Research Article |
Corresponding author: Rui-Qing Ji ( jiruiqingjrq@126.com ) Academic editor: Olivier Raspé
© 2020 Meng-Le Xie, Tie-Zheng Wei, Yong-Ping Fu, Dan Li, Liang-Liang Qi, Peng-Jie Xing, Guo-Hui Cheng, Rui-Qing Ji, Yu Li.
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:
Xie M-L, Wei T-Z, Fu Y-P, Li D, Qi L-L, Xing P-J, Cheng G-H, Ji R-Q, Li Y (2020) Three new species of Cortinarius subgenus Telamonia (Cortinariaceae, Agaricales) from China. MycoKeys 69: 91-109. https://doi.org/10.3897/mycokeys.69.49437
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Cortinarius is an important ectomycorrhizal genus that forms a symbiotic relationship with certain trees, shrubs and herbs. Recently, we began studying Cortinarius in China and here we describe three new species of Cortinarius subg. Telamonia based on morphological and ecological characteristics, together with phylogenetic analyses. Cortinarius laccariphyllus sp. nov. (section Colymbadini) is associated with broadleaf trees, with strongly hygrophanous basidiomata, special Laccaria-like lamellae and white and extremely sparse universal veil. Cortinarius neotorvus sp. nov. (section Telamonia) is associated with broadleaf trees and is easily confused with C. torvus, but can be distinguished by the colour of the fresh basidiomes and the stipe usually somewhat tapering towards the base. Cortinarius subfuscoperonatus sp. nov. (section Fuscoperonati) is associated with coniferous trees, with subglobose to broadly ellipsoid spores and is closely related to C. fuscoperonatus. A key to the new species and similar species in sections Colymbadini, Telamonia and Fuscoperonati is provided.
Ectomycorrhizal fungi, morphology, phylogeny, taxonomy
Cortinarius (Pers.) Gray is one of the most species-rich agaric genera, with reportedly more than 2250 species worldwide (
Previously, phylogenetic studies of Cortinarius have shown that many traditional infrageneric groups are artificial (Høiland & Holst-Jensen 2000;
The diverse ecosystems in China provide a conducive environment for the growth of Cortinarius species. Research, dedicated to the phylogeny and taxonomy of Chinese Cortinarius, was initiated in recent years. During field trips in the past years, many specimens of Cortinarius were collected from China. However, only two new Cortinarius species have been described and reported, based on Chinese specimens until now (
We collected specimens from northeast China and northwest China, two important floristic areas of China. Fresh basidiomata were photographed and noted under daylight in the field, dried in an oven at about 50 °C and deposited in the Herbarium of Mycology, Jilin Agricultural University (
The macroscopic characters were described from fresh basidiomata. Colour codes were taken from
We extracted the DNA from fresh tissue dried in silica gel by the NuClean PlantGen DNA Kit (CWBIO, China) and amplified the ITS region with primers ITS1F and ITS4 (
BLAST searches with the newly-generated ITS sequences were performed against NCBI (https://www.ncbi.nlm.nih.gov/) and UNITE (https://unite.ut.ee/) databases to retrieve similar sequences for the phylogenetic analyses (Table
Species | Voucher | GenBank accession No. | Locality | Reference |
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C. agathosmus TYPE | CFP536 | KC608590 | Sweden |
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C. ahsii TYPE | MM19650703 (IB) | KX882644 | USA |
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C. ahsii | JFA10303 (WTU) | KX882649 | USA |
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C. alboviolaceus | HMJAU44214 | MK552393 | China | This study |
C. alboviolaceus | HMJAU44245 | MK234572 | China |
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C. alboviolaceus | HMJAU44347 | MK552392 | China | This study |
C. alboviolaceus | F15809 | FJ157005 | Canada |
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C. armeniacus | HMJAU44408 | MK552394 | China | This study |
C. armeniacus | F16352 | FJ039573 | Canada |
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C. armillatus TYPE | F256861 (S) | NR131891 | Sweden |
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C. bulliardii | CFP499 (S) | JX114942 | Sweden |
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C. caesioarmeniacus TYPE | H7000901 | KP137498 | Canada |
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C. caesioarmeniacus | HMJAU44409 | MK552396 | China | This study |
C. caesioarmeniacus | HMJAU44403 | MK552395 | China | This study |
