Research Article |
Corresponding author: Zuo-Hong Chen ( chenzuohong@263.net ) Academic editor: Bao-Kai Cui
© 2024 Pan Long , Song-Yan Zhou, Sai-Nan Li, Fei-Fei Liu, Zuo-Hong Chen.
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:
Long P, Zhou S-Y, Li S-N, Liu F-F, Chen Z-H (2024) Three new species of Cortinarius section Delibuti (Cortinariaceae, Agaricales) from China. MycoKeys 101: 143-162. https://doi.org/10.3897/mycokeys.101.114705
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Three new species of Cortinarius section Delibuti, namely C. fibrillososalor, C. pseudosalor, and C. subtropicus are described as new to science based on morphological and phylogenetic evidences. Cortinarius pseudosalor is extremely morphologically similar to C. salor, but it differs from the latter by smaller coarsely verrucose basidiospores. Cortinarius fibrillososalor can be easily differentiated by its fibrillose pileus. The pileus of C. subtropicus becomes brown without lilac tint at maturity comparing with other members of section Delibuti. A combined dataset of ITS and LSU sequences was used for phylogenetic analysis. The phylogenetic reconstruction of section Delibuti revealed that these three new species clustered and formed independent lineages with full support respectively. A key to the three new species and related species of section Delibuti is provided in this work.
Morphology, new taxa, phylogeny, taxonomy
The genus Cortinarius (Pers.) Gray (Cortinariaceae, Agaricales), which is known for its high species diversity, comprises more than 3000 taxa and exhibits a global distribution (
Cortinarius sect. Delibuti (Fr.) Sacc., typified by C. delibutus Fr., is widely distributed (
The research on Cortinarius has mainly been conducted in Europe and North America, while it is still lacking in East Asia (
The specimens were collected from central and southwestern China during 2012–2022. The vouchers are all deposited in the Mycological Herbarium of Hunan Normal University (
List of sequences of Cortinarius used for phylogenetic analyses. The sequences newly generated in this study are in bold, and all type specimens are highlighted with an asterisk.
Species | Voucher | Locality | GenBank Accession No. | Reference | |
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ITS | LSU | ||||
Cortinarius anomalus | TUB011883 | Europe, Germany | AY669645 | AY669645 |
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C. anomalus * | CFP1154 (S) | Europe, Ångermanland | KX302224 | – |
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C. barlowensis * | JFA13140 | North America | FJ717554 | – |
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C. bolaris | T40 | Europe, Norway | KC842426 | KC842496 |
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C. bolaris * | CFP1008 | Europe | KX302233 | – |
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C. bolaris | TUB0118524 | Europe, Germany | AY669596 | AY669596 |
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C. calaisopus * | PDD 94050 | New Zealand, Dunedin | NR157880 | MH108373 | Genbank |
C. calaisopus | PDD103678/CO2106 | New Zealand | KF727395 | KF727338 |
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C. camphoratus | SMI193 | North America, Canada | FJ039626 | – |
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C. delibutus | F17048 | North America, Canada | FJ717515 | FJ717515 |
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C. delibutus | SAT01-301-12 | North America, USA | FJ717513 | – |
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C. dysodes | PDD70499/CO1038 HT | New Zealand | GU233340 | GU233394 |
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C. ferrusinus * | JB8106 13 | Europe | KY657254 | – | Genbank |
C. fibrillososalor * |
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East Asia, China, Hunan | OR647481 | OR647506 | This study |
C. fibrillososalor |
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East Asia, China: Hunan | OR647485 | OR647507 | This study |
C. fibrillososalor |
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East Asia, China, Hunan | OR647483 | – | This study |
C. fibrillososalor |
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East Asia, China, Hunan | OR647355 | OR647497 | This study |
C. fibrillososalor |
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East Asia, China, Hunan | OR660685 | OR647503 | This study |
C. illibatus | HMJAU48760 | East Asia, China, Heilongjiang | MW911735 | – |
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C. illibatus | OS574 | Europe | KC842441 | KC842511 |
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C. pseudocamphoratus * | HMJAU48694 | East Asia, China, Xizang | NR_176776 | – |
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C. putorius | TN07411 HT | North America, USA | KR011124 | – |
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C. rotundisporus | PDD96298/ JAC12057 | New Zealand | MH101550 | MH108389 |
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C. rotundisporus | PERTH 05255074 | Australia | AY669612 | AY669612 |
