中国丝膜菌属李玉丝膜菌组一新种:李玉丝膜菌
Cortinarius liyui sp. nov., a new species of Cortinarius sect. Liyuorum sect. nov. from China
收稿日期: 2022-10-19 接受日期: 2022-10-31
Corresponding authors:
Received: 2022-10-19 Accepted: 2022-10-31
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作者简介 About authors
ORCID:XIEMengle(0000-0002-7798-7048) 。
魏铁铮,中国科学院微生物研究所真菌学国家重点实验室副研究员长期从事大型经济真菌的系统学、生物多样性及保护的研究 , E-mail:weitiezheng@163.com
本研究报道了丝膜菌属的一个新种:李玉丝膜菌,该种采自中国吉林省蒙古栎林中,其主要特征为担子体小型、幼时具蓝紫色调,菌盖强水浸状、具条纹,担孢子近球形。该种在系统发育树中形成一个独立的分支,与斑丝膜菌组形成姐妹关系,但遗传关系较远,属于新组李玉丝膜菌组。本文对新分类单元进行了详细描述,并对其系统发育上相近的物种进行了比较。
关键词:
A telamonioid species new to science, Cortinarius liyui, found from Jilin Province, Northeast China is presented based on morphological characteristics and molecular phylogenetic analysis. The species is characterized by its small and violet tinged basidiomata, hygrophanous and striate pileus, and subglobose to broadly ellipsoid basidiospores. The phylogeny inferred from the ITS and 28S sequences demonstrates that C. liyui belongs to Cortinarius sect. Liyuorum, a section new to science and probably related to sect. Spilomei. Detailed descriptions of the new taxon and a comparison with its phylogenetically related taxa are provided.
Keywords:
本文引用格式
谢孟乐, 李熠, 王科, 冀瑞卿, 魏铁铮.
XIE Mengle, LI Yi, WANG Ke, JI Ruiqing, WEI Tiezheng.
INTRODUCTION
Cortinarius (Pers.) Gray is an important ectomycorrhizal genus with agaricoid to sequestrate basidiomata, rust brown basidiospore deposit and verrucose basidiospores (Peintner et al. 2001; Frøslev et al. 2006; Bhunjun et al. 2022). Many Cortinarius species have been used as indicator species for valuable natural environments in Europe due to their narrow ecological preferences and sensitivity to environmental change (Vesterholt 1991; Hallingbäck & Aronsson 1998). Many Cortinarius species are edible mushrooms, for example C. armillatus (Fr.) Fr., C. emodensis Berk. and C. sinensis L.H. Sun, T.Z. Wei & Y.J. Yao (Dai et al. 2010; Li et al. 2015; Xie 2018). Some species, for example C. collinitus (Pers.: Fr.) Fr. and C. violaceus (L.) Gray, with potential value of anti-tumor and antioxidant (Dai & Yang 2008; Dai et al. 2009; Li & Bao 2020). In addition, several species are notoriously poisonous, e.g. C. orellanus Fr. and C. speciosissimus Kühner & Romagn (Bau et al. 2014; Wu et al. 2019).
The latest phylogenomic study by Liimatainen et al. (2022) showed that the genus Cortinarius s.l. could be divided into ten genera. The recent research based on the ITS region in GenBank showed the species of Cortinarius s.s. maybe over 5 000 in the world, including more than one thousand undescribed species (Bhunjun et al. 2022). In China, 309 Cortinarius s.l. taxa have been reported, including nineteen taxa that originally described from China (Zang 1987; Yang 1992; Wei & Yao 2013; Xie 2018; Xie et al. 2019, 2020, 2021a, 2021b, 2022; Yuan et al. 2020; Luo & Bau 2021; Wang et al. 2022).
In this study, samples of one telamonioid species, C. liyui, were collected and described as a new species from Jilin Province, China. The morphological characters and phylogenetic analysis supported the recognition of this species within Cortinarius and confirmed this species belongs to a novel section Liyuorum.
