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MycoKeys 83: 85–103 (2021)

doi: 10.3897/mycokeys.83.71405 RESEARCH ARTICLE
https://mycokeys.pensoft.net Launched to accelerate biodiversity research

Two new psathyrelloid species of Coprinopsis

(Agaricales, Psathyrellaceae) from China

Gu Rao1, Dan Dai1, Hui-Nan Zhao1, Yi Liang1, Yu Li1, Bo Zhang1

1 Engineering Research Center of Edible and Medicinal Fungi, Ministry of Education, Jilin Agricultural Uni-
versity, Changchun, Jilin 130118, Changchun, China

Corresponding authors: Yu Li ([email protected]), Bo Zhang ([email protected])

Academic editor: María P. Martín | Received 10 July 2021 | Accepted 20 August 2021 | Published 8 September 2021

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

Abstract
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.

Keywords
Asia, molecular systematics, morphology, new taxa, taxonomy

Introduction

Coprinoid mushrooms are fascinating fungal taxa with the characteristic of deliques-
cent lamellae. Coprinus sensu lato is not monophyletic (Johnson and Vilgalys 1998;
Hopple and Vilgalys 1999; Park et al. 1999a, 1999b; Moncalvo et al. 2002). Based on
molecular studies, Redhead et al. (2001) subdivided Coprinus s. l. into four genera,

Copyright Gu Rao et al. 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.
86 Gu Rao et al. / MycoKeys 83: 85–103 (2021)

Coprinus Pers. (Agaricaceae), Corinopsis P. Karst. (Psathyrellaceae), Coprinellus P. Karst.

(Psathyrellaceae) and Parasola Redhead, Vilgalys & Hopple (Psathyrellaceae). Com-
bined with anatomical characteristics, 100 species were divided from Coprinus s. l.
into Coprinopsis, which were then widely accepted. The subdivision of the Coprinus s.
l. was full of controversies at the time. Noordeloos et al. (2005) argued that there were
many unresolved problems in the phylogeny and it was too early to subdivide it into
four genera.
Schafer (2010) used the classification system of Redhead et al. (2001) to divide
Coprinopsis into five sections, Atramentarii, Lanatuli, Alachuani, Narcotici and Nivei,
corresponding to Coprinus sect. Atramentarii (Fries 1836), C. sect. Lanatuli (Fries
1836), C. C. subsect. Alachuani (Singer 1949), C. C. subsect. Narcotici (Uljé and
Noordeloos 1993) and C. C. subsect. Nivei (Citerin 1992), but the traditional sec-
tion C. sect. Picacei (Kauffman 1918) was not covered. Wächter and Melzer (2020)
subdivided the genus Coprinopsis into 20 sections according to the subclades of the
phylogenetic tree (Cinereae, Filamentiferae, Melanthinae, Alopeciae, Xenobiae, Phlyc-
tidosporae, Krieglsteinerorum, Erythrocephalae, Geesteranorum, Mitraesporae, Radiatae,
Subniveae and Canocipes).
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 (Redhead et al. 2001). According to Kirk et al. (2008), 200 species
of Coprinopsis were known so far the world over. At the time of submission, there are
228 records of Coprinopsis in Index Fungorum (www.indexfungorum.org), including
synonyms, varieties, forms and names. There are few studies on Coprinopsis in China.
According to the report of Huang (2019), there were only 16 species of Coprinopsis
in China, amongst which eight species were newly recorded in China and reported in
2019. Based on traditional morphology, sequence data and phylogenetic analyses, two
new species of Coprinopsis were found in Jilin Province, China. They both belong to
the C. sect. Melanthinae and will be reported as follows.

