Revealing Brownish Mycena Diversity in China: New Discoveries and Taxonomic Insights

Abstract

Within the genus Mycena, species exhibiting brownish basidiomata present considerable challenges in identification due to similar coloration. This study underscores the significance of pileipellis types and cheilocystidia characteristics as critical in delimiting brownish Mycena species. To clarify the principal taxonomic characters and their utility in distinguishing between brownish Mycena species, a morphological taxonomy and phylogenetic analysis were performed. Five new species from China were introduced and characterized through a comprehensive morphological anatomy and phylogenetic substantiation: M. campanulatihemisphaerica sp. nov., M. digitifurcata sp. nov., M. kunyuensis sp. nov., M. limitis sp. nov., and M. oryzifluens sp. nov. Discussions of these taxa are supplemented with morphological illustrations. The phylogenetic relationships were inferred using Bayesian Inference and Maximum Likelihood methods based on sequences from the internal transcribed spacer and the large subunit regions of nuclear ribosomal RNA. With the addition of these five new species, the worldwide count of brownish Mycena increases to 94, and a key to the 29 known species of brownish Mycena from China is presented.

Full text

Citation: Wei, R.; Ge, Y.; Qi, L.; Han,
M.; Zeng, H.; Hu, Y.; Zou, L.; Cheng,
X.; Wu, X.; Na, Q. Revealing Brownish
Mycena Diversity in China: New
Discoveries and Taxonomic Insights. J.
Fungi 2024,10, 439. https://doi.org/
10.3390/jof10060439
Academic Editor: Jadson Diogo
Pereira Bezerra
Received: 5 May 2024
Revised: 17 June 2024
Accepted: 17 June 2024
Published: 20 June 2024
Copyright: © 2024 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
Fungi
Journal of
Article
Revealing Brownish Mycena Diversity in China: New Discoveries
and Taxonomic Insights
Renxiu Wei
1, †
, Yupeng Ge
1,2,3,†
, Liangliang Qi
4
, Menghui Han
1
, Hui Zeng
2,3
, Yaping Hu
5
, Li Zou
6
,
Xianhao Cheng 1, Xiaoming Wu 7and Qin Na 1,*
1Institute of Mycological Science and Technology, School of Agriculture, Ludong University,
Yantai 264025, China; wrx_1260@163.com (R.W.); gaiyupeng@126.com (Y.G.);
han_mh8128@163.com (M.H.); chengxianhao@sohu.com (X.C.)
2Institute of Edible Fungi, Fujian Academy of Agricultural Sciences, Fuzhou 350014, China; zenghui@faas.cn
3National and Local Joint Engineering Research Center for Breeding & Cultivation of Features Edible Fungi,
Fuzhou 350014, China
4Microbiology Research Institute, Guangxi Academy of Agricultural Sciences, Nanning 530007, China;
qiliangliang529@126.com
5Nanjing Institute of Environmental Sciences, Ministry of Ecology and Environment, State Environmental
Protection Scientific Observation and Research Station for Ecological Environment of Wuyi Mountains,
Nanjing 210042, China; huyap9009@163.com
6College of Forestry, Northeast Forestry University, Harbin 150040, China; 13903650896@163.com
7Kunyushan National Nature Reserve, Yantai 264112, China; w3215892@163.com
*Correspondence: naqin19890317@163.com
†These authors contributed equally to this work.
Abstract: Within the genus Mycena, species exhibiting brownish basidiomata present considerable
challenges in identification due to similar coloration. This study underscores the significance of
pileipellis types and cheilocystidia characteristics as critical in delimiting brownish Mycena species.
To clarify the principal taxonomic characters and their utility in distinguishing between brownish
Mycena species, a morphological taxonomy and phylogenetic analysis were performed. Five new
species from China were introduced and characterized through a comprehensive morphological
anatomy and phylogenetic substantiation: M. campanulatihemisphaerica sp. nov., M. digitifurcata sp.
nov., M. kunyuensis sp. nov., M. limitis sp. nov., and M. oryzifluens sp. nov. Discussions of these taxa
are supplemented with morphological illustrations. The phylogenetic relationships were inferred
using Bayesian Inference and Maximum Likelihood methods based on sequences from the internal
transcribed spacer and the large subunit regions of nuclear ribosomal RNA. With the addition of
these five new species, the worldwide count of brownish Mycena increases to 94, and a key to the
29 known species of brownish Mycena from China is presented.
Keywords: biodiversity; species complex; new taxon; taxonomy; phylogeny
1. Introduction
Mycena (Pers.) Roussel was established in the early 18th century, standing as one
of the earliest genera among fungi [
1
]. Currently, there are over 500 known species,
with 2418 listed in the Index Fungorum (https://www.indexfungorum.org, accessed on
27 February 2024), highlighting a remarkable species diversity [
2
]. Mycena species are
widely distributed all over the world, and play a crucial role in the decomposition of
dead branches, fallen leaves, and rotting wood, which facilitates the material circulation
of forest ecosystems [
3
–
7
]. Moreover, some species have demonstrated a propensity for
enhancing Orchidaceae seed germination, which contributes to the resilience and diversity
of ecosystems [
8
–
10
]. The diverse morphology and color of Mycena contribute a high
species diversity, while they also present significant taxonomic challenges due to subtle
morphological variations and complex microstructures among species [
11
–
16
]. Taxonomists
J. Fungi 2024,10, 439. https://doi.org/10.3390/jof10060439 https://www.mdpi.com/journal/jof

J. Fungi 2024,10, 439 2 of 36
have proposed several different classification systems, reflecting a varied emphasis on
different feature prioritizations [
11
,
12
,
17
–
20
]. Presently, the classification system proposed
by Maas Geesteranus (1992a, 1992b) has garnered considerable consensus, which divides
Mycena into 44 sections and considers the color of the pileus and stipe as a primary grouping
criterion, for example, sect. Oregonenses Maas Geest., sect. Cyanocephalae Singer, etc. [
11
,
12
].
Brownish Mycena, characterized by a yellowish-brown, grayish-brown to dark brown
pileus or stipe, predominantly inhabit the northern temperate zone, with fewer occurrences
in tropical and subtropical areas [
11
,
12
,
20
–
27
]. They are categorized into at least nine
sections, namely: sect. Amictae A.H. Sm. ex Mass Geest., sect. Basipedes (Fr.) Quél., sect.
Cinerellae Singer ex Maas Geest., sect. Exornatae Maas Geest., sect. Filipedes (Fr.) Quél.,
sect. Fragilipedes (Fr.) Quél., sect. Hiemalis Konrad & Maubl., sect. Mycena (Pers.) Roussel,
and sect. Rubromarginatae Singer ex Maas Geest. [
17
,
21
,
23
,
26
,
28
–
30
]. Among them, sect.
Mycena and sect. Fragilipedes stand out due to substantial species diversity and typical
morphological characteristics. The former is characterized by its light brown, yellowish-
brown to sepia pileus, with both the cystidia and pileipellis covered with excrescences. M.
galericulata (Scop.) Grayis, serving as the type species in genus Mycena and as well as in the
type sect. Mycena, is a prime example of brownish Mycena’s characteristic traits. The other
section, Fragilipedes, is distinguished by its grayish-brown to dark brown pileus, along with
smooth cystidia and a pileipellis typically covered with excrescences [11,12,21].
In recent years, the species diversity of Mycena has received heightened attention,
leading to the publication of many new species, and marking substantial advances in
research [13–15,26,27,30–32]. However, brownish Mycena have remained relatively under-
explored, with their phylogenetic relationships still unclear and a considerable number of
species unknown. Notably, fewer than 30 brownish Mycena species have been identified
in Asia, with a mere 24 documented in China, compared to the 75 cataloged in Europe
and North America [
17
,
20
–
23
,
28
–
30
]. The considerable diversity, similar color, and small
basidiomata present significant challenges in studying morphological structures, leading
to a dominant reason for underestimating the species diversity of brownish Mycena in
China and across Asia [
11
–
13
,
16
,
23
]. At the same time, the abundant types of cystidia and
pileipellis hyphae found in brownish Mycena reflect their diverse shapes and suggest a
complex evolutionary history. This indicates the species’ adaptability to a wide range of
ecological pressures and its ability to thrive in diverse habitats. The diversity of ecological
niches and displayed adaptability are concrete manifestations of its complex evolutionary
background, reflecting the species’ versatility and flexibility in adapting to environmental
changes [21,23].
During our ongoing research and fieldwork aimed at exploring the diversity of Mycena
in China, five novel species were identified among 14 new specimens through comparative
studies with known species. The identification of these novel species was further supported
through comprehensive morphological and anatomical research, alongside molecular
phylogenetic analyses. This work will expand the variety of brownish Mycena in China and
provide fundamental information for the systematic study of this group.
2. Materials and Methods
2.1. Sample Collection and Morphological Description
Specimens used in this study were collected from a broad spectrum of locations across
China, spanning Fujian Province, the Guangxi Zhuang Autonomous Region, Heilongjiang
Province, Jilin Province, Jiangxi Province, Shandong Province, and Zhejiang Province. Dur-
ing fieldwork, meticulous records were maintained, noting location information, collectors,
the collection date, and the ecological habits of each specimen. Simultaneously, the odor
and taste of the fresh specimens were recorded. The specimens’ photos were taken using
a Canon EOS 90D digital camera (Canon, Tokyo, Japan) equipped with an EF-S60 mm
f/2.8 Macro USM (Canon, Tokyo, Japan) and an Olympus E-M1 Mark III camera (Olympus,
Tokyo, Japan), coupled with an M. Zuiko Digital Ed 12–40 mm or 60 mm lens (Olympus,
Tokyo, Japan). Macro-morphological features, encompassing basidiomata size, color, shape,

