Research Article |
Corresponding author: Chang-Lin Zhao ( fungichanglinz@163.com ) Academic editor: Bao-Kai Cui
© 2024 Kai-Yue Luo, Jiang-Qing Su, Chang-Lin Zhao.
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:
Luo K-Y, Su J-Q, Zhao C-L (2024) Morphological and molecular identification for four new wood-inhabiting species of Trechispora (Basidiomycota) from China. MycoKeys 105: 155-178. https://doi.org/10.3897/mycokeys.105.120438
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Four new wood-inhabiting fungi, Trechispora albofarinosa, T. bisterigmata, T. pileata and T. wenshanensis spp. nov., are proposed based on a combination of morphological features and molecular evidence. Trechispora albofarinosa is characterized by the farinose basidiomata with flocculence hymenial surface, a monomitic hyphal system with clamped generative hyphae, and ellipsoid, warted basidiospores. Trechispora bisterigmata is characterized by the membranous basidiomata with odontioid hymenial surface, rhizomorphic sterile margin, barrelled basidia and subglobose to broad ellipsoid, smooth basidiospores. Trechispora pileata is characterized by the laterally contracted base, solitary or imbricate basidiomata, fan shaped pileus, radially striate-covered surface with appressed scales, odontioid hymenophore surface, and subglobose to broad ellipsoid, thin-walled, smooth basidiospores. Trechispora wenshanensis is characterized by a cottony basidiomata with a smooth hymenial surface, and ellipsoid, thin-walled, warted basidiospores. Sequences of ITS and LSU marker of the studied samples were generated, and phylogenetic analyses were performed with the maximum likelihood, maximum parsimony, and Bayesian inference methods. The phylogenetic tree inferred from the ITS+nLSU sequences highlighted that four new species were grouped into the genus Trechispora.
East Asia, macrofungi, molecular systematics, taxonomy, 4 new taxa
Fungi represent one of the most diverse groups of organisms on earth, with an indispensable role in the processes and functioning of ecosystems (
Trechispora P. Karst. (Hydnodontaceae) typified by T. onusta P. Karst., which is characterized by resupinate to effused basidiomata; a smooth to hydnoid to poroid hymenophore; ampullaceous septa; short cylindric basidia; and smooth to verrucose or aculeate basidiospores (
There have been many studies on the phylogeny of this genus in recent years. A high phylogenetic diversity on the corticioid Agaricomycetes based on two genes, 5.8S and 28S showed that nine taxa of Trechispora nested into trechisporoid clade (
During investigations into the wood-inhabiting fungi in the Yunnan-Guizhou Plateau of China, samples representing four additional species belonging to genus Trechispora were collected. To clarify the placement and relationships of the four species, we carried out a phylogenetic and taxonomic study on Trechispora, based on the ITS+nLSU.
The specimens studied were deposited at the herbarium of Southwest Forestry University (SWFC), Kunming, Yunnan Province, China. The macromorphological descriptions were based on field notes and photos captured in the field and laboratory. Color, texture, taste and odor of basidiomata were mostly based on authors’ field trips. Color terminology followed
The CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd, Beijing) was used to obtain genomic DNA from the dried specimens following the manufacturer’s instructions (
List of species, specimens and GenBank accession numbers of sequences used in this study.