1 MATERIALS AND METHODS
1.1 Specimens and morphological description
Specimens were collected in Yanji, Jilin Province. Fresh fruit bodies were photographed in the field, and dried in an oven at 50 °C. Specimens have been deposited in the Herbarium of Mycology, Jilin Agricultural University (HMJAU), Changchun, China. The macroscopic characteristics were recorded from the fresh basidiomata. Microscopic characteristics were examined and described in 5% KOH, Congo Red or Melzer’s reagent under a Zeiss AX10 light microscope. The length/width ratios (Q) were calculated for individual basidiospores. Xav. and Qav. refer to the average value of basidiospores of per specimen. The ornamentation of basidiospores was studied using a Hitachi, model SU8010, field emission scanning electron microscope (FESEM) in Jilin Agricultural University.
1.2 DNA extraction, PCR and sequencing
The DNA was extracted from dried specimens, using a NuClean PlantGen DNA Kit (CWBIO). Primers ITS1F and ITS4 were used to amplify internal transcribed spacer (ITS) region (White et al. 1990; Gardes & Bruns 1993). The nuclear large subunit rDNA (28S) region was amplified with primers LR0R and LR7 (Vilgalys & Hester 1990). The PCR amplification progress followed Xie et al. (2022) and amplicons were sequenced by Sangon Biotech (Shanghai) Co. Ltd.
1.3 Phylogenetic analysis
The taxon sampling strategy for the selection of sequences for phylogenetic trees was to choose related taxa based on a BLASTn search in GenBank within Cortinarius (Table 1). Two partition datasets (ITS, 28S) were separately aligned and manually adjusted with BioEdit 7.1.3.0 (Hall 1999). Phyutility 2.2 was used to concatenate the aligned datasets (Smith & Dunn 2008). Section Orellani M.M. Moser was selected as an outgroup for phylogenetic analysis due to sect. Orellani separated from other lineages (Soop et al. 2019).
Table 1 Sequences used in the phylogenetic analysis
Species | Voucher number | Origin | ITS | 28S |
---|---|---|---|---|
C. anomalus | CFP1154 (TYPUS) | Sweden | KX302224 | - |
C. bolaris | TUB 0118524 | Germany | AY669596 | AY669596 |
C. bolaris | 3861 | Canada | KJ705110 | - |
C. bolaris | CFP1008 (TYPUS) | Sweden | KX302233 | - |
C. camphoratus | EH23 | Canada | FJ717505 | FJ717505 |
C. caninus | HMJAU44372 | China | OP620657 | OP620671 |
C. caninus | CFP627 (TYPUS) | Sweden | KX302250 | - |
C. cotoneus | 19XML11153 | China | OP620655 | OP620666 |
C. cruentoides | PDD101864 (TYPUS) | New Zealand | KJ635217 | KJ635217 |
C. dysodes | PDD70499 (TYPUS) | New Zealand | GU233340 | GU233394 |
C. eartoxicus | MEL2351137 | Australia | KP311432 | KP311376 |
C. epsomiensis | KM74963 (TYPUS) | United Kingdom | MK010952 | - |
C. ferrusinus | JB-8106/13 | Spain | KY657254 | - |
C. illibatus | HMJAU48760 | China | MW911735 | OP620668 |
C. indotatus | PDD88257 | New Zealand | KJ421110 | KJ421110 |
C. liyui | HMJAU58936 | Jilin, China | OP620658 | - |
C. liyui | HMJAU58937 | Jilin, China | OP620659 | - |
C. liyui | ECM86 | Zhejiang, China | JQ991711 | - |
C. liyui | HMJAU58939 (TYPUS) | Jilin, China | OP620660 | OP620672 |
C. liyui | HMJAU58938 | Jilin, China | OP620661 | - |
C. liyui | HMJAU58935 | Jilin, China | OP620662 | - |
C. orellanoides | TUB011828 | Germany | AY669595 | AY669595 |
C. orellanus | IB19980580 | Austria | AF389164 | AF388773 |
C. pseudocamphoratus | HMJAU48698 (holotype) | China | OM001483 | OM001524 |
C. sommerfelti | HMJAU44457 | China | OP620652 | OP620663 |
C. spilomeus | CFP1137 (TYPUS) | Sweden | KX302267 | - |
C. subsalor | HMJAU48759 (TYPUS) | China | MW911734 | OP620670 |
C. subsanguineus | HMJAU48961 | China | OP620653 | OP620664 |
C. tabularis | CFP949 (TYPUS) | Sweden | KX302275 | - |
C. tasmacamphoratus | HO-A20606A0 | Tasmania | AY669633 | AY669633 |
C. tibeticisalor | HMJAU48764 (TYPUS) | China | MW911729 | OP620669 |
C. uliginosus | KH7 | Norway | KC842412 | KC842482 |
C. veronicae | PDD68468 (TYPUS) | New Zealand | KC017355 | - |
C. xiaojinensis | HMJAU58895 | China | OP620654 | OP620665 |
C. zakii | UBC-F19609 | Canada | HQ604650 | HQ604650 |
C. sp. | SWUBC741 | Canada | DQ481671 | - |
C. sp. | TU105220 | Sweden | UDB015906 | - |
C. sp. | T21468 | China | OP620656 | OP620667 |
C. sp. | MEL2089705 | Australia | GQ890326 | JX544951 |
Newly generated sequences are in bold. - Indicate no data.