J. Fungi 2024,10, 439 3 of 36
and surface characteristics were detailed in records based on photographs and field notes.
The nomenclature of color descriptions refers to Ridgway (1912) [
33
]. The extent of pileus
marginal striations or sulcations was quantitatively assessed by the ratio of the striation’s or
sulcation’s length to the pileus’s radius (n R) [
34
]. The specimens were dried in an electric
oven (Stöckli, A. & J. Stöckli AG, Netstal, Switzerland) at 40
◦
C and were then placed in a
ziplock bag with allochroic silica gel for preservation. All specimens are now preserved in
the Fungarium of the Fujian Academy of Agricultural Sciences (FFAAS), China.
Microscopical features were examined with a Lab A1 microscope (Carl Zeiss AG,
Jena, Germany) using dried specimens rehydrated in a 5% KOH aqueous solution and
stained with a 1% Congo red aqueous solution when necessary. Melzer’s reagent was
used to test the amyloidity of basidiospores and lamellar trama [
35
]. Basidiospores were
photographed using ZEN 2.3 (Blue Edition) software (Carl Zeiss Microscopy GmbH, Jena,
Germany), with at least 20 basidiospores measured per specimen. For the holotype, a more
extensive examination entailed measuring 40 or more basidiospores from each basidiomata.
The measured results were presented in the form [a/b/c] (d)e–f–g(h)
×
(i)j–k–l(m)
µ
m
[
Q=(n)o–p
(q), Q
m
=r
±
s]. The abbreviation [a/b/c] represents anumber of basidiospores
measured from bnumber of basidiomata of cnumber of collections; dand hrepresent
the minimum and maximum lengths (5% extremum), while the range e–gencompasses
the remaining 90% of the measured values, with fas the mean length. Width (i–m) and
Q values (n–q) are analogous representations. If the difference between dand e,gand
h,iand j, or land mis less than 0.2
µ
m, and the difference between nand oor pand
qis less than 0.01
µ
m, d,h,i,m,n, or qwere omitted accordingly. Q represents the
“length/width” ratio of a basidiospore. Q
m
presents the average Q of all basidiospores
±
the sample standard deviation [
36
]. Other microstructures examined include basidia,
cheilocystidia, pleurocystidia, pileocystidia, caulocystidia, and the respective compositions
of the pileipellis and stipipellis, along with lamellar trama. For each specimen, 20 of these
structures were recorded, detailing their sizes, shapes, contents, and other features. The
pileipellis samples were specifically cut from the mid-zone between the center and the
margin of the pileus, while stipitipellis samples were procured from the middle of the stipe.
The delineation of macro- and microstructures commenced with manual sketches on
A4 paper using a pencil and subsequently transposed to tracing paper using a Sakura needle
pen (Sakura Color Products Corporation, Osaka, Japan). The sketches were converted
to digitalization in TIF format using a Canon LiDE120 scanner (Canon, Tokyo, Japan).
Photoshop was then utilized for further refinement and typesetting.
2.2. DNA Extraction, PCR Amplification, and Sequencing
DNA extraction from the dried specimens utilized the NuClean Plant Genomic DNA
kit (Kangwei Century Biotechnology Co., Beijing, China). PCR amplification followed in a
25
µ
L reaction mix, comprising 12.5
µ
L 2
×
Utaq PCR MasterMix (ZomanBio, Beijing, China),
8.5
µ
L ddH
2
O, 1
µ
L each of forward and reverse primers, and 2
µ
L DNA template [
14
].
To amplify the nuclear ITS rDNA region (ITS1–5.8S–ITS2), the primer pairs ITS1/ITS4
were employed with the following PCR thermocycling protocol: 94
◦
C for 4 min, then
34 cycles of 94
◦
C for 45 s, 52
◦
C for 45 s, and 72
◦
C for 1 min, with a final extension of
72
◦
C for 10 min [
37
,
38
]. The large subunit (LSU) of rDNA was amplified using LR0R/LR7
primer pairs with the PCR cycle set to 95
◦
C for 5 min; 30 cycles of 95
◦
C for 30 s, 55
◦
C
for 30 s, and 72
◦
C for 1 min; followed by 72
◦
C for 10 min [
16
,
39
]. PCR amplification
products underwent agarose gel electrophoresis for validation before the Sanger dideoxy
was sequenced by the Beijing Genomics Institute (Beijing, China) [
40
]. The sequence quality
was checked using BioEdit 7.0.9.0, with chromatograms inspected for accuracy [
41
,
42
].
Low-quality sequences were enhanced by applying plasmid vectors using VELUTE Gel
Mini Purification and pBLUE-T kits (Beijing Zoman Biotechnology Co., Beijing, China) for
optimal sequencing results [
43
]. All newly obtained sequences in this study were submitted
to GenBank.

J. Fungi 2024,10, 439 4 of 36
2.3. Molecular Phylogeny
The newly obtained sequences were subject to BLAST searches within the NCBI
GenBank database (https://www.ncbi.nlm.nih.gov/, accessed on 27 March 2024) to retrieve
and download homologous sequences (with nucleotide identities >90%). Furthermore,
sequences of species bearing a morphological resemblance to the species in this study were
also sourced from GenBank (https://www.ncbi.nlm.nih.gov/genbank/, accessed on 27
March 2024), selecting one or more representatives per species that share a morphological
or phylogenetic affinity with the newly described species for preliminary phylogenetic
analysis. M. tenuispinosa J. Favre and M. mucoroides Aronsen were used as outgroup
taxa to the root tree. Gaps were treated as missing data after aligning the ITS and LSU
datasets independently using the auto strategy (FFT-NS-1, FFT-NS-2, FFT-NS-i or L-INS-i)
by MAFFT v.7.110, with gene fragments manually refined and amalgamated in MEGA
5 [
44
–
46
]. Both Bayesian Inference (BI) and Maximum Likelihood (ML) were utilized for
phylogenetic inference. For BI analysis, the optimal nucleotide evolution was estimated
using Modeltest 2.3 with the BIC criterion [
47
]. MrBayes 3.2.6 executed the analysis over
5,000,000 MCMC generations; four chains were conducted with the sampling every 500th
generation. The first 25% of trees were discarded as burn-in after the average standard
deviation of split frequencies under 0.01, with results aggregated through the “sump”
and “sumt” commands [
48
]. Tracer v.1.7.2 served in the visualization and examination of
MCMC trace files from the Bayesian phylogenetic inference [
49
]. Meanwhile, ML analysis
was conducted by RAxMLGUI 2.0 with the GTRGAMMA model and 1000 bootstrap (BS)
replicates using default parameters [
50
]. Finally, the phylogenetic tree was visualized in
FigTree v.1.4.3 and refined in Photoshop CS4.
3. Results
3.1. Phylogenetic Analysis
The molecular analyses dataset consisted of 152 sequences in total, including 28 newly
obtained sequences (14 spanning ITS and LSU) and 124 sequences downloaded from
GenBank (98 ITS and 26 LSU). Detailed information is provided in Table 1. The aligned
dataset culminated in a total of 1743 aligned sites, including gaps (824 ITS, 919 LSU).
The average deviation of split frequencies was 0.006, the ESS (effective sample size) was
1908.3, and the average Potential Scale Reduction Factor (PSRF) parameter
values = 1.000
after 5,000,000 MCMC generations. The ML phylogenetic analysis yielded a final log-
likelihood value of
−
13,761.724187. The average bootstrap value was 78.5%, indicating a
generally high confidence in the tree’s branches. The lowest bootstrap value was 50.0%,
suggesting some branches may require further verification, while the highest value was
99.0%, reflecting very high confidence in certain branches. The tree topology effectively
depicted the evolutionary relationships among the major taxa, with samples from each
novel species clustering together as expected.
Table 1. DNA sequences of Mycena used in the phylogenetic analysis in this study.
GenBank NO.
Species Voucher/Strain NO. Locality ITS LSU Reference
M. abramsii HMJAU43606 China MH396629 MK629355 Direct sub.
M. abramsii HMJAU43282 China MH396626 MK629348 Direct sub.
M. aetites CBS 221.47 Netherlands MH856225 MH867752 [51]
M. aetites CBS 222.47 Netherlands MH856226 MH867753 [51]
M. aff. discobasis DED 8211 (SFSU) USA MH414555 MH385331 [52]
M. aff. discobasis BAP 658 (SFSU) USA MH414554 MH385330 [52]

J. Fungi 2024,10, 439 5 of 36
Table 1. Cont.
GenBank NO.
Species Voucher/Strain NO. Locality ITS LSU Reference
M. aff. murina D1.1c Italy KJ093496 — [53]
M. albiceps RA705-6 USA MK234177 — [54]
M. albiceps MGW1504 USA KY744173 MF797661 Direct sub.
M. algeriensis HMJAU43798 China MK733295 MK722347 Direct sub.
M. algeriensis HMJAU59791 China OR468696 — Direct sub.
M. amicta CBS:257.53 Netherlands MH857184 MH868722 [51]
M. atkinsoniana S.D. Russell
iNaturalist # 8545962
USA MN906081 — Direct sub.
M. atkinsoniana S.D. Russell
MycoMap # 7612 USA MN906082 — Direct sub.
M. bulliformis MF74930 USA MT636954 — Direct sub.
M. bulliformis JLF9168 USA MZ007503 — Direct sub.
M. bulliformis SFSU:BAP 547 USA KX513844 KX513848 [55]
M. campanulati-
hemisphaerica
FFAAS1047
Holotype China PP706092 PP704692 This study
M. campanulati-
hemisphaerica FFAAS1048 China PP706093 PP704693 This study
M. campanulati-
hemisphaerica FFAAS1049 China PP706094 PP704694 This study
M. campanulati-
hemisphaerica FFAAS1050 China PP706095 PP704695 This study
M. chlorophos CT15101401 China MH400938 — Direct sub.
M. chlorophos CT151014 China MH400939 — Direct sub.
M. cinerella O73656 Netherlands GU234146 — [56]
M. cinerella H6018219 Hungary MW540670 — Direct sub.
M. citrinomarginata SHXG China OM228755 OM228763 Direct sub.
M. citrinomarginata
S.D. Russell
iNaturalist #
17474783
USA ON416970 — Direct sub.
M. clavicularis HMJAU43611 China ON791480 ON791547 Direct sub.
M. coralliformis ACL306 Malaysia — KJ206962 [57]
M. cristinae JS347 Brazil MT921381 MT921384 [26]
M. digitifurcata FFAAS1054 China PP706099 PP704699 This study
M. digitifurcata FFAAS1055
Holotype China PP706100 PP704700 This study
M. limitis FFAAS1056 China PP706101 PP704701 This study
M. limitis FFAAS1057 China PP706102 PP704702 This study
M. limitis FFAAS1058
Holotype China PP706103 PP704703 This study
M. fulgoris ACP1690 Malaysia MG926694 — [58]
M. fulgoris
XAL A. Cortes-Perez
1690 Mexico NR_163300 — [58]
M. galericulata HMJAU43035 China MW222635 — Direct sub.
M. galericulata HMJAU43845 China MW222634 — Direct sub.
M. galopus 461E Poland MZ078480 — [59]
M. galopus 69 Norway MW576941 — [60]
M. globulispora ACP1765 Mexico MG926696 — [58]
M. globulispora ACP1704 Mexico MG926697 — [58]
M. haematopus 1055 Canada KJ705181 — Direct sub.
M. haematopus BIOUG24046-B04 Canada KT695316 — [61]
M. inclinata Champ-36 Spain KX449443 — [62]
M. inclinata AH56011 Spain ON113881 — [63]