C. cinnabarinus | IK85-1517 (H) | JX114943 | Finland |
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C. cinnabarinus TYPE | CFP379 (S) | JX114944 | Sweden |
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C. coccineus TYPE | 435745 (GK) | JX114945 | France |
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C. colynbadinus | CFP1130 (S) | JX127302 | Sweden |
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C. colynbadinus TYPE | F248443 (S) | NR131819 | Sweden |
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C. fructuodorus | TN09-113 | KC608582 | USA |
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C. fructuodorus TYPE | H7001104 | NR131827 | USA |
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C. fuscoperonatus | SSt16-046 | MF139754 | Sweden |
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C. fuscoperonatus | CFP1470 | JX407330 | France |
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C. fuscoperonatus | CFP505 | EU433390 | Sweden | GenBank/Liimatainen |
C. laccariphyllus | HMJAU44449 | MK552380 | China | This study |
C. laccariphyllus | HMJAU44450 | MK552381 | China | This study |
C. millaresensis | XC2011-200 | MH784748 | France |
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C. millaresensis | XC2013-163 | MH784752 | France |
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C. nolaneiformis | DB886 (BP) | KJ206487 | Hungary |
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C. nolaneiformis TYPE | PRM857042 | NR131833 | Czech Republic |
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C. paragaudis TYPE | F256858 (S) | NR131814 | Norway |
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C. privignofulvus TYPE | AB00-10-128 (PC) | MH784703 | France |
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C. privignofulvus | AB04-09-192 | MH784714 | France |
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C. neotorvus | HMJAU44438 | MK552383 | China | This study |
C. neotorvus | HMJAU44441 | MK552384 | China | This study |
C. neotorvus | HMJAU44442 | MK552385 | China | This study |
C. neotorvus | HMJAU44443 | MK552386 | China | This study |
C. neotorvus | HMJAU44437 | MK552382 | China | This study |
C. rigidipes TYPE | MM1962/0062 (IB) | KJ206504 | Switzerland |
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C. rigidipes | IK95-1873 (H) | KJ206506 | Germany |
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C. subargyronotus TYPE | H7018127 | KP137494 | Sweden |
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C. subfuscoperonatus | HMJAU44446 | MK552389 | China | This study |
C. subfuscoperonatus | HMJAU44447 | MK552390 | China | This study |
C. subfuscoperonatus | HMJAU44445 | MK552388 | China | This study |
C. subfuscoperonatus | HMJAU44444 | MK552387 | China | This study |
C. subfuscoperonatus | HMJAU44448 | MK552391 | China | This study |
C. torvus | TUB 011515 | AY669668 | Germany |
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C. torvus | IK98-1973 | JX407337 | Denmark |
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C. torvus | TF01-035 | AJ889977 | Denmark | GenBank/Kjoller |
C. turgidoides | AB15-09-37 | MH784723 | France |
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C. turgidoides | AB07-09-121 | MH784717 | France |
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C. uraceomajalis | DB2291 (BP) | KJ206511 | Hungary |
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C. uraceomajalis | DB2283 (BP) | KJ206510 | Hungary |
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C. uraceomajalis TYPE | DB1623 (BP) | NR131835 | Hungary |
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C. uraceonemoralis | ORS-ERDO99-15-1 (BP) | KJ206520 | Hungary |
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C. uraceonemoralis TYPE | H7017739 | NR131836 | Italy |
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C. uraceus TYPE | TN04-872 (H) | NR131837 | Finland |
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C. uraceus | IK98-1607 (H) | KJ206525 | Finland |
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C. vernalisierraensis TYPE | DBB33386 (UC) | KX882652 | USA |
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C. vernalisierraensis | DBB15144 (UC) | KX882653 | USA |
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All ITS sequences were aligned and edited with BioEdit 7.0.9 (
The dataset for phylogenetic analyses contained 60 ITS sequences, representing 27 species (Table
The phylogenetic analyses recovered seven sections, including outgroup (Fig.