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C. salor | TUB011838 | Europe, Germany | AY669592 | AY669592 |
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C. spilomeus * | S: CFP1137 | Europe | KX302267 | – |
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C. spilomeus | TUB011523 | Europe | AY669654 | AY669654 |
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C. pseudosalor |
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East Asia, China, Hunan | OR647352 | – | This study |
C. pseudosalor * |
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East Asia, China, Hubiei | OR660686 | OR647504 | This study |
C. pseudosalor |
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East Asia, China, Hubiei | OR660688 | OR647505 | This study |
C. subsalor | HMJAU48758 | East Asia, China, Zhejiang | MW911733 | – |
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C. subsalor * | HMJAU48759 | East Asia, China, Zhejiang | MW911734 | – |
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C. subtortus | F16111 | North America | FJ157044 | FJ157044 |
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C. subtortus | TUB011382 | Europe | AY174857 | AY174857 |
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C. subtropicus |
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East Asia, China, Hunan | OR647356 | OR647498 | This study |
C. subtropicus |
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East Asia, China, Guangxi | OR647491 | OR647509 | This study |
C. subtropicus |
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East Asia, China, Hunan | OR660684 | OR647501 | This study |
C. subtropicus |
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East Asia, China, Hunan | OR660687 | OR647502 | This study |
C. subtropicus * |
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East Asia, China, Hunan | OR647488 | OR647508 | This study |
C. tasmacamphoratus | HO A20606A0 | Australia, Tasmania | AY669633 | AY669633 |
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C. tessiae | PDD107517/CO1450 | New Zealand | MG019356 | MG019356 |
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C. tessiae | PDD72611 | New Zealand | HM060317 | HM060316 | Genbank |
C. tetonensis * | JFA10350 | North America | MZ580436 | – |
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C. tibeticisalor * | HMJAU48764 | East Asia, China, Xizang | MW911730 | – |
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C. tibeticisalor | HMJAU48762 | East Asia, China: Xizang | MW911731 | – |
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C. tibeticisalor | HMJAU48763 | East Asia, China, Xizang | MW911732 | – |
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C. viridipileatus | OTA61977 | New Zealand | MK546592 | MK546595 |
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C. viridipileatus | OTA64087 | New Zealand | MK546593 | MK546596 |
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The descriptions of macromorphological characters were based on field records and photographs. Color codes were used following
Total genomic DNA was extracted by a Fungal DNA Mini Kit (Omega, USA). ITS 4 and ITS 5 (
The sequences newly generated in this study and downloaded from GenBank were used for phylogenetic analysis (Table
In the concatenated dataset (ITS+LSU), a total of 78 sequences (48 ITS, 30 LSU) from 48 samples were used for phylogenetic analyses among sect. Delibuti, sect. Subtorti, sect. Camphorati, sect. Bolares, sect. Spilomei, and sect. Anomali, of which 24 sequences (13 ITS, 11 LSU) were newly yielded in this study (Table
Phylogenetic tree of Cortinarius sect. Delibuti inferred from a combined matrix of ITS and LSU through maximum likelihood and Bayesian inference. Bayesian posterior probabilities (PP) > 0.90 and bootstrap values (BP) >85% are reported at the nodes (PP/BP); “–” indicates that the support value was less than the respective threshold. The three newly described species are highlighted in bold. Aus: Australia; EA: East Asia; Eur: Europe; NAm: North America; NZ: New Zealand.
The phylogenetic relationship of sections within the genus Cortinarius in the present study was unclear and weakly supported. In the multi-locus tree, the monophyly of sect. Delibuti was supported with well-supported values (BP = 99%, PP = 1.00), including 12 species. section Camphorati was also monophyletic with fully supported values (BP = 100%, PP = 1.00), emcompassing 5 species. In sect. Delibuti, C. delibutus, C. illibatus, C. salor, C. subsalor, C. tibeticisalor, and three novel species, namely C. fibrillososalor, C. pseudosalor, and C. subtropicus, formed a monophyletic lineage (BP = 95%, PP = 1.00). Cortinarius fibrillososalor, C. subtropicus, and C. tibeticisalor formed a clade only be found in East Asia (BP = 99%, PP = 1.00), while C. tibeticisalor has a special olive-green tint, was only distributed in Tibetan Plateau (
Fibrillososalor (Latin) refers to the species morphologically similar to Cortinarius salor, but with fibrils on the pileus.
Differs from the other species of sect. Delibuti from its fibrillose pileus.