Materials and methods

Collecting and morphological studies

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 Au-
tonomous 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 (HMJAU).
Photos of fresh basidiomata were taken in the field. The macromorphology was ob-
served from fresh basidiomata and the observation of microstructure was based on dry
specimens under a light microscope (LEICA DM1000). The mainly used reagents
are 5% potassium hydroxide (KOH) solution, 1% Congo Red and Melzer’s Reagent.
 Two new Psathyrelloid species Coprinopsis 87

The morphological description referred to Largent et al. (1977 and 1978). The surface
of the basidiospores was observed and photographed under a scanning electron mi-
croscope (SEM) (Hitachi SU8000) at 2.0 kV, with a working distance of 8 mm. The
following symbols were used in the description: [n/m/p] indicates that ‘n’ randomly se-
lected basidiospores from ‘m’ basidiomata of ‘p’ collections were measured, ‘avl’ means
the average length of basidiospores, except the extreme values, ‘avw’ means the average
width of the basidiospores, except the extreme values, ‘Q’ represents the quotient of
the length and width of a single basidiospores in side view, ‘Qm’ refers to the average Q
value of all basidiospores ± standard deviation. Dimensions for basidiospores are given
as (a) b–c (d). The range of b–c contains a minimum of 90% of the measured values.
Extreme values (i.e. a and b) are given in parentheses.

Research methods of molecular systematics

The total DNA of the specimens was extracted by the new plant genomic DNA extrac-
tion kit from Jiangsu Kangwei Century Biotechnology Company Limited. The am-
plification primers of LSU nrDNA (LSU) were LROR and LR5 (Vilgalys and Hester
1990), the ITS nrDNA (ITS) regions were ITS1 and ITS4 (White et al. 1990; Gardes
and Bruns 1993). The amplification reactions were carried out in a 25 μl system and
the total amount of the reactions was as follows: ddH2O 13.5 μl, 10×Taq Buffer 5 μl,
10 mM dNTPs 1 μl, 10 mM upstream primer 1 μl, 10 mM downstream primer 1 μl,
DNA sample 2 μl, 2 U/μm Taq Polymerase 1.5 μl. The cycle parameters were as fol-
lows: 4 min at 94 °C for 1 cycle; 40 s at 94 °C, 40 s at 54 °C, 1 min at 72 °C for 35
cycles; 10 min at 72 °C for 1 cycle; storage at 4 °C. The PCR product was subjected to
0.5% agarose gel electrophoresis to test strips. The sequencing work was entrusted to
Shenggong Bioengineering (Shanghai) Company Limited and the sequencing results
were clipped with Seqman 7.1.0 (Swindell and Plasterer 1997) and then submitted to
GenBank (https://www.ncbi.nlm.nih.gov/genbank/). The newly-obtained sequences
are shown in Table 1. The sequences of relevant taxa were downloaded from GenBank
and from the related articles (Larsson and Örstadius 2008; Nagy et al. 2010; Nagy
2012; Örstadius et al. 2015; Crous 2017; Melzer et al. 2017).
The ‘auto’ strategy and normal alignment mode of MAFFT (Katoh et al. 2005)
were used for Sequence alignment and Gblocks (Castresana et al. 2000; Talavera and
Castresana 2007) was used to obtain the conservative segments of sequences with
the following parameters: the minimum number of sequences for a conserved/flank

Table 1. Taxa, vouchers and sequence accession numbers of newly generated sequences.

Taxon Voucher ITS nrDNA LSU nrDNA

Coprinopsis pusilla HMJAU 58779 MZ398012 MZ398067
C. pusilla HMJAU 58780 MZ398013 MZ398068
C. pusilla HMJAU 58781 MZ398014 MZ398069
C. jilinensis HMJAU 58782 MZ398015 MZ398070
C. jilinensis HMJAU 58783 MZ398016 MZ398071
88 Gu Rao et al. / MycoKeys 83: 85–103 (2021)

Table 2. The best models, based on ModelFinder and MegaX.