J. Fungi 2024,10, 439 6 of 36
Table 1. Cont.
GenBank NO.
Species Voucher/Strain NO. Locality ITS LSU Reference
M. interrupta HMJAU43791 China MK733300 — Direct sub.
M. interrupta HMJAU43849 China MK733301 — Direct sub.
M. jingyinga LE-BIN 4556 Russia OP997504 — Direct sub.
M. jingyinga OMDL00049 USA OR572512 — Direct sub.
M. kunyuensis FFAAS1045
Holotype China PP706090 PP704690 This study
M. kunyuensis FFAAS1046 China PP706091 PP704691 This study
M. laevigata HMJAU43187 China MK733302 — Direct sub.
M. laevigata HMJAU43618 China MK733304 — Direct sub.
M. laevigata HMJAU43604 China MK733303 MK722354 Direct sub.
M. maculata CBS:235.47 Netherlands MH856231 — [51]
M. maculata 140 m Norway MW576905 — [60]
M. maculata CBS:239.47 Netherlands MH856234 MH867763 [51]
M. megaspora H6036833 Hungary MW540680 — Direct sub.
M. megaspora 5702 Russia MZ754456 — Direct sub.
M. meliigena NSK 1014985 Russia OQ216533 — Direct sub.
M. meliigena
S.D. Russell ONT
iNaturalist #
98315803
USA OP455719 — Direct sub.
M. metata HAY-F-003966 USA OR858777 — Direct sub.
M. metata HAY-F-005148 USA OR882691 — Direct sub.
M. mucoroides OF76006 Sweden NR176117 — Direct sub.
M. mucoroides AAronsen9-060913 Sweden KU861561 — [28]
M. nebula ACP1659 Mexico MG926685 — [58]
M. niveipes DA-18015 France OM368073 — Direct sub.
M. overholtsii iNAT:11409406 USA MZ146337 — Direct sub.
M. overholtsii
S.D. Russell
iNaturalist #
37566563
USA ON561746 — Direct sub.
M. oryzifluens FFAAS1051
Holotype China PP706096 PP704696 This study
M. oryzifluens FFAAS1052 China PP706097 PP704697 This study
M. oryzifluens FFAAS1053 China PP706098 PP704698 This study
M. pasvikensis AAronsen86-12 Sweden KU861556 — [28]
M. pasvikensis AAronsen45-13 Sweden KU861557 — [28]
M. picta
CAFUNDIS
iNaturalist
171114596
USA OR858681 — Direct sub.
M. picta TUR194167 Hungary MW540717 — Direct sub.
M. plumbea AFTOL-ID 1631 USA DQ494677 DQ470813 [64]
M. pluteoides HMJAU43771 China MK733307 MK722357 Direct sub.
M. polycystidiata FFAAS0418 China ON427732 — [14]
M. polycystidiata FFAAS0421 China ON427733 — [14]
M. polygramma CBS 243.47 Netherlands MH856238 MH867767 [51]
M. pura IS10/11/2000 Denmark FN394611 — [65]
M. renati CBS:358.50 Netherlands MH856658 MH868174 [51]
M. romagnesiana 388a USA JF908421 — Direct sub.
M. romagnesiana 388f USA JF908422 — Direct sub.
M. rubromarginata UBC F16259 Canada EF530939 — Direct sub.
M. rubromarginata CBS:265.48 Netherlands MH856335 MH867890 [51]
M. rubromarginata CBS:268.48 Netherlands MH856338 MH867891 [51]

J. Fungi 2024,10, 439 7 of 36
Table 1. Cont.
GenBank NO.
Species Voucher/Strain NO. Locality ITS LSU Reference
M. rufobrunnea FFAAS0414 China ON427728 — [14]
M. rufobrunnea FFAAS0415 China ON427729 — [14]
M. sanguinolenta CBS:367.50 Netherlands MH856662 — [51]
M. sanguinolenta KUBOT-KRMK-
2020-57 India MW446185 MW446189 Direct sub.
M. semivestipes HMJAU43825 China MK733308 MK722358 Direct sub.
M. semivestipes HMJAU43830 China MK733309 — Direct sub.
M. seynesii 71l USA JF908469 — Direct sub.
M. seynesii 71h USA JF908470 — Direct sub.
M. shengshanensis FFAAS0424 China ON427739 — [14]
M. shengshanensis FFAAS0425 China ON427740 — [14]
M. silvae-nigrae HMJAU43815 China MK733310 MK722359 Direct sub.
M. sinar ACL092 Malaysia KF537247 — [66]
M. sinar ACL135 Malaysia KF537249 — [66]
M. subulata FFAAS0419 China ON427735 — [14]
M. subulata FFAAS0423 China ON427737 — [14]
M. tenuispinosa LE 321750 Russia MK478466 — Direct sub.
M. tintinnabulum H6008524 Hungary MW540664 — Direct sub.
M. tintinnabulum NSK 1017255 Russia OR242685 — Direct sub.
M. venus CT20160127 China MG324368 — [30]
M. venus CT20160218 China MG324369 — [30]
M. yuezhuoi FFAAS0347 China MW581493 — [15]
M. yuezhuoi FFAAS0344 China MW581490 — [15]
M. zephirus CBS:270.48 Netherlands MH856339 MH867892 [51]
Remarks: New generated sequences are emphasized in bold; “—” show missing sequence.
The phylogenetic evaluations using ML and BI indicated generally similar topologies
for the five newly identified species, although some discrepancies between the methods
were observed. Consequently, the BI tree was selected to be shown in Figure 1, incor-
porating only those nodes supported by bootstrap values of at least 75% and posterior
probabilities exceeding 0.95. The phylogenetic structure has resolved into 13 distinct clades,
each delineated by shared morphological characteristics of pileipellis hyphae or cheilo-
cystidia among the species within them. While some clades received lower support rates,
the topology of the phylogenetic tree still effectively reflects the shared characteristics of
groups within these clades, such as smooth or ornamented characteristics.
The five newly identified species demonstrated high support within the phylogenetic
tree, with each being robustly placed within clades 1, 2, 9, and 10. These results reinforce
the taxonomic standing of these clades and confirm the systematic placement of these
new species.

J. Fungi 2024,10, 439 8 of 36
J. Fungi 2024, 10, x FOR PEER REVIEW 9 of 37
Figure 1. Bayesian Inference tree based on concatenated ITS + LSU dataset. Only branch nodes with
both Maximum Likelihood bootstrap support values above 75% and Bayesian posterior probabili-
ties exceeding 0.95 are indicated. Red dots and text represent new taxa, black dots indicate the pres-
ence of both ITS and LSU sequences, and white dots signify the presence of only LSU sequences.
Figure 1. Bayesian Inference tree based on concatenated ITS + LSU dataset. Only branch nodes with
both Maximum Likelihood bootstrap support values above 75% and Bayesian posterior probabilities
exceeding 0.95 are indicated. Red dots and text represent new taxa, black dots indicate the presence
of both ITS and LSU sequences, and white dots signify the presence of only LSU sequences.
Clade 1 emerges as a monophyletic clade, incorporating M. oryzifluens sp. nov. sup-
ported by three specimens (FFAAS1051, FFAAS1052, and FFAAS1053). In the BI phyloge-