Basidiomes small to medium-sized, telamonia-like, development type stipiocarpic. Pileus 2.9–5.2 cm, at first broadly convex, then lower convex to plane, broadly umbonate at the centre, margin incurved or decurved to upturned; at first violaceous (17B6–17B8), tinged brown (5B4–5C6) at the centre then becoming whitish mauve (16A1–16A2), finely fibrillose, with brown (5A5–5C7) universal veil remains at margin; surface silky when dry or glutinous when wet. Context thin, creamy white, soft, beige (3A1–A2) when bruised. Lamellae adnate to adnexed, lilac (17A2–17B2) to brownish (6C5–6D7), moderately distant, sometimes margin wavy. Stipe cylindrical to clavate, bend, gradually slender to the apex, 3.4–5.9 cm long, 0.4–0.8 cm wide, violaceous (17A4–17B5) when young then fading to whitish mauve (16A2–16A3) tint, leaving an ochraceous (5B6–5D8) ringon the upper stem, hollow. Odour indistinct.
Basidiospores [100/5/5] (6.5–) 7.0–8.8 (–9.2) × (5.0–) 5.9–7.2 (–8.1) μm, av. 8.1 × 6.5 μm, Q = 1.14 (1.16) – 1.31 (1.45), Qm = 1.24 ± 0.02, broadly globose to long ellipsoid, rarely subglobose, yellowish brown, moderately verrucose, without amyloid and dextrinoid reaction. Basidia (27–) 28–35 × (8–) 9–11 μm, 4-spored, sterigmata up to 2.4–3.7 μm, clavate to subcylindrical, colourless or with amber yellow oily inclusions or granules. Pileipellis duplex, hyphae 4–8 μm wide, epicutis strongly gelatinous, 68–128 μm thick, composed of colourless or amber yellow, irregularly arranged and strongly interwoven hyphae, hypocuits 25–38 μm thick, composed of colourless or amber yellow, nearly parallel cylindrical hyphae. Lamellar edges fertile. Cystidia absent. Lamellar trama regular, 40–80 μm thick, composed of parallel arranged hyphae, hyphae 3–6 μm wide. Stipitipellis gelatinous, stipe hyphae 3–6 μm wide, thin-walled, cylindrical, interwoven. Clamp connections present in all tissues.
Solitary to gregarious on soil in evergreen broad-leaved forest, known from Hunan, China; July to September.
China, Hunan Province: Sangzhi County, Badagongshan National Nature Reserve, at 29.769154°N, 110.086577°E, alt. 1405 m, 31 July 2020, Z.H. Chen, P. Long and S.N. Li, (
Cortinarius fibrillososalor can be differentiated from other species of section Delibuti for its fibrillose pileus, usually under evergreen broad-leaved forest at 1405–1500m. In addition, basidiospores broadly globose to long ellipsoid, rarely subglobose while other members in this section usually subglobose to broadly ellipsoid.
Pseudosalor (Latin) refers to the species morphologically similar to Cortinarius salor.
This species differs from other species in sect. Delibuti for its high morphological similarity with C. salor, but having smaller coarsely verrucose basidiospores.
Basidiomes small to medium-sized, development type stipiocarpic. Pileus 2.8–6.5 cm, at first broadly convex, then lower convex to plane, margin incurved when young, decurved to upturned at maturity; bluish violaceous (18A3–18C5) when young, tinge of white at the centre when chapped, later fading to ochraceous grey (5B6–5C7) when old with brown (5B8–5C8) universal veil remains at margin; dry, viscid. Context dirty white, soft. Lamellae adnexed, pale yellow (1A2) with lilac tint (16A1–16A2) then brownish (5B6–5D7), moderately distant, sometimes margin wavy. Stipe clavate, gradually slender to the apex, 4–8.4 cm long, 0.4–1.0 cm wide, violaceous (16A2–16A4) when young then fading to upper dirty white, whitish mauve (16A2) at base, leaving an ochraceous ring (5B8–5C8) on the upper stem, hollow in centre. Odour indistinct.
Basidiospores [60/3/3] (7.3–) 7.4–8.4 × (5.7–) 6.0–7.4 (–7.5) μm, av. 7.9 × 6.7 μm, Q = (1.11) 1.12– (1.26) 1.27, Qm = 1.18 ± 0.11, subglobose to broadly ellipsoid, yellowish brown, coarsely verrucose, without amyloid and dextrinoid reaction. Basidia (29–) 30–38 × (8–) 9–12 μm, 4-spored, sterigmata up to 3.7–5.0 μm, clavate to subcylindrical, colourless or with amber yellow granules. Pileipellis duplex obviously, hyphae 2–6 μm wide, epicutis gelatinous, 50–75 μm thick, composed of colourless or amber yellow, moderately interwoven hyphae, hypocuits 50–75 μm thick, composed of colourless or amber yellow, hyphae nearly parallel cylindrical. Lamellar edges fertile. Cystidia absent. Lamellar trama regular, 45–55 μm thick, composed of hyphae and inflated cells, hyphae 2–5 μm wide, inflated cells 14–24 × 5–9 μm. Stipitipellis gelatinous, stipe hyphae 2–7 μm wide, thin-walled, cylindrical, weakly interwoven. Clamp connections present in all tissues.