Pileus 2.2–6.6 cm in diam., strongly hygrophanous, translucently striate. Lamellae distant, Laccaria-like when young. Universal veil white, extremely sparse. Basidiospores 7.7–9.7 × 4.5–5.8 μm. The ITS sequences differ from the sequences of other species of section Colymbadini by at least fifteen substitutions and eight indel positions.
China. Jilin Province: Antu County, Liangjiang Town, Dongfanghong Village, broadleaf forest (Quercus mongolica dominated forest with some Juglans and Acer), 42°42'51"N, 128°01'10"E, alt. 640 m, 5 August 2017, M.L. Xie, HMJAU44449, GenBank No. (ITS) MK552380.
The name refers to the Laccaria-like lamellae when young.
Pileus 2.2–6.6 cm in diam., conical when young, then convex, strongly hygrophanous, reddish-brown (9E6–8), dark brown at the centre (8F6–8), margin to half-way translucently striate, rarely fibrillose, margin thin and wavy. Lamellae subadnate to emarginated, distant, Laccaria-like (Laccaria laccata (Scop.) Cooke) when young, reddish-brown (9E6–8) to rusty brown (6E8), edge slightly serrate. Stipe 4.2–6.6 cm long, 0.4–0.8 cm thick at apex, 0.2–0.5 cm thick at base, cylindrical to tapering towards base, dark brown (7F6) to black brown (7F3), surface with white fibrillose when young, these disappearing with age (excluding the base of stipe). Universal veil white, extremely sparse, soon disappearing. Context dark brown (7F6–8), strongly hygrophanous (pileus and stipe). Odour indistinct. Exsiccata brown (5F8) to black brown (7F5). UV fluorescence yellow on stipe, pileus and lamellar edge, strong at stipe base.
Basidiocarps of three newly-described species. a, b Cortinarius laccariphyllus (a, b HMJAU44449, holotype); c, d Cortinarius neotorvus (c HMJAU44441, holotype; d HMJAU44439); e, f Cortinarius subfuscoperonatus (e HMJAU44444, holotype; f HMJAU44445). Scale bars: 2 cm (a, b, d–f). Photographs by Meng-Le Xie.
Basidiospores 7.7–9.7 × 4.5–5.8 μm, Q = 1.43–1.84, `X = 8.0–8.5 × 4.8–5.2 μm, `Q = 1.66–1.70 (60 spores, 2 specimens), ellipsoid to amygdaloid, moderately and sharply verrucose, moderately dextrinoid. Basidia 4-spored, clavate, 23–39 × 7–9 μm, thin-walled, hyaline to olivaceous brown in 5% KOH. Lamellar edge fertile, with cylindrical-clavate sterile cells, 14–41 × 7–17 μm, thin-walled, hyaline in 5% KOH. Lamellar trama hyphae regular, pale olivaceous to olivaceous brown in 5% KOH, finely and densely encrusted. Pileipellis: epicutis hyphae cylindrical, 4–9.5 μm wide, dark olivaceous brown in 5% KOH, encrusted; hypocutis well developed, hyphae 11.5–53 μm wide, sub-cellular to cylindrical, slightly olivaceous in 5% KOH, finely encrusted. Pileus trama hyphae thin-walled, hyaline to slightly olivaceous in 5% KOH, smooth to finely encrusted. Clamp connections present.
The ITS sequences of two specimens are 534 bp long and 100% identical. They differ from the sequences of other species of section Colymbadini (
In broadleaf forest (Quercus mongolica dominated forest). Gregarious. Known from Jilin Province, China.
China. Jilin Province: Antu County, Liangjiang Town, Dongfanghong Village, broadleaf forest (Quercus mongolica dominated forest with some Juglans and Acer), 42°42'51"N, 128°01'10"E, alt. 640 m, 5 August 2017, M.L. Xie, HMJAU44450, GenBank No. (ITS) MK552381.