BI ML NJ
ITS nrDNA SYM+G4 TVMe+G4 T92+G5
LSU nrDNA K2P+I TIM2+F+I K2+G5

position (12/12), the maximum number of contiguous non-conserved positions (8),

minimum length of a block (10) and allowed gap positions (with half ). ModelFinder
(Kalyaanamoorthy et al. 2017) was used to select the best-fit models using the Bayes-
ian Information Criterion (BIC). The Maximum Likelihood (ML) analyses were per-
formed in IQTree 1.6.8 (Nguyen et al. 2015) and the Bayesian Inference phylogenies
were performed in MrBayes 3.2.6 (Ronquist et al. 2012) (2 parallel runs, 2000000
generations), in which the initial 25% of sampled data were discarded as burn-in.
The above software was integrated into PhyloSuite 1.2.2 (Zhang et al. 2020). The
neighbour-joining (NJ) tree was carried out in Mega X (Kumar et al. 2018). The ML
and NJ trees were evaluated by bootstrap analysis with 1000 replicates and the best
models are shown in Table 2.

Results
BLASTn results

In the BLASTn alignment, based on ITS sequences, Coprinopsis pusilla and C. me-
lanthina KC992961 (Örstadius et al. 2015) had the highest sequence identities
(94.92%–95.26%), with 34–36 base differences. Corpinopsis jilinensis and C. uliginicola
MG712323 (Zhang 2019) had the highest sequence identity (99.27%), with five base
differences. The sequence identities between C. jilinensis and C. uliginicola KC992960
Type (Örstadius et al. 2015) were 93.98%–94.35%, with 39–42 base differences. In
BLASTn alignment, based on LSU sequences, C. pusilla and C. melanthina KC992961
(Örstadius et al. 2015) had the highest sequence identities (99.14%–99.25%), with
7–8 base differences. Corpinopsis jilinensis and C. uliginicola KC992960 Type (Örstadi-
us et al. 2015) had the highest sequence identity (99.13%), with eight base differences.

Phylogenetic analyses
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 1, 2),
Coprinopsis pusilla and C. jilinensis both belong to C. sect. Melanthinae. Coprinopsis
pusilla and C. melanthina formed a sister clade, C. jilinensis and C. uliginicola formed a
sister clade, both of which were strongly supported. In the phylogenetic trees, based on
 Two new Psathyrelloid species Coprinopsis 89

100/100/1 Coprinopsis pannucioides DQ389727

77/90/0.99 Coprinopsis pannucioides FN396143
Coprinopsis canoceps KC992964
Coprinopsis marcescibilis DQ389728
Coprinopsis marcescibilis FM878020
71/97/0.97 Coprinopsis marcescibilis FM878021
33/43/0.68
Coprinopsis pseudomarcescibilis KY698007
99/97/1
Coprinopsis pseudomarcescibilis NR_158341 Type
Coprinopsis musae KC992965 Type
48/44/0.79
100/100/1 Coprinopsis musae KC992966
89/83/0.99
100/100/1 Coprinopsis udicola KC992967 Type
Coprinopsis udicola KY698004
34/*/0.5 100/100/1 Coprinopsis submicrospora HQ847053
Coprinopsis submicrospora KC992959 Type
100/100/1 Coprinopsis aesontiensis NR_158339 Type
Coprinopsis jilinensis MZ398015 Type
Coprinopsis jilinensis sp. nov
86/43/0.99 Coprinopsis jilinensis MZ398016
Coprinopsis uliginicola KC992960 Type
Coprinopsis cineraria KC992962 Type
79/99/0.99 Coprinopsis cineraria KC992963
Coprinopsis sp. MF163181
86/39/0.99 Coprinopsis sp. MG966286
Coprinopsis melanthina HQ872017
67/52/0.94
Coprinopsis melanthina KC992961
Coprinopsis melanthina KY698010
Coprinopsis melanthina KY698011
81/87/0.98
Coprinopsis pusilla MZ398013
100/99/1
Coprinopsis pusilla MZ398012 Coprinopsis pusilla sp. nov
Coprinopsis pusilla MZ398014 Type
100/100/1 Parasola conopilea DQ389725
Parasola auricoma FM163185

0.04

Figure 1. 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 in-
dicates 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.

