Research Article |
Corresponding author: Yu Li ( yuli996@126.com ) Corresponding author: Bo Zhang ( zhangbofungi@126.com ) Academic editor: María P. Martín
© 2021 Gu Rao, Dan Dai, Hui-Nan Zhao, Yi Liang, Yu Li, Bo Zhang.
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:
Rao G, Dai D, Zhao H-N, Liang Y, Li Y, Zhang B (2021) Two new psathyrelloid species of Coprinopsis (Agaricales, Psathyrellaceae) from China. MycoKeys 83: 85-103. https://doi.org/10.3897/mycokeys.83.71405
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In this study, Coprinopsis jilinensis and Coprinopsis pusilla were introduced, based on their morphological characteristics, the internal transcribed spacer (ITS) and large subunit ribosomal (LSU) region sequences of nrDNA. These new psathyrelloid species were found in Jilin Province, China. Coprinopsis jilinensis has brown pileus, utriform pleurocystidia, brown, smooth, dextrinoid basidiospores and tiny pore. It mainly grows on humus. Coprinopsis pusilla has small basidiomata, greyish-white pileus, thick and distinct veil at edges, subcolourless and verrucose basidiospores. It is poreless and it grows on the decaying wood of broad-leaved trees. Both of them belong to the C. sect. Melanthinae. A supplementary description of C. sect. Melanthinae was given in combination with the newly-discovered taxa and an identification key to the fourteen psathyrelloid species of Coprinopsis is provided. Coprinopsis sect. Canocipes and C. sect. Quartoconatae were evaluated and the phylogenetic position of the psathyrelloid species of Coprinopsis was discussed. Psathyrella subagraria, as a confusing species, was also discussed in this study.
Asia, molecular systematics, morphology, new taxa, taxonomy
Coprinoid mushrooms are fascinating fungal taxa with the characteristic of deliquescent lamellae. Coprinus sensu lato is not monophyletic (
Coprinopsis is a worldwide fungal taxon that includes some well-known species, such as C. atramentaria, characterised by hyphal pileus cuticle, abundant powdery to floccose veil covering the whole pileus, coprophilous, growing in a terrestrial or lignicolous habitat (
The fresh basidiomata were collected from the Red Leaves Valley in Hanchongling (approximate 43°02'1.67"N, 127°59'36.55"E), Dunhua City, Yanbian Korean Autonomous Prefecture, Jilin Province, China. After dried at 45 °C for 1‒2 days, they were stored in the Herbarium of Mycology of Jilin Agricultural University (
The total DNA of the specimens was extracted by the new plant genomic DNA extraction kit from Jiangsu Kangwei Century Biotechnology Company Limited. The amplification primers of LSU nrDNA (LSU) were LROR and LR5 (
Taxa, vouchers and sequence accession numbers of newly generated sequences.
The ‘auto’ strategy and normal alignment mode of MAFFT (
In the BLASTn alignment, based on ITS sequences, Coprinopsis pusilla and C. melanthina KC992961 (
After Gblocks clipping, the ITS data matrix included 31 sequences of 588 nucleotide sites from 16 taxa and the data matrix included 20 sequences of 921 nucleotide sites from 14 taxa (gaps included). In the ITS and LSU phylogenetic trees (Figs
The phylogenetic tree of Coprinopsis by ITS nrDNA, based on the Maximum Likelihood method (ML). The three values of internal nodes respectively represent Maximum Likelihood bootstrap (MLBP)/neighbour-joining bootstrap (NJBP)/Bayesian posterior probability (BIPP). The thick node indicates the significantly-supported branch in at least two analyses (MLBP ≥ 70, NJBP ≥ 70, BIPP ≥ 95%). The GenBank accession number is marked after the species name. At the same time, the sequence from the type specimen is also marked at the end. Two new species from China are expressed in bold font and Parasola conopilea (Fr.) Örstadius & E. Larss and P. auricoma (Pat.) Redhead, Vilgalys & Hopple are selected as the outgroups.