Cortinarius laccariphyllus has strongly hygrophanous basidiomata, Laccaria-like (when young), with distantly-spaced lamellae and an extremely sparse, white veil. Morphologically, C. nolaneiformis (Velen.) Dima, Niskanen & Liimat. is similar to C. laccariphyllus due to the strongly hygrophanous pileus, similar colouration and similar size of spores. Cortinarius uraceomajalis Dima, Liimat., Niskanen & Bojantchev is also similar to C. laccariphyllus because of the black brown stipe and the striate pileus. However, both C. nolaneiformis and C. uraceomajalis have a yellowish veil and medium-spaced lamellae and lamellae not Laccaria-like. Furthermore, C. nolaneiformis is associated with broadleaf trees and also occurs in coniferous forest; C. uraceomajalis has a somewhat lighter brown pileus as well as generally smaller (av. 7.8–8.1 × 4.6–4.7 μm) and narrower (Qav. > 1.7) spores (
Pileus 2–4.4 cm in diam., weakly hygrophanous, orange grey. Lamellae greyish-red when young. Stipe cylindrical to somewhat tapering towards base. Universal veil greyish-yellow. Context white, sometimes with violet tinge at the stipe apex. Basidiospores 8.5–10.2 × 5.8–6.9 μm. Lamellar edge sterile. The ITS sequence of the holotype differs from the sequences of other species in section Telamonia by at least six substitutions and five indels.
China. Jilin Province: Antu County, Liangjiang Town, Dongfanghong Village, broadleaf forest (Quercus mongolica dominated forest with some Juglans and Acer), 42°42'51"N, 128°01'10"E, alt. 640 m, 5 August 2017, M.L. Xie, HMJAU44441, GenBank No. (ITS) MK552384.
The name refers to Cortinarius torvus.
Pileus 2–4.4 cm in diam., hemispherical when young, then convex to almost plane with a low, broad umbo, weakly hygrophanous, orange grey (5B2), paler at the margin, surface with greyish-white fibrillose. Lamellae emarginate, medium-spaced, yellowish-grey (4B2), greyish-red (9B4–6) when young, sometimes with violet tinge when young, margin paler, slightly serrate. Stipe 4.1–10.5 cm long, 0.5–0.7 cm thick at apex, 0.3–0.5 cm thick at base, cylindrical to somewhat tapering towards base, orange grey (5B2) when moist, sometimes with violet tinge at the apex when young, surface with richly whitish fibrillose. Universal veil greyish-yellow (4B3), copious, usually forming a girdle on the upper stipe, cortina white. Context white (A1), marbled watery when moist, sometimes with violet tinge at the apex of the stipe. Odour indistinct. Exsiccata brown (6E5) to dark brown (6F6).
Basidiospores 8.5–10.2 × 5.8–6.9 μm, Q = 1.31–1.67, `X = 9.0–9.9 × 6.1–6.5 μm, `Q = 1.45–1.61 (130 spores, 6 collections), ellipsoid, moderately verrucose, moderately dextrinoid. Basidia 4-spored, cylindrical to clavate, 27–53 × 7–12 μm, thin-walled, hyaline to olivaceous brown in 5% KOH. Lamellar edge sterile, sterile cells cylindrical-clavate, 11–26 × 3–9 μm, thin-walled, hyaline in 5% KOH. Lamellar trama hyphae regular, pale olivaceous in 5% KOH, smooth. Universal veil hyphae thin-walled, hyaline to pale olivaceous yellow in 5% KOH. Pileipellis: epicutis hyphae cylindrical, 2–6 μm wide, olivaceous brown in 5% KOH, smooth; hypocutis well developed, hyphae 15–38 μm wide, sub-cellular, thin-walled, hyaline in 5% KOH, smooth. Pileus trama hyphae thin-walled, hyaline to slightly olivaceous in 5% KOH, smooth. Clamp connections present.