ITS sequences (Fig. 1), C. submicrospora and C. marcescibilis/C. pseudomarcescibilis/C.

musae/C. udicola formed a sister clade in the ML and BI trees (MLBP/BIPP: 34/0.5),
but C. submicrospora and C. canoceps/C. pannucioides formed a sister clade in the NJ
tree (NJBP: 39). In the phylogenetic trees, based on LSU nrDNA sequences (Fig. 2),
C. marcescibilis and C. pseudomarcescibilis formed a sister clade with strong support in
the NJ and BI trees (NJBP/BIPP: 99/0.8). However, in the ML tree, C. marcescibilis
formed a sister clade with all taxa, except the outgroups (MLBP: 100) and C. pseu-
domarcescibilis formed a sister clade with all taxa, except the outgroups and C. marcesci-
90 Gu Rao et al. / MycoKeys 83: 85–103 (2021)

Coprinopsis marcescibilis FM876277

*/99/0.80

Coprinopsis pseudomarcescibilis NG_059042 Type

Coprinopsis musae KC992965 Type

100/100/1
30/100/1
Coprinopsis musae KC992966

Coprinopsis pannucioides FN396202

Coprinopsis svubmicrospora HQ847133

*/99/1 100/100/1
43/77/0.36
*/*/0.62 Coprinopsis submicrospora KC992959 Type

Coprinopsis udicola KC992967 Type

*/*/0.56
Coprinopsis canoceps KC992964
54/77/0.77
Coprinopsis cineraria KC992963

Coprinopsis jilinensis MZ398070 Type

100/97/0.99
100/100/1 Coprinopsis jilinensis sp. nov
98/98/0.99 Coprinopsis jilinensis MZ398071

Coprinopsis uliginicola KC992960 Type

Coprinopsis melanthina HQ847137

84/86/0.97 100/99/1

Coprinopsis melanthina KC992961

100/100/1
Coprinopsis pusilla MZ398068

99/99/0.83 Coprinopsis pusilla MZ398067 Coprinopsis pusilla sp. nov

Coprinopsis pusilla MZ398069 Type

Parasola conopilea DQ389725

100/100/1

Parasola auricoma JQ045871

0.009

Figure 2. 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.

bilis (MLBP: 30). In the analyses, based on different methods, the sister branch relation-
ships of C. pannucioides, C. submicrospora, C. uliginicola and C. canoceps were different.
In the BI tree, C. pannucioides and C. submicrospora/C. udicola/C. canoceps formed a
sister clade with strong support (BIPP: 1), C. canoceps and C. submicrospora/C. udicola
formed a sister clade (BIPP: 0.56), C. submicrospora and C. udicola formed a sister
clade (BIPP: 0.62). In the ML tree, C. submicrospora and C. udicola/C. canoceps/C.
pannucioides formed a sister clade with strong support (MLBP: 98), C. udicola formed
a sister clade with C. pannucioides/C. canoceps (MLBP: 30) and C. pannucioides and
C. canoceps formed a sister clade (MLBP: 50). In the NJ tree, C. pannucioides and C.
submicrospora/C. udicola/C. canoceps formed a sister clade with strong support (NJBP:
99), C. submicrospora formed a sister clade with C. udicola /C. canoceps (NJBP: 38) and
C. udicola and C. canoceps formed a sister clade (NJBP: 41).
 Two new Psathyrelloid species Coprinopsis 91

Taxonomy
Coprinopsis jilinensis G. Rao, H.N. Zhao, B. Zhang & Y. Li, sp. nov.
MycoBank No: 840297
Figures 3, 4, 7C, D

