW191573 | MW191544 |
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C. submicroporus | Cui 18156 (holotype) | China | MW182186 | MW182241 | MW182222 | MW182205 | MW191574 |
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C. subungulatus | Cui 18046 (holotype) | China | MW448566 * | MW448563 * | MW448560 * | MW448559 * | MW452598 * | MW452603 | Present study | |
C. subungulatus | Zhao 10833 | China | MW742586 * | OL423616 * | OL437203 * | OL423628 * | OL444992 * | OL445002 * | Present study | |
C. subviridis | Viacheslav Spirin 8774a | United States | MG137120 | MG137166 |
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C. subviridis | Reijo Penttilä 14376 | Finland | MG137165 |
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C. tenuicontextus | Cui 16280 (holotype) | China | OL423607 * | OL423618 * | OL437205 * | OL423630 * | OL445004 * | Present study | ||
C. tenuicontextus | Zhao 813 | China | MG231802 * | OL423619 * | OL437206 * | OL423631 * | OL445005 * | Present study | ||
C. tenuis | Cui 10788 (holotype) | China | KX900885 | KX900955 | KX901023 | KX901098 | KX901161 | KX901208 |
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C. tenuis | Dai 12974 | China | KX900884 | KX900954 | KX901022 | KX901097 | KX901160 | KX901207 | KX901258 |
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C. tricolor | Cui 12233 (holotype) | China | KX900876 | KX900946 | KX901014 | KX901089 | KX901199 | KX901253 |
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C. tricolor | Cui 10790 | China | KX900875 | KX900945 | KX901013 | KX901088 | KX901198 | KX901252 |
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C. ungulatus | Cui 10778 | China | KX900870 | KX900940 | KX901008 | KX901083 | KX901193 | KX901247 |
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C. ungulatus | Dai 12897 (holotype) | China | KX900869 | KX900939 | KX901007 | KX901082 | KX901154 | KX901192 | KX901246 |
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C. yanae | Heikki Kotiranta 27606 | Russia | MG137122 | MG137168 |
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C. yanae | Heikki Kotiranta 27454 (holotype) | Russia | MG137121 | MG137167 |
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Cystidiopostia hibernica | Cui 2658 | China | KX900905 | KX900975 | KX901045 | KX901118 | KX901218 |
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C. inocybe | LY BR 3703 | France | KX900903 | KX900973 | KX901044 | KX901116 | KX901267 |
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C. pileata | Cui 10034 | China | KX900908 | KX900956 | KX901050 | KX901122 | KX901170 | KX901222 | KX901269 |
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Fuscopostia duplicate | Dai 13411 (holotype) | China | KF699125 | KJ684976 | KR606027 | KR605928 | KX901174 | KR610845 | KR610756 |
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F. fragilis | JV 0610-8 | Czech | JF950573 |
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F. lateritia | Dai 2652 | China | KX900913 | KX900983 |
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F. leucomallella | Cui 9599 | China | KF699123 | KJ684983 | KX901056 | KX901129 | KX901176 | KX901228 | KX901272 |
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Jahnoporus brachiatus | X 3232 | Russia | KU165781 |
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J. hirtus | Spinosa 10 X 2014 | United States | KU165784 | KY949044 |
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J. oreinus | X 3241 | Russia | KU165785 |
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Oligoporus rennyi | TN-6645 | Finland | KC595929 | KC595929 |
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O. sericeomollis | Cui 9870 | China | KX900920 | KX900990 | KX901068 | KX901141 | KX901184 |
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Osteina obducta | Cui 10074 | China | KX900924 | KX900994 | KX901071 | KX901144 | KX901240 |
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O. undosa | Dai 7105 | China | KX900921 | KX900991 | KX901069 | KX901142 | KX901238 |
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Postia amurensis | Dai 903 (holotype) | China | KX900901 | KX900971 | KX901042 |