The ITS sequences of C. neotorvus are 513–515 bp long (5 collections, Table
In broadleaf forest (Quercus mongolica dominated forest). Solitary or gregarious. Known from Jilin and Heilongjiang Province, China.
China. Heilongjiang Province: Heihe City, Wudalianchi Scenic Area, broadleaf forest (Quercus mongolica), 48°39'15"N, 126°28'18"E, alt. 290 m, 16 August 2017, M.L. Xie, HMJAU44442, GenBank No. (ITS) MK552385; 12 August 2018, P.J. Xing, HMJAU44440; Heihe City, Shengshan National Nature Reserve, broadleaf forest (Quercus mongolica dominated forest with some Tilia and Alnus), 49°30’N, 126°43’E, alt. 300 m, 11 September 2017, G.H. Cheng, HMJAU44443, GenBank No. (ITS) MK552386. Jilin Province: Yanji City, Sandaowan Town, broadleaf forest (Quercus mongolica), 43°16'10"N, 129°07'19"E, alt. 580 m, 8 September 2018, M.L. Xie, HMJAU44437, GenBank No. (ITS) MK552382, HMJAU44438, GenBank No. (ITS) MK552383, HMJAU44439.
Cortinarius neotorvus is easily confused with C. torvus due to highly similar morphology. Morphologically, the lamellae of C. torvus are adnate to subdecurrent and distant (
Pileus 1.6–4.4 cm in diam. Context white, greyish-brown when moist. Basidiospores 9.5–12.1 × 7.9–9.7 μm. The ITS sequence of the holotype differs from other species in section Fuscoperonati by at least six substitutions and six indels.
China. Gansu Province: Zhangye City, Minle County, Gansu Qilianshan National Nature Reserve, coniferous forest (Picea crassifolia), 38°17'55"N, 100°45'54"E, alt. 2860 m, 9 August 2018, M.L. Xie, HMJAU44444, GenBank No. (ITS) MK552387.
The name refers to its affinity to Cortinarius fuscoperonatus.
Pileus 1.6–4.4 cm in diam., hemispherical when young, then low convex, weakly hygrophanous, pale greyish-brown (6C3), sometimes reddish-brown (9E5–9E6) to dark brown (6F6–6F8), surface with greyish-brown fibrillose, margin wavy with age. Lamellae emarginate, medium-spaced, reddish-brown to rusty brown (7D6–7E7), margin even when young, then slightly serrate. Stipe 2.3–7.5 cm long, 0.8–1.3 cm thick at apex, 1.5–2.5 cm thick at base, clavate, white to pale grey (E2), mycelium white at the base. Universal veil greyish-brown (6C2), rich, usually forming an annular band on the middle part and distinct belts or zones lower down. Context white (A1), greyish-brown (7F8) and marbled watery when moist, strongly hygrophanous near pileus and lamellae. Odour somewhat radish-like. Chemical reaction: pileus and context (fresh basidiomata) are dark black brown (8F3) with 10% KOH. Exsiccata brown (6E5) to dark brown (6F5).
Basidiospores 9.5–12.1 × 7.9–9.7 μm, Q = 1.10–1.45, `X = 10.3–11.2 × 8.0–8.6 μm, `Q = 1.24–1.33 (135 spores, 5 collections), subglobose to broadly ellipsoid, moderately to strongly verrucose, strongly dextrinoid. Basidia 4-spored, clavate, 35–58 × 10–13 μm, thin-walled, hyaline to olivaceous brown in 5% KOH. Lamellar edge fertile, with cylindrical-clavate sterile cells, 13–27 × 6–11 μm, thin-walled, hyaline to slightly olivaceous yellow in 5% KOH. Lamellar trama hyphae regular, pale olivaceous to olivaceous brown in 5% KOH, smooth. Pileipellis: epicutis hyphae cylindrical, 4–12 μm wide, slightly olivaceous brown to olivaceous brown in 5% KOH, some hyphae finely encrusted; hypocutis well developed, hyphae 24–89 × 15–29 μm, sub-cellular to sub-cylindrical, thin-walled, hyaline to slightly olivaceous brown in 5% KOH, smooth. Pileus trama hyphae almost thin-walled, hyaline in 5% KOH, smooth. Clamp connections present.