Typification. China. Red Leaves Valley in Hanchongling, Dunhua City, Yanbian Ko-
rean Autonomous Prefecture, Jilin Province, 22 August 2019, G. Rao & H.N Zhao
(HMJAU 58782 Holotype!).
Sequences ex holotype. MZ398015 (ITS nrDNA), MZ398070 (LSU nrDNA).
Etymology. The epithet “jilinensis” refers to this species that was first discovered
in Jilin Province, China.
Description. Basidiomata small to medium-sized. Pileus 33–52 mm broad, coni-
cal 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 ad-
nexed, not the same length and width, edges slightly toothed, concolorous, not deli-
quescent. 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-sterig-
mate 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.
Habitat and distribution. On humus of broad-leaved forest or coniferous and
broad-leaved mixed forests in autumn.
Additional specimens examined. China. Red Leaves Valley in Hanchongling,
Dunhua City, Yanbian Korean Autonomous Prefecture, Jilin Province, 14 September
2019, Gu Rao (HMJAU 58783).
Notes. Coprinopsis jilinensis is characterised by its small to medium-sized basidi-
omata, brown pileus with white hairs, smooth and dextrinoid basidiospores with small
92 Gu Rao et al. / MycoKeys 83: 85–103 (2021)

Figure 3. Basidiomata and microscopic features of Coprinopsis jilinensis A collection HMJAU 58783
B collection HMJAU 58782 C basidiospores D basidia E pleurocystidia F Cheilocystidia G pileipellis.
Scale bars: 5 mm (A, B); 10 µm (C–G).

pores, pleurocystidia and cheilocystidia present. C. jilinensis forms a strongly-support-

ed independent clade in ITS and LSU phylogeny trees (Figs 1, 2).
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 (Smith 1972). Other similar
species, C. cineraria is characterised by grey, hygrophanous and striate pileus, little short
 Two new Psathyrelloid species Coprinopsis 93

Figure 4. Scatter plot of basidiospores size in Coprinopsis jilinensis.

basidiospores (6.5–8.5 µm), pleurocystidia absent, pileipellis an epithelium (Takahashi

2000); C. melanthina is characterised by little long and subcolourless basidiospores (avl
× avw = 10.5 × 5.8 µm), pleurocystidia absent (Kits van Waveren 1985); C. pusilla has
small basidiomata, grey pileus, subcolourless and verrucose basidiospores (this study)
and Psathyrella subagraria has hygrophanous pileus, thick flesh and caulocystidia pre-
sent (Smith 1972), both of which could be clearly distinguished from C. jilinensis in
terms of morphology.

Coprinopsis pusilla G. Rao, B. Zhang & Y. Li, sp. nov.

MycoBank No: 840298
Figures 5, 6, 7A, B

Typification. China. Red Leaves Valley in Hanchongling, Dunhua City, Yanbian

Korean Autonomous Prefecture, Jilin Province, 21 August 2019, Gu Rao (HMJAU
58781 Holotype!).
Sequences ex holotype. MZ398014 (ITS nrDNA), MZ398069 (LSU nrDNA).
Etymology. The epithet “pusilla” refers to this species having small basidiomata.
94 Gu Rao et al. / MycoKeys 83: 85–103 (2021)

Figure 5. Basidiomata and microscopic features of Coprinopsis pusilla A collection HMJAU 58780
B collection HMJAU 58781 C collection HMJAU 58779 D basidiospores E basidia and basidioles
F cheilocystidia G pileipellis. Scale bars: 5 mm (A–C); 10 µm (D–G).

Description. 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 eas-
ily disappearing. Lamellae close or crowded, subwhite, greyish-white or coffee brown,
 Two new Psathyrelloid species Coprinopsis 95

Figure 6. Scatter plot of basidiospores size in Coprinopsis pusilla.

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 floccu-
lent 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. Cheilocys-
tidia (25) 27–53 (55) × (11) 13–21 µm, variable-shaped, subcylindrical, utriform,
lageniform, reverse gourd-shaped and subcapitate, sphaeropedunculate elements pre-
sent 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)
96 Gu Rao et al. / MycoKeys 83: 85–103 (2021)

Figure 7. Scanning electron micrograph of basidiospores A, B Coprinopsis pusilla C, D C. jilinensis. Scale

bars: 2 µm (A–D).