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P. hirsuta | Cui 11237 (holotype) | China | KJ684970 | KJ684984 | KX901038 | KX901113 | KX901266 |
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P. lactea | Cui 12141 | China | KX900892 | KX900962 | KX901029 | KX901104 | KX901163 | KX901211 | KX901260 |
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P. lowei | Cui 9585 | China | KX900898 | KX900968 | KX901035 | KX901110 |
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P. ochraceoalba | Cui 10802 (holotype) | China | KM107903 | KM107908 | KX901041 | KX901115 | KX901216 | Shen et al. 2015 | ||
P. sublowei | Cui 9597 (holotype) | China | KX900900 | KX900970 | KX901037 | KX901112 | KX901265 |
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P. tephroleuca | Dai 12610 | Finland | KX900897 | KX900967 | KX901034 | KX901109 | KX901166 | KX901214 | KX901263 |
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Spongious gloeoporus | Cui 10401 | China | KX900915 | KX900985 | KX901060 | KX901133 | KX901232 | |||
S. floriformis | Cui 10292 | China | KM107899 | KM107904 | KX901058 | KX901131 | KX901178 | KX901230 | KX901274 | |
S. floriformis | Dai 13887 | China | KX900914 | KX900984 | KX901057 | KX901130 | KX901177 | KX901229 | KX901273 |
In the phylogenetic trees, Cyanosporus flavus grouped together with C. fusiformis, C. subungulatus and C. ungulatus (Figs
Phylogenetically, Cyanosporus rigidus form a separate lineage different from other species in the genus. Morphologically, C. submicroporus share similar pores and basidiospores with C. rigidus, but C. submicroporus differs by having cream to pinkish buff pileal surface and white to smoke grey pore surface when fresh, buff to buff-yellow pileal surface and buff to olivaceous buff pore surface when dry. Cyanosporus auricomus and C. luteocaesius resemble C. rigidus in morphology by producing yellow-colored basidiomata, but C. auricomus differs from C. rigidus by having a hirsute pileal surface and larger basidiospores (4.4–5.6 × 1.5–1.8 μm;
Phylogenetically, C. tenuicontextus is closely related to C. caesiosimulans, C. cyanescens, C. populi, C. subviridis and C. yanae (Figs
The natural distribution of plant-associated fungi across broad geographic ranges is determined by a combination of the distributions of suitable hosts and environmental conditions (
In the current study, 77 samples of Cyanosporus throughout China and 11 samples outside of China have been morphologically examined in detail. The specimens collected from China representing 21 species were sequenced here and referred to in our phylogeny, viz., C. alni, C. auricomus, C. bifarius, C. bubalinus, C. coeruleivirens, C. comatus, C. flavus, C. fusiformis, C. hirsutus, C. magnus, C. microporus, C. piceicola, C. populi, C. rigidus, C. subhirsutus, C. submicroporus, C. subungulatus, C. tenuicontextus, C. tenuis, C. tricolor and C. ungulatus. Another two species reported in a previous study, viz., C. glauca (=Postia glauca Spirin & Miettinen) and C. simulans (=Postia simulans (P. Karst.) Spirin & Rivoire;
In summary, we performed a comprehensive study on the species diversity and phylogeny of Cyanosporus with an emphasis on Chinese collections. So far, 35 species are accepted in the Cyanosporus around the world, including 23 species from China. Currently, Cyanosporus is characterized by an annual growth habit, resupinate to effused-reflexed or pileate, soft corky, corky, fragile to hard corky basidiomata, velutinate to hirsute or glabrous pileal surface with blue-tinted, white to cream or yellow-colored, white to cream pore surface with round to angular pores, a monomitic hyphal system with clamped generative hyphae, and hyaline, thin- to slightly thick-walled, smooth, narrow, allantoid to cylindrical basidiospores that are usually weakly cyanophilous; it grows on different angiosperm and gymnosperm trees, causes a brown rot of wood and has a distribution in Asia, Europe, North America, Argentina in South America and Australia in Oceania (
We express our gratitude to the curators of herbaria of