Species name | Specimen No. | GenBank accession No. | References | |
---|---|---|---|---|
ITS | LSU | |||
Fibrodontia alba | TNM F24944 | NR153983 | NG060401 |
|
F. brevidens | Wu 9807-16 | KC928276 | KC928277 |
|
Trechispora alba | CH21384 | OR557258 | – |
|
T. albofarinosa | CLZhao 4356 | OQ241383 | OQ282703 | This study |
T. amianthina | CBS 202.54 | – | MH868822 |
|
T. araneosa | KHL 8570 | AF347084 | – |
|
T. bambusicola | CLZhao 3302 | MW544021 | MW520171 |
|
T. bambusicola | CLZhao 3305 | MW544022 | MW520172 |
|
T. bispora | CBS 142.63 | MH858241 | MH869842 |
|
T. bisterigmata | CLZhao 2522 | OQ241386 | – | This study |
T. bisterigmata | CLZhao 7870 | OQ241387 | – | This study |
T. byssinella | UC 2023068 | KP814481 | – | Unpublished |
T. chartacea | FLOR56185 | MK458775 | – |
|
T. clancularis | FRDBI 4426619 | MW487976 | – | Unpublished |
T. cohaerens | HHB-19445 | MW740327 | – | Unpublished |
T. copiosa | AMO427 | MN701015 | MN687973 |
|
T. copiosa | AMO450 | MN701017 | MN687974 |
|
T. crystallina | LWZ 20170729-2 | OM523419 | OM339238 |
|
T. cyatheae | FR0219443 | UDB024016 | UDB024017 |
|
T. cyatheae | FR0219446 | UDB024020 | UDB024021 |
|
T. dentata | Dai 22565 | OK298491 | OM049408 |
|
T. dimitiella | Dai 21181 | OK298493 | OK298949 |
|
T. dimitiella | Dai 21931 | OK298492 | OK298948 |
|
T. echinospora | E11/37-10 | JX392850 | JX392851 |
|
T. echinospora | E11/37-12 | JX392853 | JX392854 |
|
T. farinacea | 356 | AF347089 | – |
|
T. farinacea | MA-Fungi 79474 | JX392855 | JX392856 |
|
T. fimbriata | CLZhao 7969 | MW544024 | MW520174 |
|
T. fimbriata | CLZhao 9006 | MW544025 | MW520175 |
|
T. foetida | FLOR 56315 | MK458769 | – |
|
T. fragilis | Dai 20535 | OK298494 | OK298950 |
|
T. gelatinosa | AMO824 | MN701020 | MN687977 |
|
T. gelatinosa | AMO1139 | MN701021 | MN687978 |
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T. gracilis | LWZ 20170814-17 | OM523435 | OM339253 |
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T. havencampii | DED8300 | NR154418 | NG059993 |
|
T. hondurensis | HONDURAS19-F016 | NR178152 | NG081479 |
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T. hondurensis | HONDURAS19-F016a | MT571523 | MT636540 |
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T. hymenocystis | KHL 8795 | AF347090 | – | Unpublished |
T. hymenocystis | KHL 16444 | MT816397 | – | Unpublished |
T. incisa | GB0090521 | KU747093 | KU747086 | Unpublished |
T. incisa | GB0090648 | KU747095 | KU747087 | Unpublished |
T. invisitata | 5425_537 | ON963772 | – | Unpublished |
T. invisitata | UC2023088 | KP814425 | – | Unpublished |
T. kavinioides | KGN 981002 | AF347086 | – | Unpublished |
T. laevispora | Dai 21655 | OK298495 | OM108710 |
|
T. larssonii | LWZ 20190817-11a | OM523442 | OM339259 |
|
T. longiramosa | HG 140168 | OM523448 | OM339264 |
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T. mellina | URM85756 | – | MH280000 | Unpublished |
T. microspora | FRDBI 18772216 | OL828778 | – | Unpublished |
T. mollis | URM85884 | MK514945 | MK514945 | Unpublished |
T. mollis | URM85885 | – | MT423667 | Unpublished |
T. mollusca | iNAT 30809943 | MZ269232 | – | Unpublished |
T. mollusca | CFMR:DLL2011-186 | KJ140681 | – | Unpublished |
T. nivea | MA-Fungi 76238 | JX392824 | JX392825 |
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T. nivea | MA-Fungi 76257 | JX392826 | JX392827 |
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T. pallescens | FLOR56184 | MK458767 | – | Unpublished |
T. pallescens | FLOR56188 | MK458774 | – | Unpublished |
T. papillosa | AMO713 | MN701022 | MN687979 |
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T. papillosa | AMO795 | MN701023 | MN687981 |
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T. patawaensis | VPapp-GF1901 | OL314550 | OL314546 | Unpublished |
T. perminispora | LWZ2019081639a | OM523525 | OM339329 |
|
T. pileata | CLZhao 4456 | OQ241388 | OQ282715 | This study |
T. praefocata | FRDBI 18819116 | OL828784 | – | Unpublished |
T. regularis | KHL 10881 | AF347087 | – | Unpublished |
T. rigida | URM85754 | MT406381 | MH279999 | Unpublished |
T. sinensis | LWZ 20170816-35 | OM523479 | OM339287 |
|