J. Fungi 2024,10, 439 9 of 36
netic reconstruction, M. oryzifluens sp. nov. clusters closely with other members of Clade 1,
indicating a distinct genetic lineage. Although in the ML analysis, M. oryzifluens sp. nov.
and M. amicta (Fr.) Quél. span a broad clade from Clade 2 to Clade 13, their monophyletic
status is firmly supported (BS/BPP = 100/1.00). Additionally, species in Clade 1 share
the characteristic of non-smooth pileipellis hyphae. Clade 2 incorporates the new species
M. digitifurcata sp. nov. (FFAAS1054, FFAAS1055), all members sharing the feature of
non-smooth pileipellis hyphae. In the BI tree, M. digitifurcata sp. nov. is closely related
to M. cristinae, although this relationship is not as strong in the ML analysis. Despite
variations between the BI and ML trees, M. digitifurcata sp. nov. is consistently recognized
as a monophyletic entity with high statistical support (BS/BPP = 99/1.00). Species of Clade
3 exhibit smooth pileipellis and cheilocystidia, receiving unanimous support (100%) and
demonstrating high phylogenetic consistency. Clade 4, although supported at a lower rate
(BS/BPP = 81/1.00), presents species with cylindrical excrescences on the cheilocystidia,
providing key taxonomic insights. Clades 5 and 6 both exhibit very high support rates
(BS/BPP = 100/1.00), where Clade 5 species feature smooth cheilocystidia, while Clade 6
species display excrescences on the cheilocystidia. Clades 7 and 8 also showcase specific
morphological characteristics, with Clade 7 having non-smooth pileipellis and Clade 8
featuring non-smooth pileipellis, along with some species having gelatinous layers. Clade
9 includes M. limitis sp. nov. (FFAAS1056, FFAAS1057, and FFAAS1058), sharing a close
phylogenetic relationship with M. niveipes (Murrill) Murrill, along with other species such
as M. seynii Quél., M. bulliformis B.A. Perry & Desjardin, and M. fulgoris Cortés-Pérez
& Desjardin. Although Clades 9 and 12 form independent groups in BI analysis, Clade
12 nests within Clade 9 in ML analysis. Nonetheless, the position of M. limitis sp. nov.
remains well-supported (BS/BPP = 100/1.00). Clade 10 comprises the novel species M.
campanulatihemisphaerica sp. nov. (FFAAS1047, FFAAS1048, and FFAAS1049) and M. kun-
yuensis sp. nov. (FFAAS1045, FFAAS1046), sharing characteristics of smooth cheilocystidia
with other species. Clade 11, though not highly supported, includes species that exhibit
smooth cheilocystidia. Clade 12 is fully supported (100%), with species displaying smooth
cheilocystidia and non-smooth pileipellis, showing clear phylogenetic consistency. Clade
13, similarly supported fully (100%) in the BI tree, shares the characteristic of non-smooth
pileipellis among its species.
3.2. Taxonomy
Mycena campanulatihemisphaerica R.X. Wei, X.M. Wu, J. Yu, Y.P. Ge & Q. Na, sp. nov.,
Figures 2–4.
MycoBank no: 853739
Etymology: The epithet campanulatihemisphaerica combines the Latin feminine adjec-
tives campanulata (bell-shaped) and hemisphaerica (half-spherical). This name accurately
describes the shape of the pileus, which transitions from bell-shaped to nearly hemispherical.
Holotype: CHINA. Shandong Province, Kunyushan National Nature Reserve, Yantai
City, 19 July 2019, leg. Renxiu Wei, Liming Xue, Ruichen Liu, Qin Na, and Yupeng Ge,
516 m asl, FFAAS1047 (collection no. MY022).
Diagnosis: Pileus fuscous brown, obviously sulcate, basidiospores narrowly ellipsoid
to cylindrical, cheilocystidia ovate, ellipsoid, fusiform to sublageniform, some slightly
tapered or forked apices, pileipellis and stipitipellis covered with cylindrical excrescences.
Differs from M. venus by unbranched cheilocystidia and larger basidiospores.
Description: Pileus 3–12 mm diam., ovoid, hemispherical to campanulate, sometimes
apex with an inconspicuous umbo, margin smooth; *Hair Brown (XLVI17””i) at center,
paler towards the margin to Pale Drab-Gray (XLVI17””f), Pale Smoke Gray (XLVI21””f)
to White (LIII); pruinose at center, obviously sulcate towards the center up to 0.5–0.8 R,
Light Drab (XLVI17””b) to *Hair Brown (XLVI17””i), dry. Context White (LIII), fragile, thin.
Lamellae emarginate to sinuate, 0.2–0.6 mm wide, L= 12–16, l= 1–2, White (LIII), edge
concolorous with face. Stipe 19–60
×
0.9–2.0 mm, central, cylindrical, hollow, fragile, *Hair

J. Fungi 2024,10, 439 10 of 36
brown (XLVI17””i) at the base, paler toward the apex to White (LIII), finely pubescent, base
covered with short, dense, White (LIII) fibrils. Odor and taste indistinctive.
J. Fungi 2024, 10, x FOR PEER REVIEW 11 of 37
Figure 2. Basidiomata of Mycena campanulatihemisphaerica sp. nov. (a,d–i) FFAAS1047 (holotype). (b)
FFAAS1049; (c) FFAAS1050; (f) Pileus striate-sulcate; (g) pubescence on stipe; (h) lamellae; (i) fibrils
length in base of stipe. Bars: (a–e) = 10 mm; (f–i) = 5 mm. Photos by Qin Na.
Figure 2. Basidiomata of Mycena campanulatihemisphaerica sp. nov. (a,d–i)FFAAS1047 (holotype).
(b)FFAAS1049; (c)FFAAS1050; (f) Pileus striate-sulcate; (g) pubescence on stipe; (h) lamellae; (i) fibrils
length in base of stipe. Bars: (a–e) = 10 mm; (f–i) = 5 mm. Photos by Qin Na.

J. Fungi 2024,10, 439 11 of 36
J. Fungi 2024, 10, x FOR PEER REVIEW 12 of 37
Figure 3. Microscopical features of Mycena campanulatihemisphaerica (FFAAS1047, holotype). (a–e)
Basidiospores; (f) basidium; (g–r) cheilocystidia; (s) pileipellis and upper part of pileipellis hypha
with cylindrical excrescences; (t) hymenium and lamellar trama; (u) stipitipellis and stipitipellis hy-
pha with cylindrical excrescences. Bars: (a–e) = 5 µm; (f–u) = 10 µm. Structures (a–e) were rehydrated
in 5% KOH aqueous solution, and (f–u) were stained in 1% Congo red aqueous solution.
Figure 3. Microscopical features of Mycena campanulatihemisphaerica (FFAAS1047, holotype).
(a–e) Basidiospores
; (f) basidium; (g–r) cheilocystidia; (s) pileipellis and upper part of pileipellis
hypha with cylindrical excrescences; (t) hymenium and lamellar trama; (u) stipitipellis and stipitipel-
lis hypha with cylindrical excrescences. Bars: (a–e) = 5
µ
m; (f–u) = 10
µ
m. Structures (a–e) were
rehydrated in 5% KOH aqueous solution, and (f–u) were stained in 1% Congo red aqueous solution.

J. Fungi 2024,10, 439 12 of 36
J. Fungi 2024, 10, x FOR PEER REVIEW 13 of 37
Figure 4. Morphological features of Mycena campanulatihemisphaerica (FFAAS1047, holotype). (a) Ba-
sidiomata; (b) basidia; (c) basidiospores; (d) cheilocystidia; (e) stipitipellis; (f) pileipellis. Bars: (a) =
10 mm; (b–f) = 10 µm. Drawing by Renxiu Wei.
Habit and habitat: Fascicled or scaered on roen branches in mixed broadleaf–co-
nifer forests, mainly under trees of Picea, Pinus, and Quercus.
Known distribution: Fujian Province, Jiangxi Province, Shandong Province, China.
Additional material examined: China. Fujian Province, Junzifeng National Nature
Conservation Area, Mingxi County, Sanming City, 2 May 2021, leg. Renxiu Wei, Binrong
Ke, Hui Zeng, Junqing Yan, Qin Na, and Yupeng Ge, 824 m asl, FFAAS1049 (collection no.
MY0267); Jiangxi Province, Jiangxi Agricultural University, Nanchang City, 22 April 2021,
Figure 4. Morphological features of Mycena campanulatihemisphaerica (FFAAS1047, holotype). (a) Ba-
sidiomata; (b) basidia; (c) basidiospores; (d) cheilocystidia; (e) stipitipellis; (f) pileipellis. Bars:
(a) = 10 mm; (b–f) = 10 µm. Drawing by Renxiu Wei.
Basidiospores [107/5/4] (8.5)8.8–10.0–12.1
×
4.5–5.3–6.5
µ
m, [Q =1.74–2.05, Q
m
=
1.87 ±0.09
]
[holotype (40/2/1) 8.8–10.0–12.1
×
4.5–5.3–6.5
µ
m, Q = 1.74–2.05, Q
m
=
1.88 ±0.09
], nar-
rowly ellipsoid to cylindrical, hyaline, smooth, thin-walled, amyloid. Basidia 21–30
×
7–10
µ
m,
four-spored, rarely two-spored, clavate, hyaline, sterigmata 2–4
µ
m in length. Cheilocystidia
18–36
×
9–16
µ
m, ovate, ellipsoid, fusiform to sublageniform, apex 1.5–4.2
×
1.2–2.4
µ
m,

J. Fungi 2024,10, 439 13 of 36
tapered, occasionally forked, hyaline, thin-walled. Pleurocystidia absent. Pileipellis a cutis
composed of two layers of hyphae, upper part of hyphae 2.0–5.2
µ
m diam., parallel, hyaline,
thin-walled, covered with cylindrical excrescences, 2.0–7.0
×
1.0–2.4
µ
m; lower part of
hyphae 4.3–6.8
µ
m diam., interwoven, hyaline, thin-walled, smooth; pileocystidia absent.
Lamellae trama subcellular, composed of subglobose, ellipsoid, and narrowly ellipsoid cells,
23–43
×
17–25
µ
m, hyaline, dextrinoid. Stipitipellis a cutis composed of a hypha,
1.9–4.8 µm
diam., hyaline, thin-walled, covered with cylindrical excrescences, 1.7–4.1
×
0.6–1.6
µ
m;
caulocystidia absent. Clamps present in all tissues, but rarely observed in context.
Habit and habitat: Fascicled or scattered on rotten branches in mixed broadleaf–conifer
forests, mainly under trees of Picea,Pinus, and Quercus.
Known distribution: Fujian Province, Jiangxi Province, Shandong Province, China.
Additional material examined: China. Fujian Province, Junzifeng National Nature
Conservation Area, Mingxi County, Sanming City, 2 May 2021, leg. Renxiu Wei, Binrong
Ke, Hui Zeng, Junqing Yan, Qin Na, and Yupeng Ge, 824 m asl, FFAAS1049 (collection no.
MY0267); Jiangxi Province, Jiangxi Agricultural University, Nanchang City, 22 April 2021,
leg. Renxiu Wei, Junqing Yan, and Qin Na, 59 m asl, FFAAS1048 (collection no. MY056);
Shandong Province, Mengshan Scenic Area, Mengyin County, Linyi City, 20 July 2021,
leg. Renxiu Wei, Yulan Sun, Zewei Liu, Qin Na, and Yupeng Ge, 962 m asl, FFAAS1050
(collection no. MY0389).
Notes: M. venus exhibits morphological affinities with M. campanulatihemisphaerica, par-
ticularly in the characteristics of the pileus and stipe. However, M. venus is distinguished by
its branched cheilocystidia and notably smaller basidiospores, measuring (4.8) 6.8–7.7 (8.7)
×
(3.2) 3.8–4.8 (5.4)
µ
m, with a Q
m
of 1.69 [
30
]. M. silvae-nigrae Maas Geest. & Schwöbel and
M. leptocephala (Pers.) Gillet share similarities with M. campanulatihemisphaerica in pileus
color. Nonetheless, M. silvae-nigrae is set apart by its uniformly dark brown stipe and the
presence of pleurocystidia [
21
,
23
,
28
]; M. leptocephala is differentiated by presenting caulocys-
tidia and pleurocystidia [
20
,
21
,
23
,
28
,
29
]. Microscopically, M. abramsii (Murrill) Murrill and
M. scirpicola M. Villarreal, Heykoop, Esteve-Rav. & Maas Geest. also display cheilocystidia
of a similar morphology, such as clavate, fusiform, and sublageniform to cylindrical, but M.
campanulatihemisphaerica is distinguishable from these species by its fuscous brown pileus
and the absence of both pleurocystidia and
caulocystidia [20,21,23,28,29,67,68]
. Addition-
ally, some cheilocystidia were observed with characteristics such as slightly tapered or
forked apices, resembling subglobose and subclavate cells, suggesting they are in an early
phase of cheilocystidia.
Mycena kunyuensis R.X. Wei, X.M. Wu, M.Z. Chen, Y.P. Ge & Q. Na, sp. nov.,
Figures 5–7
.
MycoBank no: 853782
Etymology: The epithet kunyuensis refers to the Kunyu mountain where the holotype
was discovered.
Holotype: CHINA. Shandong Province, Kunyushan National Nature Reserve, Yantai
City, 19 July 2019, leg. Renxiu Wei, Liming Xue, Ruichen Liu, Qin Na, and Yupeng Ge,
513 m asl, FFAAS1045 (collection no. MY021).
Diagnosis: Pileus pale olive-buff to olive-buff, basidiospores ellipsoid to narrowly
ellipsoid, cheilocystidia lageniform, obpyriform, obclavate to fusiform, with a narrow neck,
apex tapered, pileipellis and stipitipellis covered with cylindrical excrescences. Differs
from M. abramsii by smaller basidiomata and absent pleurocystidia.