10–19 (20) µm, without encrusting pigment, thick wall, colourless to yellowish, cy-
lindrical, 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 cylindri-
cal protrusions. Clamp connections present in all tissues.
Habitat and distribution. On the dead and rotten wood of broad-leaved forest or
coniferous and broad-leaved mixed forests in autumn.
Additional specimens examined. China. Red Leaves Valley in Hanchongling,
Dunhua City, Yanbian Korean Autonomous Prefecture, Jilin Province, 6 August 2019,
G. Rao (HMJAU 58779, HMJAU 58780).
Notes. Coprinopsis pusilla has a variable macromorphology, but stable micromor-
phology, 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 1, 2).
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 van Waveren 1985). C. uliginicola is characterised by large basidiomata,
brown-black basidiospores, pore present (Smith 1972). C. cineraria is characterised by
fibrous veil at the edges easily disappearing, smaller basidiospores (6.5–8.5 × 4–5 µm),
pileipellis an epithelium (Takahashi 2000).
 Two new Psathyrelloid species Coprinopsis 97

Discussion
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
(Redhead et al. 2001). Örstadius et al. (2015) understood that “psathyrelloid” means
that certain species of fungi with the morphology of Psathyrella, but belong to other
genera in the phylogenetic analysis, so the species of Psathyrella were not included.
These fungi were mainly distributed in Coprinopsis (Örstadius et al. 2015), Typhrasa
(Örstadius et al. 2015; Wang et al. 2021), Parasola (Ganga and Manimohan 2019),
Homophron (Örstadius et al. 2015), Cystagaricus (Örstadius et al. 2015), Kauffmania
(Örstadius et al. 2015), Lacrymaria (Yan 2018) and so on. According to Redhead et
al. (2001), coprinoid mushrooms were distributed in four genera: Coprinus, Parasola,
Coprinopsis and Coprinellus. In Agaricaceae, Coprinoid mushrooms should also in-
clude Montagnea, Podaxis and Xerocoprinus (Moncalvo et al. 2002; Keirle et al. 2004).
As there was no uniform standard, some fungi, such as Parasola conopilea (Ganga and
Manimohan 2019), were intermediate between psathyrelloid and coprinoid species.
With the introduction of molecular systematics, the species of coprinoid and psath-
yrelloid intersected with each other. Örstadius et al. (2015) studied the morphology of
many psathyrelloid species of Psathyrellaceae and believed that the lack of the typical
pattern of pseudoparaphyses was their main morphological characteristic. Since psath-
yrelloid species belong to different genera, their distributions were not indicated phy-
logenetically in their study. Worldwide, there were, currently, at least fourteen psath-
yrelloid species of Coprinopsis (Picón 2010; Portugalete 2011; Örstadius et al. 2015;
Crous 2017; Larsson and Örstadius 2017; Melzer et al. 2017), which belong to C. sect.
Lanatulae in Schafe’s grouping system (Schafe 2010) and to three sections in Wächter’s
grouping system, Melanthinae (including C. cineraria, C. melanthina, C. uliginicola,
C. jilinensis and C. pusilla), Canocipes (including C. aesontiensis, C. canoceps, C. loti-
nae, C. pannucioides, C. submicrospora and C. udicola) and Quartoconatae (including
C. marcescibilis, C. musae and C. pseudomarcescibilis). Currently, the known sequences
of these species in these three sections are all the psathyrelloid species of Coprinopsis.
According to the results of BLASTn analyses, ITS showed higher interspecific variabili-
ty, 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 pro-
posed by Wächter and Melzer (2020). In the phylogenetic analyses (Figs 1, 2), whether
in the phylogenetic trees, based on ITS or LSU, the tree shape of C. sect. Melanthinae
obtained was consistent and the branches were stable. However, C. sect. Canocipes and C.
sect. Quartoconatae were somewhat different from the grouping described by Wächter and
Melzer (2020), based on different sequences and analyses. In the phylogenetic trees, based
98 Gu Rao et al. / MycoKeys 83: 85–103 (2021)

on ITS, C. udicola and C. submicrospora, which originally belonged to C. sect. canocipes,