T. stellulata | 14153 | MW023104 | – | Unpublished |
T. stellulata | 33962903 | ON364078 | – | Unpublished |
T. stellulata | UC2023099 | KP814451 | – | Unpublished |
T. stellulata | UC2023230 | KP814491 | – | Unpublished |
T. stevensonii | MA-Fungi 70669 | JX392841 | JX392842 |
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T. stevensonii | MA-Fungi 70645 | JX392843 | JX392844 |
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T. subfarinacea | LWZ2020092133a | OM523528 | OM339331 |
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T. subhelvetica | 7089 | JN710601 | – | Unpublished |
T. subhymenocystis | LWZ 20190818-29b | OM523492 | OM339299 |
|
T. subregularis | VPapp-GF2103 | OL331097 | OL314548 | Unpublished |
T. subsinensis | LWZ 20190611-9 | OM523497 | OM339304 |
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T. subsphaerospora | KHL 8511 | AF347080 | – | Unpublished |
T. termitophila | AMO396 | MN701025 | MN687983 |
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T. termitophila | AMO893 | MN701026 | MN687984 |
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T. torrendii | URM85886 | MK515148 | MH280004 | Unpublished |
T. tropica | LWZ 20170613-16 | OM523503 | OM339311 |
|
T. tuberculata | Dai17433 | OM523507 | OM339314 |
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T. wenshanensis | CLZhao 11649 | OQ241389 | OQ282716 | This study |
T. wenshanensis | CLZhao 11715 | PP712100 | – | This study |
T. wenshanensis | CLZhao 22940 | PP712101 | – | This study |
T. yunnanensis | CLZhao 210 | NR177488 | MN654918 |
|
T. yunnanensis | CLZhao 214 | MN654922 | MN654919 |
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The sequences were aligned in MAFFT version 7 (
Maximum parsimony analysis in PAUP* version 4.0a169 (http://phylosolutions.com/paup-test/) was applied to ITS+nLSU following a previous study (
MrModeltest 2.3 (
The ITS+nLSU dataset comprised sequences from 88 fungal specimens representing 64 taxa. The dataset had an aligned length of 2271 characters, of which 1376 characters were constant, 190 were variable and parsimony-uninformative and 705 were parsimony-informative. Maximum parsimony analysis yielded 300 equally parsimonious tree (TL = 5543, CI = 0.2979, HI = 0.7021, RI = 0.5278 and RC = 0.1572). The best model of nucleotide evolution for the ITS+nLSU dataset estimated and applied in the Bayesian analysis was found to be GTR+I+G. Bayesian analysis and ML analysis resulted in a similar topology as in the MP analysis. The Bayesian analysis had an average standard deviation of split frequencies = 0.012925 (BI) and the effective sample size (ESS) across the two runs is double the average ESS (avg. ESS) = 389. The phylogenetic tree inferred from the ITS+nLSU sequences highlighted that four new species were grouped into the genus Trechispora (Fig.
Maximum parsimony strict consensus tree illustrating the phylogeny of the four new species and related species in Trechispora, based on ITS+nLSU sequences. Branches are labelled with maximum likelihood bootstrap values > 70%, parsimony bootstrap values > 50% and Bayesian posterior probabilities > 0.95, respectively.
China. Yunnan Province, Pu’er, Jingdong County, Huangcaoling, Wuliangshan National Nature Reserve, 24°23′N, 100°45′E, altitude 2350 m a.s.l., on the fallen branch of Pinus, leg. C.L. Zhao, 5 October 2017, CLZhao 4356 (SWFC).
Albofarinosa (Lat.): referring to the farinose basidiomata with white hymenial surface.
Basidiomata annual, resupinate, farinose, without odor or taste when fresh, up to 3.5 cm long, 1.5 cm wide, and 300–500 µm thick. Hymenial surface flocculence, white when fresh, white to cream on drying. Sterile margin indistinct, white, and up to 0.5 mm wide.
Hyphal system monomitic, generative hyphae with clamp connections with ampullaceous septa, colorless, thick-walled, frequently branched, interwoven, 2–3.5 µm in diameter; IKI–, CB–, tissues unchanged in KOH.
Cystidia and cystidioles absent; basidia clavate, with four sterigmata and a basal clamp connection, 6.5–10 × 3.5–5 µm.
Basidiospores ellipsoid, colorless, thin-walled, aculeate, IKI–, CB–, 2.5–3.5 (–4) × 2–2.5 (–3.5) μm, L = 3.18 µm, W = 2.44 µm, Q = 1.3 (n = 30/1).