For phylogenetic analysis, Bayesian inference (BI) and maximum likelihood (ML) methods were used. For BI analysis, the best-fit models for ITS and 28S partitions were estimated using the Akaike information criterion (AIC), implemented in MrModeltest 2.3 (Nylander 2004). The BI analysis was performed with MrBayes 3.2.6 (Ronquist & Huelsenbeck 2003). Four Markov chains were run for 2 runs from random starting trees for 500 000 generations, sampling every 100th generation. The first 25% of trees were discarded to build the 50% majority rule consensus tree. RAxML 8.2.12, implemented in raxmlGUI, was used for ML analysis, with a rapid bootstrapping algorithm of 1 000 replicates (Silvestro & Michalak 2012; Stamatakis 2014). All default parameters with the GTRGAMMA model were used in the ML analysis.
2 RESULTS
2.1 Phylogenetic analysis
The phylogenetic analysis includes 39 specimens, of which 13 are type specimens. In total, 21 sequences were newly generated. The best-fit models for BI analysis of both ITS and 28S were GTR+I+G. The ML tree is selected to represent the phylogeny (Fig. 1). The ML bootstrap values (ML) ≥60% and Bayesian posterior probabilities (BPP)≥0.90 are shown on the branches (ML/BPP).
Fig. 1
Fig. 1
ML phylogram inferred from the ITS+28S dataset. The tree is rooted with sect. Orellani. The ML bootstrap values (ML)≥60% and Bayesian posterior probabilities (BPP)≥0.90 are shown on the branches (ML/BPP). New species are marked in black bold font.
The phylogenetic analysis recovered 12 sections of Cortinarius, including the outgroup. Every section formed a separately monophyletic lineage with strong statistical support. The specimens of Cortinarius liyui clustered in sect. Liyuorum (100%/1). Section Liyuorum was separated into single lineage and formed a sister relationship (65%/0.96) with sect. Spilomei (Moënne-Locc. & Reumaux) Consiglio, D. Antonini & M. Antonini (93%/1).
2.2 Taxonomy
Cortinarius sect. Liyuorum M.L. Xie, R.Q. Ji & T.Z. Wei sect. nov.
Fungal Name: FN571217
Diagnosis. Basidiomata small. Pileus brown with violet tinge, hygrophanous. Lamellae grayish violet to rusty brown. Stipe slender, cylindrical or slightly clavate, with violet tinge, then brown. Universal veil white and spares. Basidiospores rather small, subglobose to broadly ellipsoid, rarely ellipsoid. Pileipellis with an epicutis and a developed hypodermium.
Type species. Cortinarius liyui M.L. Xie, R.Q. Ji & T.Z. Wei
Etymology. Named after the type species of the section.
Cortinarius liyui M.L. Xie, R.Q. Ji & T.Z. Wei sp. nov.Fig. 2
Fungal Name: FN571211
Fig. 2
Fig. 2
Cortinarius liyui. A: Basidiomata (HMJAU58937). B: Basidiomata (HMJAU58938). C: Basidiomata (HMJAU58939, holotype). D: Basidiospores under FESEM (HMJAU58939). E: Basidiospores (HMJAU58939). F: Pileipellis including epicuits (upper portion) and hypodermium (lower portion) (HMJAU58939).