formed a branch with three species belonging to C. sect. Quartoconatae in ML and BI
trees. However, in the NJ tree, C. udicola formed a sister clade with three species belonging
to C. sect. Quartoconatae. In the phylogenetic trees, based on LSU, C. sect. Canocipes and
C. sect. Quartoconatae were well separated in the NJ tree, while in ML and BI trees, C.
musae belonging to C. sect. Quartoconatae formed a sister clade with four species of C. sect.
Canocipes. The branching relationships of phylogenetic trees, based on different molecular
sequences and analyses may vary. Over-subdivision of sections would cause intersection.
Subdivision of C. sect. Canocipes and C. sect. Quartoconatae may need to be reconsidered.
Coprinopsis sect. Melanthinae has a relatively clear systematic differentiation
(Wächter and Melzer 2020), which is consistent with the results of this study. Cur-
rently, there are only six species in this section, including the C. lignicola nom. prov.
(GenBank no.: MG966286 and MF163181). Combined with the two newly-discov-
ered species, the expression of this section is modified as follows:

Coprinopsis sect. Melanthinae Wächter & A. Melzer

Description. Basidiomata very small to large, on humus or lignicolous. Pileus not ra-
dially sulcate, lamellae not deliquescent. Veil strongly developed, consisting of chains
of subcylindrical, sometimes encrusted cells. Basidiospores medium to large-sized, el-
lipsoid 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 col-
lection 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 HMJAU 58779
and collection HMJAU 58780 were collected at approximately the same time in the same
forest, which were not far apart from each other. However, due to the significant difference
in macroscopic morphology, they were made into two collections. The macroscopic mor-
phology of Collection HMJAU 58781, collected later, was also significantly different from
the two specimens collected previously. Through the observation of the microscopic mor-
phology of these collections and the phylogenetic analysis, combined with ITS and LSU
molecular sequences, the results showed that the three collections were the same species.
Pleurocystidia is present in many species of Coprinopsis, such as C. cinerea, C. jonesii
and C. pseudoradiata (Redhead et al. 2001), but in psathyrelloid species of Coprinopsis, spe-
 Two new Psathyrelloid species Coprinopsis 99

cies with pleurocystidia are rare. The pleurocystidia are sparsely distributed in the hymeni-
um of C. jilinensis, but not present in C. uliginicola, which is closely related to C. jilinensis.
In psathyrelloid species of Coprinopsis, only C. pannucioides (Larsson and Örstadius 2008),
C. udicola (existing but rare) (Örstadius et al. 2015) and C. jilinensis have pleurocystidia.
Psathyrella subagraria is a confusing species, described by Smith (1972) as it is very
similar to P. uliginicola morphologically, with the main difference being that this species
has pleurocystidia, which are mainly growing on humus. Since Smith (1972) introduced
P. subagraria, no further reports have been made. Örstadius et al. (2015) moved P. ulig-
inicola to Coprinopsis, based on molecular studies, the taxonomic status of P. subagraria
being questionable. There are some differences between P. subagraria and C. jilinensis in
macroscopic morphology, but they are similar in microscopic morphology. Smith (1972)
described P. subagraria as having two basidiospores sizes [8–10 × 4–5 (10–12 × 4.5–5.5)
µm], one of which was very close to the size of C. jilinensis [(8) 8.5–10 (10.2) × 4.5–5.9
(6) µm], requiring re-examination of the type specimens of P. subagraria in future studies.

Key to fourteen psathyrelloid species of Coprinopsis

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 epithe-
lium........................................................................................... 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
100 Gu Rao et al. / MycoKeys 83: 85–103 (2021)

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 scat-
tered...........................................................................................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 fusi-
form........................................................................................... C. canoceps
– Stipe > 60 mm long; basidia 1-, 2-, 4-sterigmate; cheilocystidia part inflated
fusiform................................................................................. C. aesontiensis

Acknowledgements
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 Pro-
ject (No. 20190201026JC), Modern Agroindustry Technology Research System (No.
CARS20) and Scientific and Technological Tackling Plan for the Key Fields of Xinji-
ang Production and Construction Corps (No. 2021AB004).

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