China. Yunnan Province, Yuxi, Xinping County, Mopanshan National Forestry Park, 23°56′N, 101°29′E, altitude 2200 m a.s.l., on the trunk of Albizia julibrissin, leg. C.L. Zhao, 20 Aguest 2017, CLZhao 2522 (SWFC).
Bisterigmata (Lat.): referring to the basidia mainly with two sterigmata.
Basidiomata annual, resupinate, adnate, membranous, without odor or taste when fresh, up to 2.5 cm long, 1.5 cm wide, and 4 mm thick. Hymenial surface odontioid, cream. Sterile margin indistinct, white, rhizomorphic, up to 0.5 mm wide.
Hyphal system monomitic, generative hyphae with clamp connections, colorless, slightly thick-walled, ampullate septa frequently present in subiculum and hymenium with crystals, up to 6 µm wide, branched, interwoven, 2.5–4 µm in diameter; IKI–, CB–, tissues unchanged in KOH.
Cystidia and cystidioles are absent; basidia barrelled, slightly constricted, with two or four sterigmata and a basal clamp connection, 6.5–14.5 × 3.5–5.5 µm.
Basidiospores subglobose to broad ellipsoid, colorless, slightly thick-walled, smooth, IKI–, CB–, (2–) 2.5–4 × 2–3.5 µm, L = 3.03 µm, W = 2.41 µm, Q = 1.23–1.28 (n = 60/2).
(paratype). China. Yunnan Province, Yuxi, Xinping County, Mopanshan National Forestry Park, 23°56′N, 101°29′E, altitude 2200 m a.s.l., on the living angiosperm tree, leg. C.L. Zhao, 19 August 2018, CLZhao 7870 (SWFC).
China. Yunnan Province, Pu’er, Jingdong County, Wuliangshan National Nature Reserve, 24°23′N, 100°45′E, altitude 2350 m a.s.l., on the angiosperm trunk, leg. C.L. Zhao, 6 October 2017, CLZhao 4456 (SWFC).
Pileata (Lat.): referring to the pileate basidiomata.
Basidiomata annual, with a laterally contracted base, solitary or imbricate. Pileus fan shaped, cortical to corky, up to 1.5 cm long, 1 cm wide, and 2 mm thick, yellowish to yellowish brown, the surface radially striate covered with appressed scales, azonate; the hymenophore surface odontioid, yellowish brown, up to 1 mm long. Context cream, 1 mm thick. Sterile margin indistinct, slightly buff, and 0.5 mm wide.
Hyphal system monomitic, generative hyphae with clamp connections, colorless, thick-walled, frequently branched, interwoven, hyphae in spines 2.5–4 µm in diameter, IKI–, CB–, tissues unchanged in KOH. Hyphae in context colorless, thin- to thick-walled, unbranched, interwoven, 4.5–6 µm in diameter, IKI–, CB–, tissues unchanged in KOH.
Cystidia and cystidioles absent; basidia subcylindrical, constricted, with four sterigmata and a basal clamp connection, 5–7 × 2.5–4 µm.
Basidiospores subglobose to broad ellipsoid, colorless, thin-walled, smooth, IKI–, CB–, (2.5–) 2.8–5 (–5.5) × (2.5–) 3–4.7 µm, L = 4 µm, W = 3.56 µm, Q = 1.12 (n = 30/1).
China. Yunnan Province, Wenshan, Babao Town, Balao battle site, 23°22′N, 104°15′E, altitude 1300 m a.s.l., on the fallen angiosperm branch, leg. C.L. Zhao, 19 January 2019, CLZhao 11649 (SWFC).
Wenshanensis (Lat.): referring to the locality (Wenshan) of the type specimen.
Basidiomata annual, resupinate, adnate, cottony, easily to separate from substrate, without odor or taste when fresh, up to 5.5 cm long, 4 cm wide, and 200–400 µm thick. Hymenial surface smooth, slightly cream when fresh, cream to buff on drying. Sterile margin indistinct, cream, and 1–2 mm wide.
Hyphal system monomitic, generative hyphae with clamp connections, colorless, thin- to thick-walled, branched, interwoven, 1–2 µm in diameter; IKI–, CB–, tissues unchanged in KOH.
Cystidia and cystidioles are absent; basidia barrelled, with four sterigmata and a basal clamp connection, 7–10 × 3–5 μm.