The ITS sequences of C. subfuscoperonatus are 524–525 bp long (5 collections, Table
In coniferous forest (Picea crassifolia dominated forest). Solitary or gregarious. Known from Gansu Province, China.
China. Gansu Province: Zhangye City, Minle County, Gansu Qilianshan National Nature Reserve, coniferous forest (Picea crassifolia), 38°17'55"N, 100°45'54"E, alt. 2860 m, 9 August 2018, M.L. Xie, HMJAU44445, GenBank No. (ITS) MK552388; Zhangye city, Su’nan Yugu Autonomous County, Gansu Qilianshan National Nature Reserve, coniferous forest (Picea crassifolia dominated forest, occasionally with Juniperus), 38°44'57"N, 99°47'56"E, alt. 3010 m, 10 August 2018, M.L. Xie, HMJAU44446, GenBank No. (ITS) MK552389, HMJAU44447, GenBank No. (ITS) MK552390; Zhangye city, Su’nan Yugu Autonomous County, Gansu Qilianshan National Nature Reserve, coniferous forest (Picea crassifolia dominated forest, occasionally with Juniperus), 38°33'13"N, 100°41'75"E, alt. 2700 m, 11 August 2018, M.L. Xie, HMJAU44448, GenBank No. (ITS) MK552391.
Cortinarius subfuscoperonatus corresponds well to the characteristics of section Fuscoperonati, with weak hygrophanous pileus, an annular band on the middle stipe and distinct belts or zones lower down, large spores (> 10 µm long) and grow in coniferous forests. Cortinarius fuscoperonatus was previously placed in section Bovini M.M. Moser (
1 | Basidiomata medium. Pileus more or less brown, strongly hygrophanous. Stipe usually cylindrical. Universal veil sparse. With positive yellow UV reaction. Associated with coniferous and/or broadleaf trees. Spores ellipsoid to amygdaloid | (section Colymbadini) 2 |
– | Basidiomata medium to large. Pileus more or less brown and hygrophanous. Stipe cylindrical to clavate. Universal veil white to greyish-yellow, sometimes with violet tinge, usually forming a ring at the middle stipe. Associated with coniferous and/or broadleaf trees | (section Telamonia) 3 |
– | Basidiomata medium to large. Pileus brown and weakly hygrophanous. Stipe clavate to slightly bulbous. Universal veil greyish-brown to blackish-brown. Associated with coniferous trees. Spores subglobose to ellipsoid, moderately to strongly verrucose | (section Fuscoperonati) 4 |
2 | Pileus strongly hygrophanous, reddish-brown to dark brown, surface translucently striate. Lamellae distant, Laccaria-like when young. Stipe cylindrical to tapering towards base, hollow. Universal veil white, extremely sparse. Odour indistinct. Positively yellow UV fluorescence (exsiccata). Associated with broadleaf trees. Spores ellipsoid to amygdaloid, on average 8.0–8.5 × 4.8–5.2 μm. | C. laccariphyllus |
– | Pileus strongly hygrophanous, yellowish-brown to brown, margin striate. Lamellae medium-spaced. Stipe cylindrical to tapering towards base, not hollow. Universal veil yellow, very sparse. Odour similar to raw vegetables. Usually yellow UV fluorescence at stipe. Associated with broadleaf trees. Spores amygdaloid to narrowly amygdaloid, on average 7.8–8.1 × 4.6–4.7 μm | C. uraceomajalis |
– | Pileus strongly hygrophanous, dark greyish-brown to dark brown, margin slightly striate. Lamellae medium-spaced to fairly distant, margin whitish when young. Stipe cylindrical to clavate, sometimes tapering downwards, sometimes hollow. Universal veil yellow. Strong yellow UV fluorescence at stipe, dull yellowish-brown at pileus, lamellae and context. Associated with coniferous and broadleaf trees. Spores amygdaloid to weakly ellipsoid, on average 8.1–8.6 × 4.8–5.1 μm | C. nolaneiformis |