J. Fungi 2024,10, 439 14 of 36
J. Fungi 2024, 10, x FOR PEER REVIEW 15 of 37
occasionally forked; pileocystidia absent. Lamellae trama cellular, composed of subglobose,
ellipsoid, and narrowly ellipsoid cells, 24–48 × 21–25 µm, hyaline, dextrinoid. Stipitipellis
a cutis composed of a hypha, (1.7) 2.1–4.6 µm diam., covered with cylindrical excres-
cences, 0.6–10.2 (15.3) × 0.9–2.9 µm, occasionally forked, hyaline, thin-walled; caulocystidia
absent. Clamps present in all tissues, but rarely observed in context.
Figure 5. Basidiomata of Mycena kunyuensis sp. nov. (a–d,f–j) FFAAS1045; (e) FFAAS1046 (holo-
type); (f) The surface of the stipe is pubescent; (g,i) Pileus color and striate-sulcate; (h,j) Fibrils
length in the base of the stipe. Bars: (a–d) = 10 mm; (e–j) = 5 mm. Photos by Qin Na.
Figure 5. Basidiomata of Mycena kunyuensis sp. nov. (a–d,f–j)FFAAS1045; (e)FFAAS1046 (holotype);
(f) The surface of the stipe is pubescent; (g,i) Pileus color and striate-sulcate; (h,j) Fibrils length in the
base of the stipe. Bars: (a–d) = 10 mm; (e–j) = 5 mm. Photos by Qin Na.

J. Fungi 2024,10, 439 15 of 36
J. Fungi 2024, 10, x FOR PEER REVIEW 16 of 37
Figure 6. Microscopical features of Mycena kunyuensis (FFAAS1045, holotype). (a–e) Basidiospores;
(f) basidium; (g–p) cheilocystidia; (q) pileipellis and pileipellis hypha with cylindrical excrescences;
(r) Hymenia and lamellar trama; (s) stipitipellis and stipitipellis hypha with cylindrical excrescences.
Bars: (a–f) = 5 µm; (g–s) = 20 µm. Structures (a–e) were rehydrated in 5% KOH aqueous solution,
and (f–s) were stained in 1% Congo red aqueous solution.
Figure 6. Microscopical features of Mycena kunyuensis (FFAAS1045, holotype). (a–e) Basidiospores;
(f) basidium; (g–p) cheilocystidia; (q) pileipellis and pileipellis hypha with cylindrical excrescences;
(r) Hymenia and lamellar trama; (s) stipitipellis and stipitipellis hypha with cylindrical excrescences.
Bars: (a–f)=5
µ
m; (g–s) = 20
µ
m. Structures (a–e) were rehydrated in 5% KOH aqueous solution,
and (f–s) were stained in 1% Congo red aqueous solution.

J. Fungi 2024,10, 439 16 of 36
J. Fungi 2024, 10, x FOR PEER REVIEW 17 of 37
Figure 7. Morphological features of Mycena kunyuensis (FFAAS1045, holotype). (a) Basidiomata; (b)
basidia; (c) basidiospores; (d) cheilocystidia; (e) stipitipellis; (f) pileipellis and context. Bars: (a) = 10
mm; (b–f) = 10 µm. Drawing by Renxiu Wei.
Habit and habitat: Scaered on roen branches in mixed broadleaf–conifer forests,
mainly under trees of Picea, Pinus, and Quercus.
Known distribution: Shandong Province, China.
Additional material examined: CHINA. Shandong Province, Kunyushan National
Nature Reserve, Yantai City, 19 July 2019, leg. Renxiu Wei, Liming Xue, Ruichen Liu, Qin
Na, and Yupeng Ge, 524 m asl, FFAAS1046 (collection no. MY023).
Notes: M. abramsii is the closest species to M. kunyuensis, sharing numerous macro-
scopic and microscopic characters. Nevertheless, M. abramsii is distinguished by its much
larger basidiomata and the presence of pleurocystidia [20,21,23,28,29,67,68]. In addition,
Figure 7. Morphological features of Mycena kunyuensis (FFAAS1045, holotype). (a) Basidiomata;
(b) basidia; (c) basidiospores; (d) cheilocystidia; (e) stipitipellis; (f) pileipellis and context. Bars:
(a) = 10 mm; (b–f) = 10 µm. Drawing by Renxiu Wei.
Description: Pileus 5–11 mm diam., fusiform to campanulate when young, slightly
campanulate, conical, hemispherical to plano-convex with age, margin slightly serrulate,
occasionally cracked when old; Citrine-Drab (XL21”’i), Deep Quaker Drab (LI1””’i) at
center, paler towards margin to *Olive-Buff (XL21”’d) when young, Citrine-Drab (XL21”’i),
Pale Mouse Gray (LI15””’d) at center, gradually paler towards margin to Pale Olive-
Buff (XL21”’f), Pallid Mouse Gray (LI15””’f) to White (LIII) when matured; initially pru-
inose, when matured glabrescent, undistinguished to clear striate towards the center up

J. Fungi 2024,10, 439 17 of 36
to
0.5–0.6 R
, dry. Context White (LIII), fragile, thin. Lamellae sinuate, 0.3–0.6 mm wide,
L= 14–18
,l= 1, White (LIII), edge concolorous. Stipe 33–46
×
0.8–1.1 mm, central, clavate
to cylindrical, hollow, fragile, in young stages; Deep Quaker Drab (LI1””’i), *Hair Brown
(XLVI17””i), when matured White (LIII) in the upper part, Pale Smoke Gray (XLVI21””f),
Light Grayish Olive (XLVI21””b) towards the base, pruinose when young, then apex to
middle part finely pubescent, almost glabrous in the lower part, base slightly swollen,
densely covered with long White (LIII) fibrils. Odor and taste indistinctive.
Basidiospores [65/3/2] (7.0) 7.4–9.3–11.5 (11.8)
×
4.6–5.8–6.9
µ
m, [Q = (1.40) 1.43–1.77
(1.79), Q
m
= 1.60
±
0.09] [holotype (40/2/1) (7.0) 7.4–8.8–10.4
×
4.6–5.6–6.8
µ
m, Q = (1.40)
1.43–1.77, Q
m
= 1.56
±
0.08], ellipsoid to narrowly ellipsoid, hyaline, smooth (under oil),
thin-walled, amyloid. Basidia 16–23
×
5–9
µ
m, four-spored, rarely two-spored, clavate,
hyaline, sterigmata 2.1–4.2
µ
m in length. Cheilocystidia 28–59
×
8–20
µ
m, lageniform,
obpyriform, obclavate to fusiform, with a narrow neck, apex tapered, smooth, sometimes
neck with several excrescences, and occasionally base constricted to cylindrical, hyaline,
thin-walled. Pleurocystidia absent. Pileipellis a cutis composed of parallel hyphae,
2.5–5.1 µm
diam., thin-walled, densely covered with cylindrical excrescences, 1.6–9.2
×
1.0–2.6
µ
m,
occasionally forked; pileocystidia absent. Lamellae trama cellular, composed of subglobose,
ellipsoid, and narrowly ellipsoid cells, 24–48
×
21–25
µ
m, hyaline, dextrinoid. Stipitipellis a
cutis composed of a hypha, (1.7) 2.1–4.6
µ
m diam., covered with cylindrical excrescences,
0.6–10.2 (15.3)
×
0.9–2.9
µ
m, occasionally forked, hyaline, thin-walled; caulocystidia absent.
Clamps present in all tissues, but rarely observed in context.
Habit and habitat: Scattered on rotten branches in mixed broadleaf–conifer forests,
mainly under trees of Picea,Pinus, and Quercus.
Known distribution: Shandong Province, China.
Additional material examined: CHINA. Shandong Province, Kunyushan National
Nature Reserve, Yantai City, 19 July 2019, leg. Renxiu Wei, Liming Xue, Ruichen Liu, Qin
Na, and Yupeng Ge, 524 m asl, FFAAS1046 (collection no. MY023).
Notes: M. abramsii is the closest species to M. kunyuensis, sharing numerous macro-
scopic and microscopic characters. Nevertheless, M. abramsii is distinguished by its much
larger basidiomata and the presence of pleurocystidia [
20
,
21
,
23
,
28
,
29
,
67
,
68
]. In addition,
taxa such as M. fagetorum (Fr.) Gillet, M. metata (Fr.) P. Kumm. and M. filopes (Bull.) P.
Kumm. exhibit a similar pileus color, but distinct differences are noted: M. fagetorum
is distinguished by its clavate cheilocystidia, which are prominently covered by coarse
excrescences [
28
]; M. metata is differentiated by its clavate and obovoid cheilocystidia,
and presenting pleurocystidia [
21
,
23
,
28
,
29
]; and M. filopes is recognized by its excrescence-
covered cheilocystidia and the presence of caulocystidia [
21
,
23
,
28
,
29
]. M. aetites (Fr.) Quél.
also presents lageniform cheilocystidia, but it is uniquely identified by its pileus, which
ranges from black to dark brown, and its gelatinized pileipellis [
23
,
28
]. Comparatively, M.
kunyuensis and M. campanulatihemisphaerica exhibit similarities in some macroscopic and
microscopic characters. However, M. campanulatihemisphaerica can be distinguished by its
shorter fibrils-covered stipe; fuscous brown, markedly sulcate pileus, with its margin being
slightly serrulate; larger basidiospores; and cheilocystidia that fork at the apex (Figure 8).