Basidiospores ellipsoid, colorless, thin-walled, warted, IKI–, CB–, (2–) 2.5–3.7 (–4) × (1.5–) 2–3 µm, L = 3.02 µm, W = 2.37 µm, Q = 1.25–1.30 (n = 90/3).
(paratypes). China. Yunnan Province, Wenshan, Funing county, Guying village, 23°42′N, 105°53′E, altitude 1000 m a.s.l., on the fallen angiosperm branch, leg. C.L. Zhao, 20 January 2019, CLZhao 11715; Yunnan Province, Lincang, Lancangjiang Forestry Region, 25°37′N, 97°30′E, altitude 1750 m a.s.l., on the fallen angiosperm branch, leg. C.L. Zhao, 21 July 2022, CLZhao 22940 (SWFC).
Many recently described wood-inhabiting fungal taxa have been reported in the subtropics and tropics, including in the genus Trechispora (
Based on ITS+nLSU topology (Fig.
Morphologically, Trechispora albofarinosa resembles T. olivacea K.Y. Luo & C.L. Zhao and T. yunnanensis C.L. Zhao by sharing the farinosa basidiomata. However, T. olivacea differs from T. albofarinosa by olivaceous hymenial surface and thick-walled basidiospores (
Trechispora bisterigmata is similar to T. fastidiosa (Pers.) Liberta by sharing the membranous basidiomata. However, T. fastidiosa differs from T. bisterigmata by smooth hymenial surface and larger basidiospores (6–7 × 4.5–5.5 µm;
Trechispora pileata is similar to T. byssinella (Bourdot) Liberta, T. kavinioides B. de Vries, T. silvae-ryae (J. Erikss. & Ryvarden) K.H. Larss. and T. subsphaerospora (Litsch.) Liberta by sharing smooth basidiospores. However, T. byssinella differs from T. pileata by having narrower ellipsoid basidiospores (
Trechispora wenshanensis resembles T. fastidiosa and T. laevispora Z.B. Liu, Y.D. Wu & Yuan Yuan by sharing a smooth hymenial surface. However, T. fastidiosa differs from T. wenshanensis by larger basidiospores (6–7 × 4.5–5.5 µm;
1 | Basidiomata with clavarioid | 2 |
– | Basidiomata without clavarioid | 6 |
2 | Basidiomata grayish brown to pale purple | 3 |
– | Basidiomata pure white to pale yellow | 4 |
3 | Basidiomata with dense branches and long terminal branches | T. longiramosa |
– | Basidiomata with loose branches | T. laxa |
4 | Basidiomata with flattened branches | 5 |
– | Basidiomata without flattened branches | T. tongdaoensis |
5 | Basidiomata branches polychotomous | T. alba |
– | Basidiomata branches dichotomous | T. khokpasiensis |
6 | Basidiomata pileate | T. pileata |
– | Basidiomata resupinate to effused | 7 |
7 | Hymenophore poroid | 8 |
– | Hymenophore smooth, colliculose, irpicoid, grandinioid, odontioid, hydnoid | 13 |
8 | Hyphal system dimitic | T. dimitiella |
– | Hyphal system monomitic | 9 |
9 | Subicular hyphae thick-walled | 10 |
– | Subicular hyphae thin-walled | 11 |
10 | Ampullate septa present on subicular hyphae | T. mollusca |
– | Ampullate septa absent on subicular hyphae | T. suberosa |
11 | Crystals in subiculum as numerous rodlets | T. candidissima |
– | Crystals in subiculum as rhomboidal plates or various shapes | 12 |
12 | Sphaerocysts present in cords and the adjacent part of subiculum | T. hymenocystis |
– | Sphaerocysts absent | T. subhymenocystis |
13 | Basidiospores smooth | 14 |
– | Basidiospores ornamented | 16 |
14 | Basidiomata with rhizomorph | T. bisterigmata |
– | Basidiomata without rhizomorph | 15 |
15 | Basidiospores subglobose, angular to turbinate | T. confinis |
– | Basidiospores ellipsoid | T. laevispora |
16 | Basidiomata < 50 µm thick | 17 |
– | Basidiomata > 50 µm thick | 19 |
17 | Crystals absent | T. gracilis |
– | Crystals present | 18 |
18 | Crystals aggregated, rhomboidal fakes | T. perminispora |
– | Crystals butterfly-like, easily broken into irregular shapes | T. subaraneosa |