The phylogenetic tree of Coprinopsis by LSU nrDNA, based on Bayesian Inference (BI). The three values of internal nodes respectively represent Maximum Likelihood bootstrap (MLBP)/neighbour-joining bootstrap (NJBP)/Bayesian posterior probability (BIPP). The thick node indicates the significantly-supported branch in at least two analyses (MLBP ≥ 70, NJBP ≥ 70, BIPP ≥ 95%). The GenBank accession number is marked after the species name. At the same time, the sequence from the type specimen is also marked at the end. Two new species from China are expressed in bold font and Parasola conopilea (Fr.) Örstadius & E. Larss and P. auricoma (Pat.) Redhead, Vilgalys & Hopple are selected as the outgroups.
China. Red Leaves Valley in Hanchongling, Dunhua City, Yanbian Korean Autonomous Prefecture, Jilin Province, 22 August 2019, G. Rao & H.N Zhao (
The epithet “jilinensis” refers to this species that was first discovered in Jilin Province, China.
Basidiomata small to medium-sized. Pileus 33–52 mm broad, conical to convex, dark brown or clay brown, densely covered with white hairs, not sticky when dry or wet, not hygrophanous, veil remnants flocculent at edges. Lamellae close or crowded, grey-white to fleshy brown, brownish-black after drying, sinuate or adnexed, not the same length and width, edges slightly toothed, concolorous, not deliquescent. Stipe 80–95 × 5–9 mm, white to milky white, cylindrical, down slightly rough, fibrous, a little fragile, hollow, the base with white mycelium, dense or sparse, close to the stipe surface covered with brownish-yellow pubescent, no ring. Spore print without record.
Basidiospores [60, 2, 2] (8)8.5–10(10.2) × 4.5–5.9 (6) µm, avl = 9.1 µm, avw = 5.2 µm, Q = (1.62) 1.63–1.96 (2.02), Qm = 1.77 ± 0.09, oval to long oval, brown, brownish-yellow or dark brown in 5% KOH solution, smooth, thick wall, dextrinoid, apical with small pores, 1–2 µm. Basidia 17–30 (39) × 8–10 (13) µm, clavate, 4-sterigmate up to 3–4 µm long, 2–3 sterigmate occasional. Pleurocystidia (30) 33–59 (60) × (11) 12–21 (23) µm, utriform and lageniform, sparse, smooth, hyaline. Cheilocystidia 27–56 × (10) 11–20 (22) µm, utriform and lageniform, smooth, hyaline, crowded in hymenium. Lamellar edge fertile. Pileipellis a cutis, up to 100 µm thick, hyphae (35) 42–111 (148) × (6) 7–34 (35) µm, ovoid, subcylindrical, with brownish-yellow to dark brown pigment, thick wall, encrusting pigment on the outer hyphae. Veil hyphae (5) 6–30 (33) µm wide, present dark encrusting pigment, thick wall, colourless to light yellow, cylindrical and subcylindrical. Stipitipellis a cutis, hyphae (5) 6–22 (32) µm diam., ovoid and subcylindrical, pale brown, with encrusting pigment, thick wall. Clamp connections present in all tissues.
On humus of broad-leaved forest or coniferous and broad-leaved mixed forests in autumn.
China. Red Leaves Valley in Hanchongling, Dunhua City, Yanbian Korean Autonomous Prefecture, Jilin Province, 14 September 2019, Gu Rao (
Coprinopsis jilinensis is characterised by its small to medium-sized basidiomata, brown pileus with white hairs, smooth and dextrinoid basidiospores with small pores, pleurocystidia and cheilocystidia present. C. jilinensis forms a strongly-supported independent clade in ITS and LSU phylogeny trees (Figs
Morphologically and phylogenetic similar to Coprinopsis jilinensis, C. uliginicola is characterised by long basidiospores of 10–12(–15) µm, pleurocystidia absent and caulocystidia present, pileipellis no encrusting pigment (
China. Red Leaves Valley in Hanchongling, Dunhua City, Yanbian Korean Autonomous Prefecture, Jilin Province, 21 August 2019, Gu Rao (
The epithet “pusilla” refers to this species having small basidiomata.