Solitary to gregarious on soil in coniferous and broad-leaved mixed forest or evergreen broad-leaved forest, known from Hunan and Hubei, China; August.
China, Hunan Province: Yongshun County, Xiaoxi National Nature Reserve, at 28.4215–28.5355°N, 110.650–110.2135°E, alt. 1000–1300 m, 30 August 2014, P. Zhang, (
Cortinarius pseudosalor is easily misidentified as C. salor for their high morphological similarity, except the former has smaller coarsely verrucose basidiospores. Besides, C. pseudosalor distributed in Central China under coniferous and broad-leaved mixed forest or evergreen broad-leaved forest at alt. 1000–1413 m.
Subtropicus (Latin) refers to subtropical distribution range of the species.
Differs from the other species of sect. Delibuti species in having an epicutis pileipellis that can be easily separated from the context of the pileus.
Basidiomes small, development type stipiocarpic. Pileus 2.1–4.6 cm, at first broadly convex, then lower convex to plane, broadly umbonate at the centre, margin incurved; at first violaceous (15A4–15B7), tinged brown (6A5–6C7) at the centre then becoming orange brown (5B2–5B6), brown (5A4–5B6) universal veil remains at margin; surface smooth when dry or glutinous when wet, pileipellis is easy to separate. Context thin, creamy white, soft, beige (3A1–A2) when bruised. Lamellae adnate, bluish violet (18A2–18B2) with pale greyish (18B1) to brownish (5A4–5B7), rust brown (5C7) when dry, moderately distant. Stipe cylindrical to weakly clavate, bend, gradually slender to the apex, 6.4–7.2 cm long, 0.5–1.0 cm wide, lilac (15A2–15B2) when young, dirty white at maturity, leaving an ochraceous (5D7) to orange (5B6) ring on the upper stem, hollow in centre, crumbly. Odour indistinct.
Basidiospores [120/6/6] (6.6–) 7.0–9.1 (–10.3) × (5.9–)6.1–7.9 (– 10.3) μm, av. 7.8 × 6.4 μm, Q = 1.10–1.38 (1.41), Qm = 1.24 ± 0.01, subglobose to ellipsoid, yellowish brown, moderately verrucose, without amyloid and dextrinoid reaction. Basidia 32–48 × 9–12 μm, 4-spored, sterigmata 2.8–4.9 μm, clavate to subcylindrical, colourless or with amber yellow oily inclusions. Pileipellis duplex, hyphae 4–8 μm wide, epicutis gelatinous, 30–40 μm thick, composed of colourless or amber yellow, irregularly arranged and moderately interwoven hyphae, hypocuits 130–200 μm thick, composed of colourless or amber yellow, nearly parallel cylindrical hyphae. Lamellar edges fertile. Cystidia absent. Lamellar trama regular, 40–80 μm thick, composed of parallel arranged hyphae, hyphae 3–6 μm wide. Stipitipellis gelatinous, stipe hyphae 3–6 μm wide, thin-walled, cylindrical, subparallel arranged. Clamp connections present in all tissues of the basidiome.
Solitary to gregarious on soil in under evergreen broad-leaved forest, on the ground, known from Hunan, China; July.
China, Hunan Province: Sangzhi County, Badagongshan National Nature Reserve, at 29.683144°N, 109.754104°E, alt. 1645 m, 27 July 2020, Z.H. Chen, P. Long and S.N. Li, (
Cortinarius subtropicus has an epicutis pileipellis that can be easily separated from the context of the pileus. Besides, pileus become brown without lilac or dark olive tint at maturity comparing with other members of section Delibuti.