19 | Hymenophore smooth | 20 |
– | Hymenophore colliculose, irpicoid, grandinioid, odontioid, hydnoid | 27 |
20 | Basidiospores slightly cyanophilous | T. incisa |
– | Basidiospores acyanophilous | 21 |
21 | Basidiospores > 6.5 µm long | T. yunnanensis |
– | Basidiospores < 6.5 µm long | 22 |
22 | Generative hyphae < 2 µm in diameter | T. wenshanensis |
– | Generative hyphae > 2 µm in diameter | 23 |
23 | Generative hyphae thin-walled | 24 |
– | Generative hyphae thick-walled | 25 |
24 | Hymenophore farinaceous | T. larssonii |
– | Hymenophore arachnoid | T. subfarinacea |
25 | Generative hyphae > 3.5 µm in diameter | T. latehypha |
– | Generative hyphae < 3.5 µm in diameter | 26 |
26 | Basidiospores ellipsoid, thin-walled | T. albofarinosa |
– | Basidiospores broadly ellipsoid to globose, thick-walled | T. olivacea |
27 | Hymenial surface colliculose, irpicoid or grandinioid | 28 |
– | Hymenial surface odontioid or hydnoid | 30 |
28 | Generative hyphae thick-walled | T. murina |
– | Generative hyphae thin-walled | 29 |
29 | Growth on bamboo | T. taiwanensis |
– | Growth on other plant | T. crystallina |
30 | Tramal hyphae thin-walled or slightly thick-walled | 31 |
– | Tramal hyphae distinctly thick-walled | 35 |
31 | Crystals absent in trama | T. tropica |
– | Crystals present in trama | 32 |
32 | Basidiospores subglobose to globose | T. odontioidea |
– | Basidiospores ellipsoid or broadly ellipsoid | 33 |
33 | Tramal hyphae 3–6 µm wide, spines of basidiospores constricted | T. constricta |
– | Tramal hyphae 2–4 µm wide, spines of basidiospores not constricted | 34 |
34 | Cystidia present | T. chaibuxiensis |
– | Cystidia absent | T. nivea |
35 | Hymenophore aculei > 0.4 mm long | 36 |
– | Hymenophore aculei < 0.4 mm long | 39 |
36 | Margin smooth | T. fissurata |
– | Margin fimbriate | 37 |
37 | Basidiomata irpicoid | T. dentata |
– | Basidiomata odontioid or hydnoid | 38 |
38 | Hymenophore aculei sparse, cream to buff-yellow when fresh | T. fimbriata |
– | Hymenophore aculei dense, white when fresh | T. fragilis |
39 | Generative hyphae ampullate septa absent | T. bambusicola |
– | Generative hyphae ampullate septa present | 40 |
40 | Basidiospores with sharp spines | T. subfissurata |
– | Basidiospores without sharp spines | 41 |
41 | Spines of basidiospores constricted | T. subsinensis |
– | Spines of basidiospores not constricted | T. sinensis |
The authors have declared that no competing interests exist.
No ethical statement was reported.
The research was supported by the National Natural Science Foundation of China (Project Nos: 32170004, U2102220), Forestry Innovation Programs of Southwest Forestry University (Project No: LXXK-2023Z07), and the Yunnan Province College Students Innovation and Entrepreneurship Training Program (Project no. s202310677034), and the High-level Talents Program of Yunnan Province (YNQR-QNRC-2018-111).
Conceptualization, C.–L.Z.; methodology, C.–L.Z. and K.–Y.L.; software, C.–L.Z. and K.–Y.L.; validation, C.–L.Z. and K.–Y.L.; formal analysis, C.–L.Z., K.–Y.L. and J.–Q.S.; investigation, C.–L.Z., K.–Y.L. and J.–Q.S.; resources, C.–L.Z.; writing–original draft preparation, C.–L.Z. and K.–Y.L; writing–review and editing, C.–L.Z. and K.–Y.L; visualization, C.–L.Z. and K.–Y.L; supervision, C.–L.Z.; project administration, C.–L.Z.; funding acquisition, C.–L.Z. All authors have read and agreed to the published version of the manuscript.
Jiang-Qing Su https://orcid.org/0009-0008-5480-4502
Chang-Lin Zhao https://orcid.org/0000-0002-8668-1075
All of the data that support the findings of this study are available in the main text.