J. Fungi 2024,10, 439 18 of 36
J. Fungi 2024, 10, x FOR PEER REVIEW 18 of 37
taxa such as M. fagetorum (Fr.) Gillet, M. metata (Fr.) P. Kumm. and M. filopes (Bull.) P.
Kumm. exhibit a similar pileus color, but distinct differences are noted: M. fagetorum is
distinguished by its clavate cheilocystidia, which are prominently covered by coarse ex-
crescences [28]; M. metata is differentiated by its clavate and obovoid cheilocystidia, and
presenting pleurocystidia [21,23,28,29]; and M. filopes is recognized by its excrescence-cov-
ered cheilocystidia and the presence of caulocystidia [21,23,28,29]. M. aetites (Fr.) Quél.
also presents lageniform cheilocystidia, but it is uniquely identified by its pileus, which
ranges from black to dark brown, and its gelatinized pileipellis [23,28]. Comparatively, M.
kunyuensis and M. campanulatihemisphaerica exhibit similarities in some macroscopic and
microscopic characters. However, M. campanulatihemisphaerica can be distinguished by its
shorter fibrils-covered stipe; fuscous brown, markedly sulcate pileus, with its margin be-
ing slightly serrulate; larger basidiospores; and cheilocystidia that fork at the apex (Figure
8).
Figure 8. Morphological features of Mycena kunyuensis and Mycena campanulatihemisphaerica.
(1)Mycena kunyuensis; (2)Mycena campanulatihemisphaerica. (a) Basidiomata color; (b) pileus striate-
sulcate; (c) fibrils length in base of stipe; (d–i) basidiospores shape; (j–o) cheilocystidia shape. Bars:
(a) = 10 mm; (b,c)=5mm;(d–o)=5µm.
Mycena oryzifluens R.X. Wei, L.L. Qi, Y.P. Ge & Q. Na, sp. nov., Figures 9–11.

J. Fungi 2024,10, 439 19 of 36
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Figure 9. Basidiomata of Mycena oryzifluens sp. nov. (a,c,j) FFAAS1051 (holotype); (g) FFAAS1052;
(b,d–f,h,i) FFAAS1053; (g) Lamellae; (h) The surface of the stipe is pruinose and pubescent; (i) The
surface of the pileus is pruinose and pubescent; (j) Tomentum length in the base of the stipe. Bars:
(a–i) = 2 mm; (j) = 1 mm. Photos by Yupeng Ge.
Figure 9. Basidiomata of Mycena oryzifluens sp. nov. (a,c,j)FFAAS1051 (holotype); (g)FFAAS1052;
(b,d–f,h,i)FFAAS1053; (g) Lamellae; (h) The surface of the stipe is pruinose and pubescent; (i) The
surface of the pileus is pruinose and pubescent; (j) Tomentum length in the base of the stipe. Bars:
(a–i)=2mm;(j) = 1 mm. Photos by Yupeng Ge.

J. Fungi 2024,10, 439 20 of 36
J. Fungi 2024, 10, x FOR PEER REVIEW 21 of 37
Figure 10. Microscopical features of Mycena oryzifluens (FFAAS1051, holotype). (a–f) Basidiospores;
(g,h) basidia; (i–q) cheilocystidia; (r) pileipellis and terminal cells; (s) hymenia and lamellar trama;
(t) stipitipellis and thick-walled caulocystidia. Bars: (a–h) = 5 µm; (i–t) = 20 µm. Structures (a–f) were
rehydrated in 5% KOH aqueous solution, and (g–t) were stained in 1% Congo red aqueous solution.
Figure 10. Microscopical features of Mycena oryzifluens (FFAAS1051, holotype). (a–f) Basidiospores;
(g,h) basidia; (i–q) cheilocystidia; (r) pileipellis and terminal cells; (s) hymenia and lamellar trama;
(t) stipitipellis and thick-walled caulocystidia. Bars: (a–h)=5
µ
m; (i–t) = 20
µ
m. Structures (a–f) were
rehydrated in 5% KOH aqueous solution, and (g–t) were stained in 1% Congo red aqueous solution.

J. Fungi 2024,10, 439 21 of 36
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Figure 11. Morphological features of Mycena oryzifluens (FFAAS1051, holotype). (a) Basidiomata; (b)
basidia; (c) basidiospores; (d) stipitipellis; (e) cheilocystidia; (f) pileipellis and context. Bars: (a) = 5
mm; (b,c) = 10 µm; (d–f) = 20 µm. Drawing by Renxiu Wei.
Basidiospores [96/4/3] 6.0–6.6–7.5 (7.7) × 4.6–5.3–6.3 µm, [Q = 1.10–1.45, Qm = 1.24 ± 0.07]
[holotype (40/2/1) 6.2–6.8–7.4 (7.7) × (4.8)5.0–5.6–6.3 µm, Q = 1.10–1.37, Qm = 1.20 ± 0.05],
subglobose to broadly ellipsoid, hyaline, smooth (under oil), thin-walled, amyloid. Basidia
17–27 × 7–10 µm, two- and four-spored, clavate, hyaline, sterigmata 1.7–4.7 µm in length.
Cheilocystidia 15–36 × 8–18 µm, clavate, ovoid, obpyriform, with one to several finger-like
branches at apex, 1.0–17.1 × 0.6–2.6 µm, sometimes branches with furcate excrescences,
hyaline, thin-walled. Pleurocystidia absent. Pileipellis a cutis composed of parallel hyphae,
Figure 11. Morphological features of Mycena oryzifluens (FFAAS1051, holotype). (a) Basidiomata;
(b) basidia; (c) basidiospores; (d) stipitipellis; (e) cheilocystidia; (f) pileipellis and context. Bars:
(a)=5mm;(b,c) = 10 µm; (d–f) = 20 µm. Drawing by Renxiu Wei.
MycoBank no: 853783
Etymology: The epithet oryzifluens is derived from the Latin word oryzi, meaning
rice, and fluens, meaning flowing. This name reflects the rice-like appearance of the white,
pruinose, and pubescent stipe and connects to the folklore of the type locality, where a
legend speaks of a flowing rice cave, symbolizing abundance and continuity.

J. Fungi 2024,10, 439 22 of 36
Holotype: CHINA. Guangxi Zhuang Autonomous Region, Qingxiushan Scenic Spot,
Nanning City, 14 July 2022, leg. Renxiu Wei, LiangLiang Qi, Liying Li, Yongqiang Hu, and
Yupeng Ge, 164 m asl, FFAAS1051 (collection no. MY0870).
Diagnosis: Pileus plumbeous to gray, finely white pruinose and pubescent, basid-
iospores subglobose to broadly ellipsoid, cheilocystidia clavate, ovoid, obpyriform, with
finger-like branches in the apex, pileipellis with non-smooth terminal cells, caulocystidia
thick-walled. Different from M. cretata Aronsen by clavate cheilocystidia with cylindrical
branches, thick-walled caulocystidia, and absent pleurocystidia.
Description: Pileus 2.1–6.0 mm diam., campanulate when young, then hemispherical
to oblate hemispherical; Dark Plumbeous (LII49””’i) at center, paler towards margin to
Plumbeous (LII49””’b) when young, paler towards margin to French Gray (LII49””’f),
*Lilac Gray (LIII59””’f) to White (LIII) with age; finely White (LIII) pruinose and pubescent,
especially dense White (LIII) pubescence surrounds the margin, sulcate towards the center
up to 0.5–0.6 R, Dark Vinaceous Gray (LII59””’k), Vinaceous Gray(L69””’d) when young,
Light Violet Gray(LII59””’b) to Deep Violet Gray (LII59””’i) when matured, dry. Context
White (LIII), fragile, thin. Lamellae adnate, 0.3–0.5 mm wide, L= 16–25, l= 1–2, White
(LIII), edge serrulate, concolorous with face. Stipe 5–20
×
0.3–1.0 mm, center, cylindrical,
hollow, fragile, Deep Purplish Gray (LIII67””i) when young, then paler to Light Violet-Gray
(LII59””’b), Hathi Gray (LII35””’b) with age, finely White (LIII) pruinose and pubescent,
base densely covered with White (LIII) tomentum. Odor and taste indistinctive.
Basidiospores [96/4/3] 6.0–6.6–7.5 (7.7)
×
4.6–5.3–6.3
µ
m, [Q = 1.10–1.45, Q
m
= 1.24
±
0.07]
[holotype (40/2/1) 6.2–6.8–7.4 (7.7)
×
(4.8)5.0–5.6–6.3
µ
m, Q = 1.10–1.37, Q
m
= 1.20
±
0.05],
subglobose to broadly ellipsoid, hyaline, smooth (under oil), thin-walled, amyloid. Basidia
17–27
×
7–10
µ
m, two- and four-spored, clavate, hyaline, sterigmata 1.7–4.7
µ
m in length.
Cheilocystidia 15–36 ×8–18 µm, clavate, ovoid, obpyriform, with one to several finger-like
branches at apex, 1.0–17.1
×
0.6–2.6
µ
m, sometimes branches with furcate excrescences,
hyaline, thin-walled. Pleurocystidia absent. Pileipellis a cutis composed of parallel hyphae,
2.7–5.2
µ
m diam., hyaline, thin-walled, terminal cells often swollen, 34–103
×
5–15
µ
m,
clavate to cylindrical, with sparse nodulose excrescences, 0.5–3.0
×
0.5–1.0
µ
m; pileocystidia
absent. Lamellae trama cellular, composed of subglobose, ellipsoid and narrowly ellipsoid
cells, 20–50
×
15–30
µ
m, hyaline, dextrinoid. Stipitipellis a cutis composed of a hypha,
2.8–5.7
µ
m diam., hyaline, thin-walled; caulocystidia 44–107
×
4–10
µ
m, cylindrical, with a
narrow protuberance in the apex, thick-walled (0.6–1.7
µ
m thick), smooth, hyaline. Clamps
present in all tissues, but rarely observed in context.
Habit and habitat: Scattered on litter layers and rotten branches in broad-leaved mixed
forests, mainly under trees of Ficus and Parashorea.
Known distribution: Guangxi Zhuang Autonomous Region, China.
Additional material examined: China. Guangxi Zhuang Autonomous Region, Qingx-
iushan Scenic Spot, Nanning City, 14 July 2022, leg. Renxiu Wei, LiangLiang Qi, Liying Li,
Yongqiang Hu, and Yupeng Ge, 185 m asl, FFAAS1052 (collection no. MY0871), 191 m asl,
FFAAS1053 (collection no. MY0872).
Notes: Macroscopically, M. cretata shares a similar pileus color with M. oryzifluens but
is distinguished by the presence of pleurocystidia and thin-walled caulocystidia, with its
cheilocystidia lacking finger-like extensions [
28
]. Microscopically, both M. tallangattensis
Grgur and M. scirpicola feature thick-walled caulocystidia. However, M. tallangattensis
is identified by its thick-walled cheilocystidia and the presence of pleurocystidia [
17
],
while M. scirpicola is noted for its elongated basidiospores and the absence of branched
cheilocystidia [
23
,
28
]. Additionally, taxa M. tristis Maas Geest., M. clavularis (Batsch)
Sacc., M. tenuispinosa, and M. mucor (Batsch) Quél. exhibit cheilocystidia of a similar
morphology, but M. oryzifluens is distinctively characterized by its plumbeous to gray pileus,
basidiospores that are subglobose to broadly ellipsoid, and thick-walled caulocystidia [
28
].
Mycena digitifurcata R.X. Wei, H. Zeng, Y.P. Ge & Q. Na, sp. nov., Figures 12–14.