1 | Only distributed in the Northern Hemisphere | 2 |
– | Only distributed in the Southern Hemisphere or distributed both in Northern and Southern Hemisphere | 12 |
2 | Pileus usually ochraceous yellow without bluish hue | 3 |
– | Pileus usually bluish violet, sometimes yellow brown | 4 |
3 | Lamellae usually bluish violet at first, veil yellow to pale brown | C. delibutus |
– | Lamellae pale ochraceous with tinge of pinkish violet, veil not yellowish | C. illibatus |
4 | Distributed in Europe ± North America | 5 |
– | Distributed in China, East Asia | 8 |
5 | Growing under coniferous trees and broadleaved trees; pileus bluish violet | 6 |
– | Only growing under coniferous trees (Abies and Picea); pileus usually orange | C. largodelibutus |
6 | Basidiomes small to medium-sized, pileus deep bluish violet to ochraceous; Veil violet to ochraceous | 7 |
– | Basidiomes small, pileus yellow to olive-ochre at the centre, greyish blue to violet at margin then fading quickly; veil yellow | C. betulinus |
7 | Pileus usually olive brown; growing under coniferous trees (Picea) and broadleaved trees (Betula) | C. transiens |
– | Pileus usually deep bluish violet; growing under broadleaved trees (Quercus, Fagus, Corylus) | C. salor |
8 | Pileus usually dark olivaceous to brown at maturity; distributed in Tibetan Plateau of China | C. tibeticisalor |
– | Pileus usually ochraceous yellow at maturity; distributed in Central China ± Eastern China | 9 |
9 | Pileus with fibrils, basidiospores broadly globose to long ellipsoid, rarely subglobose | C. fibrillososalor |
– | Pileus without fibrils, basidiospores subglobose to broadly ellipsoid | 10 |
10 | Basidiospores average length >8 μm | C. subsalor |
– | Basidiospores average length <8 μm | 11 |
11 | Pileipellis is easy to separate; epicutis of pileipellis less than 40 μm thick, distributed from 1625 m to 1900 m | C. subtropicus |
– | Epicutis of pileipellis more than 50 μm thick, distributed from 1000 m to 1413 m | C. pseudosalor |
12 | Growing under trees of Nothofagus | 13 |
– | Growing under trees of Myrtaceae | 15 |
13 | Pileus glutinous, greyish yellow to greyish orange, stipe violet, then becoming white to pale brownish, basidiospores ellipsoid, distributed in Northern and Southern Hemisphere | C. illitus |
– | Pileus viscid, with a green hue; basidiospores subglobose, distributed in Southern Hemisphere | 14 |
14 | Pileus blue‒green to aerugineous; stipe blue green; distributed in Australasia | C. tessiae |
– | Pileus dark green; stipe white with a purple hue; distributed in New Zealand | C. viridipileatus |
15 | Basidiomes distinctly viscid to glutinous, stipe viscid, mainly greyish blue-green | C. rotundisporus |
– | Basidiomes weakly viscid, stipe often dry, mainly yellow-green to olive | C. calaisopus |
In this study, three species of Cortinarius sect. Delibuti, namely C. fibrillososalor, C. pseudosalor, and C. subtropicus, were described as new to science based on phylogenetic analyses and morphological characteristics. The phylogenetic relationships of C. fibrillososalor and C. subtropicus in C. sect. Delibuti were resolved with close phylogenetic relationship with C. tibeticisalor. However, the phylogenetic position of C. pseudosalor is still unclear as no supported sister relationship was revealed in the phylogenetic analysis.
Cortinarius fibrillososalor
, C. pseudosalor, C. salor, C. subsalor, C. subtropicus and C. tibeticisalor have morphological homogeneity of the basidiomes. The macromorphological characters of C. pseudosalor, and C. salor are similar to basidiomes, coloured bluish violet, while C. subsalor is coloured purple to purplish red in pileus centre. Besides, C. pseudosalor has smaller coarsely verrucose basidiospores comparing C. salor and C. subsalor (
Phylogenetic analysis was first applied to the taxonomic study of Cortinarius with ITS (
We thank Professor Ping Zhang, Master Zi-Juan Jiang and Jing Wen (Hunan Normal University) and Dr. Xiao-Bin Liu (Kunming Institute of Botany, Chinese Academy of Sciences) for specimen collection. We also thank Prof. Zhu-Liang Yang (Kunming Institute of Botany, Chinese Academy of Sciences), Dr. Zheng-Mi He and Yu-Ting Su (Hunan Normal University) for improving the manuscript.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This study was financially supported by the National Science Foundation of China (Grant No. 32170016) and the Biodiversity Survey and Assessment Project of the Ministry of Ecology and Environment, China (Grant No. 2019HJ2096001006).
Conceptualization: Zuo-Hong Chen and Pan Long; methodology: Fei-Fei Liu and Pan Long; performing the experiment: Pan Long; resources: Zuo-Hong Chen, Pan Long, Sai-Nan Li, and Song-Yan Zhou; writing – original draft preparation: Pan Long; writing – review and editing: Zuo-Hong Chen and Song-Yan Zhou; supervision: Zuo-Hong Chen; project administration: Zuo-Hong Chen; funding acquisition: Zuo-Hong Chen. All authors have read and agreed to the published version of the manuscript.
All of the data that support the findings of this study are available in the main text or Supplementary Information. The sequence data generated in this study are deposited in NCBI GenBank.
Multiple sequence alignment
Data type: fas