Basidiomata very small to small. Pileus 21–29 mm broad, bell-shaped to hemispherical when young, then convex, flat to slightly reflexed at edges, with inconspicuous bulge at the middle, grey or greyish-white when dry, no record when wet, densely covered with flocculent hairs, sometimes central with blackish-grey squamous tapering to the edges, not slime, sometimes the edges crack, hygrophanous no record, veil remnants dense at edges, triangular, subtriangular or massive, not easily disappearing. Lamellae close or crowded, subwhite, greyish-white or coffee brown, flesh blond after drying, sinuate or adnexed, sometimes with vertical teeth, edges slightly toothed, concolorous, not deliquescent. Stipe 35–57 × 3–7 mm, cylindrical, subcylindrical, subequal or a little rough towards the base, white, cream white, hollow, a little fragile, not easy to detach from the cap, densely covered with white and flocculent hairs, brown, brownish-grey to brownish-yellow near the base, veil present at the stalk and cap joints, easily disappearing, no ring, the base with white mycelium. Spore print without record.
Basidiospores [90, 4, 3] 8–12 × 5–6.5 (6.8) µm, avl = 9.8 µm, avw = 5.8 µm, Q = 1.45–2.2 (2.24), Qm = 1.70 ± 0.18 µm, oval, elliptic to long elliptic, subcolourless in 5% KOH and aqueous solution, surfaces verrucose, thin wall, no pores, not amyloid. Basidia (18) 19–32 (33) × 9–11 (12) µm, clavate, 4-sterigmate up to 3–4 µm long, 2-sterigmate occasional, without pseudoparaphyses. Pleurocystidia absent. Cheilocystidia (25) 27–53 (55) × (11) 13–21 µm, variable-shaped, subcylindrical, utriform, lageniform, reverse gourd-shaped and subcapitate, sphaeropedunculate elements present on gill edges, smooth, hyaline, thin wall to thick wall. Pileipellis a cutis, terminal hyphae (30) 31–84 (98) × (7) 8–17 (18) µm, with light brown pigment, mostly thick wall in the outer hyphae, present dark encrusting pigment, terminal hyphae present small cylindrical protrusions, about 3 × 3 μm. Veil hyphae (26) 27–100 (113) × (9) 10–19 (20) µm, without encrusting pigment, thick wall, colourless to yellowish, cylindrical, subcylindrical, clavate or irregular. Stipitipellis a cutis, hyphae (21) 22–87 (88) × (9) 10–19 (20) µm, encrusting pigment not observed, colourless to light yellow, cylindrical, subcylindrical, clavate or irregular, terminal hyphae present small cylindrical protrusions. Clamp connections present in all tissues.
China. Red Leaves Valley in Hanchongling, Dunhua City, Yanbian Korean Autonomous Prefecture, Jilin Province, 6 August 2019, G. Rao (
Coprinopsis pusilla has a variable macromorphology, but stable micromorphology, which is characterised by small basidiomata, greyish-white pileus, thick and distinct veil remnants at edges, subcolourless and verrucose basidiospores, no pore, the habitat on the decaying wood of broad-leaved trees. C. pusilla forms a strongly-supported independent clade in both ITS and LSU phylogeny trees (Figs
Morphologically and phylogenetically similar to Coprinopsis pusilla, C. melanthina is characterised by larger brown pileus, fibrous veil at edges, longer basidiospores (avl = 10.5 µm) (Kits
Here we report two new psathyrelloid species of Coprinopsis from northern China. There is no unified definition of “psathyrelloid”. Previously, “psathyrelloid” was mainly regarded as a species with the morphology of Psathyrella, including Psathyrella itself (
According to the results of BLASTn analyses, ITS showed higher interspecific variability, so the ITS sequence was more advantageous in reflecting the interspecific relationship of Coprinopsis than the LSU sequence. The sequence identity between C. jilinensis from Jilin Province, China and C. uliginicola MG712323 from Hubei Province, China, was 99.27%. Based on the molecular sequence alone, C. uliginicola MG712323 and C. jilinensis could be the same species and subsequent re-examination of this specimen is recommended.
In this study, some sequences of Coprinopsis were selected, which belong to C. sect. Melanthinae, C. sect. Canocipes and C. sect. Quartoconatae in the grouping system proposed by
Coprinopsis sect. Melanthinae has a relatively clear systematic differentiation (
Coprinopsis sect. Melanthinae Wächter & A. Melzer
Description. Basidiomata very small to large, on humus or lignicolous. Pileus not radially sulcate, lamellae not deliquescent. Veil strongly developed, consisting of chains of subcylindrical, sometimes encrusted cells. Basidiospores medium to large-sized, ellipsoid to ovoid in side view, strikingly pale or brown, thin or thick-walled, germ pore absent or very indistinct, surfaces verrucose or smooth. Pseudoparaphyses absent. Basidia 4-sterigmate, 2–3-sterigmate occasional, always clavate, never polymorphic. Marginal cells of the lamellar edge predominantly utriform. Pleurocystidia present or absent. Cheilocystidia and clamps present.