J. Fungi 2024,10, 439 23 of 36
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Figure 12. Basidiomata of Mycena digitifurcata sp. nov. (a–j) FFAAS1055 (holotype); (f) Tomentum in
the base of the stipe; (g) The surface of the stipe is pruinose and pubescent; (h–j) Lamellae. Bars: (a–
j) = 5 mm. Photos by Qin Na.
Figure 12. Basidiomata of Mycena digitifurcata sp. nov. (a–j)FFAAS1055 (holotype); (f) Tomentum
in the base of the stipe; (g) The surface of the stipe is pruinose and pubescent; (h–j) Lamellae. Bars:
(a–j) = 5 mm. Photos by Qin Na.

J. Fungi 2024,10, 439 24 of 36
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Figure 13. Microscopical features of Mycena digitifurcata (FFAAS1055, holotype). (a–e) Basidiospores;
(f) basidium; (g–r) cheilocystidia; (s) pileipellis and pileipellis hypha with cylindrical excrescences;
(t) hymenia and lamellar trama; (u) stipitipellis and stipitipellis hypha with cylindrical excrescences.
Bars: (a–e) = 5 µm; (f–r) = 10 µm; (s–u) = 20 µm. Structures (a–e) were rehydrated in 5% KOH aque-
ous solution, and (f–u) were stained in 1% Congo red aqueous solution.
Figure 13. Microscopical features of Mycena digitifurcata (FFAAS1055, holotype). (a–e) Basidiospores;
(f) basidium; (g–r) cheilocystidia; (s) pileipellis and pileipellis hypha with cylindrical excrescences;
(t) hymenia and lamellar trama; (u) stipitipellis and stipitipellis hypha with cylindrical excrescences.
Bars: (a–e)=5
µ
m; (f–r) = 10
µ
m; (s–u) = 20
µ
m. Structures (a–e) were rehydrated in 5% KOH
aqueous solution, and (f–u) were stained in 1% Congo red aqueous solution.

J. Fungi 2024,10, 439 25 of 36
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Figure 14. Morphological features of Mycena digitifurcata (FFAAS1055, holotype). (a) Basidiomata;
(b) basidia; (c) basidiospores; (d) cheilocystidia; (e) stipitipellis; (f) pileipellis and context. Bars: (a)
= 5 mm; (b,d–f) = 10 µm; (c) = 5 µm. Drawing by Renxiu Wei.
Basidiospores [108/4/2] 6.3–7.3–8.8 × 3.8–4.5–5.7 µm, [Q = 1.43–1.76, Qm = 1.61 ± 0.08]
[holotype (40/2/1) 6.4–7.5–8.8 × 3.8–4.7–5.7 µm, Q = 1.43–1.76, Qm = 1.60 ± 0.08], ellipsoid
to narrowly ellipsoid, hyaline, smooth (under oil), thin-walled, amyloid. Basidia 18–24 ×
4–8 µm, two- and four-spored, clavate, hyaline, sterigmata 3.2–5.0 µm in length. Cheilocys-
tidia 10–32 × 4–18 µm, clavate, cylindrical, apex with several to fairly numerous finger-like
branches, 2.3–22.1 × 0.8–2.1 µm, branches usually with furcate excrescences at apex, 2.3–
8.9 × 0.8–1.9 µm, hyaline, thin-walled. Pleurocystidia absent. Pileipellis a cutis composed of
Figure 14. Morphological features of Mycena digitifurcata (FFAAS1055, holotype). (a) Basidiomata;
(b) basidia; (c) basidiospores; (d) cheilocystidia; (e) stipitipellis; (f) pileipellis and context. Bars:
(a)=5mm;(b,d–f) = 10 µm; (c)=5µm. Drawing by Renxiu Wei.
MycoBank no: 853784
Etymology: The epithet digitifurcata derives from the Latin words ‘digitus’, meaning
‘finger’, and ‘furcatus’, meaning ‘forked’. This name is chosen to describe the distinctive
finger-like and forked projections at the apex of the cheilocystidia.
Holotype: CHINA. Zhejiang Province, Baiyun National Forest Park, Lishui City, 2
August 2021, leg. Renxiu Wei, Zewei Liu, Qin Na, and Yupeng Ge, 212 m asl, FFAAS1055
(collection no. MY0476).

J. Fungi 2024,10, 439 26 of 36
Diagnosis: Pileus deep gray when young, light drab to drab with age, basidiospores
ellipsoid to narrowly ellipsoid, cheilocystidia with finger-like branches at apex. Differs
from M. cristinae by pruinose pileus, obviously decurrent lamellae, and being weakly
intervenose.
Description: Pileus 4.2–12.0 mm diam., hemispherical when young, oblate hemispheri-
cal to convex with age, umbilicate at the center, margin slightly wavy, revolute, sometimes
cracked with age; Castor Gray (LII35””’i) to Pale Violet-Gray (LII59””’d) at the center,
gradually paler towards margin to *Pearl Gray (LII35””’f), Smoke Gray (XLVI21””d) when
young, Light Drab (XLVI17””b) to Hair Brown (XLVI17””i) at the center, gradually paler
towards margin to Pale Smoke Gray (XLVI21””f), Pale Drab-Gray (XLVI17””f) with age;
densely covered with pruina when young, sparsely when old, striate towards the center
up to 0.5–0.8 R, Castor Gray (LII35””’i) when young, Hathi Gray (LII35””’b), Hair Brown
(XLVI17””i) when matured, dry. Context White (LIII), fragile, thin. Lamellae subdecurrent
to decurrent, 0.3–0.9 mm wide, L= 12–14, l= 1–3, White (LIII) to pale, edge concolorous
with face, sometimes non-marginate, weak and irregularly intervened, up to 1/3–1/2
lamellae wide. Stipe 10–13
×
0.8–1.0 mm, central, clavate to cylindrical, hollow, fragile,
*French Gray (LII45””’f), *Pearl Gray (LII35””’f) in the upper part, darker towards the
base to *Cinereous (LII45””’d), Hathi Gray (LII35””’b), finely White (LIII) pruinose and
pubescent, base slightly swollen, covered with masses of short and White (LIII) tomentum
when young, rarely observed when old. Odor and taste indistinctive.
Basidiospores [108/4/2] 6.3–7.3–8.8
×
3.8–4.5–5.7
µ
m, [Q = 1.43–1.76, Q
m
= 1.61
±
0.08]
[holotype (40/2/1) 6.4–7.5–8.8
×
3.8–4.7–5.7
µ
m, Q = 1.43–1.76, Q
m
= 1.60
±
0.08], ellipsoid
to narrowly ellipsoid, hyaline, smooth (under oil), thin-walled, amyloid. Basidia 18–24
×
4–8
µ
m, two- and four-spored, clavate, hyaline, sterigmata 3.2–5.0
µ
m in length. Cheilocys-
tidia 10–32
×
4–18
µ
m, clavate, cylindrical, apex with several to fairly numerous finger-like
branches, 2.3–22.1
×
0.8–2.1
µ
m, branches usually with furcate excrescences at apex, 2.3–8.9
×
0.8–1.9
µ
m, hyaline, thin-walled. Pleurocystidia absent. Pileipellis a cutis composed of par-
allel hyphae, 1.9–4.6
µ
m diam., hyaline, thin-walled, covered with cylindrical excrescences,
1.8–17.9 (29.0)
×
0.9–2.3
µ
m; pileocystidia absent. Lamellae trama subregular, hyphae
7–19 µm
diam., hyaline, dextrinoid. Stipitipellis a cutis composed of a hypha, 2.2–4.6
µ
m diam.,
hyaline, thin-walled, covered with small cylindrical excrescences, 1.1–6.3
×
0.9–2.1
µ
m;
caulocystidia absent. Clamps present in all tissues, but rarely observed in context.
Habit and habitat: Scattered on rotten branches in mixed broadleaf–conifer forests,
mainly under trees of Liriodendron,Pseudolarix, and Pinus.
Known distribution: Zhejiang Province, China.
Additional material examined: China. Zhejiang Province, Lishui City, 1 August 2021,
leg. Renxiu Wei, Binrong Ke, Zhiheng Zeng, Qin Na, and Yupeng Ge, 231 m asl, FFAAS1054
(collection no. MY0447).
Notes: M. cristinae closely resembles M. digitifurcata in pileus and stipe color, but it is
distinguished by its smooth pileus, adnate lamellae, and markedly intervenose lamellae [
26
].
M. pasvikensis Aronsen, sharing a similar pileus color with M. digitifurcata, is differentiated
by a densely fibril-covered stipe base, serrulate lamellae, and gelatinized pileipellis and
stipitipellis [
28
]. Similar cheilocystidia shapes are observed in M. pseudopicta (J.E. Lange)
Kühner and M. cinerella (P. Karst.) P. Karst., but M. pseudopicta is differentiated from
M. digitifurcata by its gelatinized pileipellis and stipitipellis, and by possessing larger
basidiospores [
23
,
28
]; M. cinerella is notable for its gelatinized pileipellis and the lighter
color of the pileus [
28
,
29
]. Microscopic observation reveals a variation in the branch lengths
of the cheilocystidia. In specimen FFAAS1054, most cheilocystidia display short (0.9–1.9
µ
m)
branched excrescences at the apex, whereas in FFAAS1055, cheilocystidia with significantly
longer (7.5–22.2
µ
m) branched excrescences at the apex are typically observed, indicating
the various forms of cheilocystidia.
Mycena limitis R.X. Wei, L. Zou, Y.P. Ge & Q. Na, sp. nov., Figures 15–17.