Diagnosis. Similar to Cortinarius subg. Telamonia species, but differs in its small and slender basidiomata, violet-tinged pileus and stipe when young, together with the small and subglobose to broadly ellipsoid basidiospores.
Holotype. Sandaowan, Yanji, Jilin Province, CHINA, on ground in Quercus mongolica forests, alt. 620 m, 23 Sept. 2020, M.L. Xie, 20XML13068 (HMJAU58939).
Etymology. liyui, in honor of Chinese mycologist Prof. Yu Li, and celebrating his 80th birthday.
Pileus 18-40 mm in diam., plano-convex, somewhat depressed at the center, strongly hygrophanous; violet brown at first, then reddish brown to grayish brown, paler at the margin, translucently striate, especially at the margin, somewhat wrinkled when mature. Lamellae up to 5 mm wide, emarginated, medium-spaced, grayish violet when young, then yellowish brown to rusty brown, edge even. Stipe 30-63×3-4 mm, cylindrical or somewhat thickened at the base, surface fibrillose, lightly purple at first, later grayish brown to dark brown, basal mycelium white, hollow when mature. Context violet tinge at first, then reddish brown. Odor radish. Universal veil white, sparse. Basidiospore deposit rust brown.
Basidiospores (150/5/5) 6.4-7.7(8.5)×5.5-6.9 (7.5) μm, Q=1.04-1.25(1.33), Xav.=6.9-7.3×6.0- 6.2 μm, Qav.=1.16-1.2, subglobose to broadly ellipsoid, moderately verrucose, moderately to strongly dextrinoid. Basidia clavate, 4-spored, colorless or lightly olivaceous brown to olivaceous brown. Lamellar trama hyphae smooth, up to 25 μm wide, lightly olivaceous brown. Lamellar edge fertile, with small clavate sterile cells. Pileipellis duplex: epicuits hyphae 2.5-8 μm wide, lightly olivaceous brown to olivaceous brown, smooth; hypodermium developed, hyphae lightly olivaceous brown, 7-23 μm wide, smooth. Clamp connections present.
Ecology and distribution. Solitary or gregarious on ground in Quercus mongolica forests. Known from Jilin Province, also from Zhejiang Province based on the ectomycorrhizal sequence (JQ991711).
Additional specimens examined. Sandaowan, Yanji, Jilin Province, on ground in Quercus mongolica forests, alt. 620 m, 23 Sept. 2020, M.L. Xie, 20XML13047 (HMJAU58935), 20XML13055 (HMJAU58936), 20XML13056 (HMJAU58937), 20XML13062 (HMJAU58938).
3 DISCUSSIONS
Cortinarius sect. Liyuorum forms a monophyletic clade in the phylogenetic tree, and is a sister to sect. Spilomei, but without robust support. The pileus of sect. Spilomei species is usually weakly hygrophanous and non-striate, and its universal veil is yellowish to reddish and usually forming characteristic rusty red squamules on the stipe (Dima et al. 2016). Our new section might belong to the subgenus Camphorati Liimat., Niskanen & Ammirati (Liimatainen et al. 2022), but the morphological features of the new species differ from those of the subgenus. At present, we are not sure which subgenus the new section belongs to. Further research is needed in the future.
Cortinarius liyui is characterized by its small and slender basidiomata with violet tinge when juvenile, additionally by its strongly hygrophanous pileus, medium-spaced lamellae, white and sparse universal veil, and small and subglobose to broadly ellipsoid basidiospores. This is a typical telamonioid species for its strongly hygrophanous basidiomata and can be easily distinguished from other telamonioid species by the violet tinge basidiomata and subglobose basidiospores. The ITS sequences of Jilin specimens are identical. In phylogenesis, a sequence (JQ991711) from the ectomycorrhizal sample from Zhejiang Province, subtropical China, clustered with the Jilin specimens, but had one bp substitution, here we treated it as C. liyui and considered this species had a distribution in subtropical China as well.
ACKNOWLEDGEMENTS
The authors are grateful to Mr. CAO Zhenyuan for his kind help in the fieldwork.
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