Coprinopsis pusilla has a variety of macroscopic morphology, including the shape and colour of the pileus, the shape of the veil and the thickness and length of the stipe and so on, which makes it difficult to determine whether it is the same species during the collection process, when the size of the basidiomata and ecological habits seem to be stable. The microscopic morphology of C. pusilla is relatively stable, including the size, shape, colour and decoration of basidiospores, the type of pileipellis and the presence or absence of cystidia. Interestingly, Coprinoid fungi are thought to be a taxon of dark-coloured basidiospores (Noordeloos et al. 2005), but C. pusilla, including C. melanthina, which is closely related to C. pusilla, have subcolourless basidiospores. Collection
Pleurocystidia is present in many species of Coprinopsis, such as C. cinerea, C. jonesii and C. pseudoradiata (
Psathyrella subagraria is a confusing species, described by
1 | Basidiospores no pore | 2 |
– | Basidiospores with pore | 6 |
2 | Basidiospores distinctly pigmented | C. submicrospora |
– | Basidiospores hyaline to very pale brown | 3 |
3 | Pileus < 20 mm, glabrous; pseudoparaphyses often present | C. musae |
– | Pileus > 20 mm, hairy; pseudoparaphyses absent or unrecorded | 4 |
4 | Pileus hygrophanous, striate; basidiospores avl < 9 μm; pileipellis an epithelium | C. cineraria |
– | Pileus not or scarcely hygrophanous, not striate; basidiospores avl > 9 μm; pileipellis a cutis | 5 |
5 | Pileus 25–80 mm, brown, fibrous veil at edges; basidiospores avl = 10.5 µm | C. melanthina |
– | Pileus 21–29 mm, grey or greyish-white, veil dense at edges; basidiospores avl = 9.8 μm | C. pusilla |
6 | Basidiospores avl > 11.5 μm | 7 |
– | Basidiospores avl < 11.5 μm | 10 |
7 | Pileus mostly brown, almost not striate, flocci at cap margin | 8 |
– | Pileus mostly white or grey, striate, no flocci at cap margin | 9 |
8 | Basidiospores 13.3–14.5 µm; irregular flocci on cap margin | C. pseudomarcescibilis |
– | Basidiospores 11.6–12.8 µm; denticulate flocci on cap margin | C. marcescibilis |
9 | Pseudoparaphyses present, pleurocystidia absent; caulocystidia present | C. lotinae |
– | Pseudoparaphyses absent; pleurocystidia rare, close to gill edge; caulocystidia absent | C. udicola |
10 | Pleurocystidia present; cheilocystidia no long neck | 11 |
– | Pleurocystidia absent; cheilocystidia part with long neck | 12 |
11 | Pileus yellowish-white, papillate-umbonate; caulocystidia present; tufty | C. pannucioides |
– | Pileus brown, not papillate-umbonate; caulocystidia absent; solitary or scattered | C. jilinensis |
12 | Basidiomata medium-sized to large-sized (pileus > 50 mm); pileus pallid to greyish | C. uliginicola |
– | Basidiomata very small to small (pileus < 50 mm); pileus not grey | 13 |
13 | Stipe < 60 mm long; basidia 4-sterigmate; cheilocystidia not inflated fusiform | C. canoceps |
– | Stipe > 60 mm long; basidia 1-, 2-, 4-sterigmate; cheilocystidia part inflated fusiform | C. aesontiensis |
We are grateful to the editors and reviewers for improving the manuscript. This study is funded by Key Project on R&D of Ministry of Science and Technology (No. 2018YFE0107800), Jilin Province Science and Technology Development Plan Project (No. 20190201026JC), Modern Agroindustry Technology Research System (No. CARS20) and Scientific and Technological Tackling Plan for the Key Fields of Xinjiang Production and Construction Corps (No. 2021AB004).