3 | Pileus pale greyish-yellow, paler at the margin, weakly hygrophanous. Lamellae emarginate, medium-spaced, greyish-red when young, sometimes with violet tinge. Stipe cylindrical to somewhat tapering towards base, pale greyish-yellow. Odour indistinct. Spores ellipsoid, on average 9.0–9.9 × 6.1–6.5 μm | C. neotorvus |
– | Pileus greyish-brown to chestnut brown, usually with violet tinge at the margin, weakly hygrophanous. Lamellae adnate, subdecurrent to distant, greyish-brown, with violet tinge. Stipe clavate, usually bulb at the base. Odour acidulous. Spores ellipsoid, 8–10.5 × 6–7 μm | C. torvus |
4 | Pileus pale greyish-brown, sometimes reddish-brown to dark brown, margin wavy with age, with greyish-brown fibrillose. Stipe clavate. Spores subglobose to broadly ellipsoid, on average 10.3–11.2 × 8.0–8.6 μm | C. subfuscoperonatus |
– | Pileus chocolate brown to blackish-brown, pale greyish-brown at the edge, fine fibrous to fine scaly. Stipe clavate, with a bulb at the base. Spores ellipsoid to broadly ellipsoid, 9.7–11.6 × 6.6–7.7 μm | C. fuscoperonatus |
Cortinarius is the most species-rich genus of Agaricales, with most of the described species distributed in the Northern Hemisphere. However, so far, little has been done on Cortinarius taxonomy in north-eastern Asia or even in the whole of Asia, leaving an important gap in our knowledge of this genus (
In this study, we described the phylogenetic relationships amongst the three new species and other species, based on the ITS sequences. However, multiple genes should be used in future studies to describe more complex phylogenetic relationships in Cortinarius, which some mycologists have conducted. Peintner et al. (2002) assessed the phylogenetic relationships of Rozites, Cuphocybe and Rapacea by molecular phylogenetic approaches, based on ITS and LSU. Frøslev et al. (2005) analysed the phylogeny of Cortinarius subgenus Phlegmacium, a taxonomically difficult group, based on ITS, RPB1 and RPB2. They speculated that the sequences from RNA polymerase II genes have the potential for resolving the phylogenetic problems of Cortinarius. Later, the study of Frøslev et al. (2007) showed that the delimitation of species, based on ITS sequences, is more consistent with a conservative morphological species concept and there is considerable potential for using ITS sequence data as a barcode for section Calochroi.
The study was supported by China Agriculture Research System (No. CARS20), Special Fund for Agro-scientific Research in the Public Interest (No. 201503137), Overseas Expertise Introduction Project for Discipline Innovation (111 Center) (No. D17014), National Natural Science Foundation of China (No. 31270072), the Special Funds for the Young Scholars of Taxonomy of the Chinese Academy of Sciences (No. ZSBR-001). We would like to express our gratitude to the people in the Engineering Research Center of Edible and Medicinal Fungi, Ministry of Education, Jilin Agricultural University, including Dr. Bo Zhang for her suggestions to improve our work, Ms. Yang Yang and Ms. Yu-Xiu Guo for their help during molecular experiments, Mr. Yang Wang, Mr. Zhu-Shan Liu and Mr. Zhi-Hui Luo for their help during the field trips in Jilin and Heilongjiang, as well as Prof. Sheng-Long Wei and Ms. Qian-Qian Liang (Gansu Engineering Laboratory of Applied Mycology, Hexi University, China) for their kind help during the field trips in Gansu. We thank Dr. Frederick Leo Sossah for his kind-hearted and excellent technical assistance with the English language correction. We also thank the reviewers, Bálint Dima, Joseph F. Ammirati and Jerry Adrian Cooper, for their suggestions and corrections to improve our work.