J. Fungi 2024,10, 439 27 of 36
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Figure 15. Basidiomata of Mycena limitis sp. nov. (a,b) FFAAS1056; (c–e) FFAAS1058 (holotype); (f–
h) FFAAS1057; (e) Lamellae; (g) lamellae and surface of stipe are glabrous; (h) fibrils in base of stipe.
Bars: (a–e,g) = 5 mm; (f,h) = 10 mm. Photos (a–e) by Qin Na; (f–h) Yupeng Ge.
Figure 15. Basidiomata of Mycena limitis sp. nov. (a,b)FFAAS1056; (c–e)FFAAS1058 (holotype);
(f–h)FFAAS1057
; (e) Lamellae; (g) lamellae and surface of stipe are glabrous; (h) fibrils in base of
stipe. Bars: (a–e,g)=5mm;(f,h) = 10 mm. Photos (a–e) by Qin Na; (f–h) Yupeng Ge.

J. Fungi 2024,10, 439 28 of 36
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Figure 16. Microscopical features of Mycena limitis (FFAAS1058, holotype). (a–e) Basidiospores; (f)
basidium; (g–l) cheilocystidia; (m–r) pleurocystidia; (s) pileipellis; (t) hymenia and lamellar trama;
(u) stipitipellis. Bars: (a–e) = 5 µm; (f) = 10 µm; (g–u) = 20 µm. Structures (a–e) were rehydrated in
5% KOH aqueous solution, and (f–u) were stained in 1% Congo red aqueous solution.
Figure 16. Microscopical features of Mycena limitis (FFAAS1058, holotype). (a–e) Basidiospores;
(f) basidium; (g–l) cheilocystidia; (m–r) pleurocystidia; (s) pileipellis; (t) hymenia and lamellar trama;
(u) stipitipellis. Bars: (a–e)=5
µ
m; (f) = 10
µ
m; (g–u) = 20
µ
m. Structures (a–e) were rehydrated in
5% KOH aqueous solution, and (f–u) were stained in 1% Congo red aqueous solution.

J. Fungi 2024,10, 439 29 of 36
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Figure 17. Morphological features of Mycena limitis (FFAAS1058, holotype). (a) Basidiomata; (b) ba-
sidia; (c) basidiospores; (d) cheilocystidia; (e) pleurocystidia; (f) stipitipellis; (g) pileipellis and con-
text. Bars: (a) = 20 mm; (b,d–g) = 20 µm; (c) = 10 µm. Drawing by Renxiu Wei.
Basidiospores [108/4/3] (8.6) 8.9–10.4–12.6 × 4.7–5.7–6.9 (7.2) µm [Q = 1.65–1.99 (2.01),
Qm = 1.82 ± 0.09] [holotype (50/2/1) 8.9–10.7–12.6 × 4.7–5.8–6.9 (7.2) µm, Q = (1.65) 1.66–1.99
(2.01), Qm = 1.83 ± 0.09], narrowly ellipsoid, hyaline, smooth (under oil), thin-walled, am-
yloid. Basidia 29–37 × 8–10 µm, clavate, hyaline, two- and four-spored, sterigmata 4.1–6.0
µm in length. Cheilocystidia 43–70 × 10–21 µm, fusiform, subfusiform, lanceolate, tapering
at apex, base constricted to cylindrical, hyaline, thin-walled, smooth. Pleurocystidia 54–123
× 14–26 µm, similar to cheilocystidia, hyaline, thin-walled, occasionally forked in the
Figure 17. Morphological features of Mycena limitis (FFAAS1058, holotype). (a) Basidiomata; (b) ba-
sidia; (c) basidiospores; (d) cheilocystidia; (e) pleurocystidia; (f) stipitipellis; (g) pileipellis and context.
Bars: (a) = 20 mm; (b,d–g) = 20 µm; (c) = 10 µm. Drawing by Renxiu Wei.
MycoBank no: 853786
Etymology: The epithet limitis is derived from the Latin word limes, which emphasizes
the morphological similarities with related species and highlights the limited distinguishing
features that set this species apart from its close relatives. Additionally, this name pays
tribute to the companionship and support provided by a forest ranger, affectionately called
brother on the border.

J. Fungi 2024,10, 439 30 of 36
Holotype: CHINA. Heilongjiang Province, Taipinggou National Nature Reserve,
Hegang City, 3 July 2023, leg. Renxiu Wei, Li Zou, Menghui Han, Nannan Geng, Tingting
Sun, Xinyu Tong, Yawei Li, Zengcai Liu, and Yupeng Ge, 621 m asl, FFAAS1058 (collection
no. GN1786).
Diagnosis: Pileus hair brown when young, olive brown when old, stipe with faint
longitudinal stripes, cheilocystidia and pleurocystidia mainly fusiform. Differs from M.
niveipes by cheilocystidia with tapered apex and stipitipellis with smooth hyphae.
Description: Pileus 10–50 mm diam., parabolic and hemispherical when young, oblate
hemispherical to plano-convex with age, slightly convex at center, margin slightly serrulate,
sometimes cracked at mature; *Hair Brown (XLVI17””i) at center, gradually paler towards
margin to Light Drab (XLVI17””i), Pale Smoke Gray (XLVI21””f) when young, Deep Olive
(XL21”’k) at center, gradually paler towards margin to Citrine Drab (XL21”’i), Pale Olive
Buff (XL21”’f) when matured; covered with finely White (LIII) pubescence when young,
then glabrescent, transparent sulcate towards the center up to 0.5–0.6 R, dry. Context
White (LIII), fragile, thin. Lamellae adnate to slightly sinuate, 2.1–5.0 mm wide,
L= 22–25
,
l= 1–3
, White (LIII), edge concolorous with face, inconspicuous intervenose. Stipe
59–85 ×
2.0–4.1 mm
, central, cylindrical, White (LIII) in the upper part, Pale Smoke Gray (XLVI21””f)
in the middle part, *Smoke Gray (XLVI21””d), *Olive-Gray (LI23””’b) towards base, hollow,
fragile, covered with finely White (LIII) pruina when young, almost glabrous when old,
with faint longitudinal stripes, base covered with masses of short and White (LIII) fibrils.
Odor and taste indistinctive.
Basidiospores [108/4/3] (8.6) 8.9–10.4–12.6
×
4.7–5.7–6.9 (7.2)
µ
m [Q = 1.65–1.99 (2.01),
Q
m
= 1.82
±
0.09] [holotype (50/2/1) 8.9–10.7–12.6
×
4.7–5.8–6.9 (7.2)
µ
m, Q = (1.65)
1.66–1.99
(2.01), Q
m
= 1.83
±
0.09], narrowly ellipsoid, hyaline, smooth (under oil), thin-
walled, amyloid. Basidia 29–37
×
8–10
µ
m, clavate, hyaline, two- and four-spored, sterig-
mata 4.1–6.0
µ
m in length. Cheilocystidia 43–70
×
10–21
µ
m, fusiform, subfusiform, lanceo-
late, tapering at apex, base constricted to cylindrical, hyaline, thin-walled, smooth. Pleu-
rocystidia 54–123
×
14–26
µ
m, similar to cheilocystidia, hyaline, thin-walled, occasionally
forked in the upper part. Pileipellis a cutis composed of parallel hyphae, 3.2–7.0
µ
m diam.,
hyaline, thin-walled, smooth; pileocystidia absent. Lamellae trama regular, hyaline, parallel,
hyphae, 6–18
µ
m diam., dextrinoid. Stipitipellis a cutis composed of a hypha, 3.2–5.0
µ
m
diam., hyaline, thin-walled, smooth; caulocystidia absent. Clamps present in all tissues, but
rarely observed in context.
Habit and habitat: Scattered on humus layer and rotten branches in deciduous broad-
leaved forests, mainly under trees of Betula,Quercus,Styphnolobium, and Tilia.
Known distribution: Heilongjiang Province, Jilin Province, China.
Additional material examined: China. Jilin Province, Changbaishan National Nature
Reserve, Yanbian Korean Autonomous Prefecture, leg. Renxiu Wei, Binrong Ke, Chi Yang,
Qin Na, and Yupeng Ge, 1 July 2021, 708 m asl, FFAAS1057 (collection no. MY0305); 3 July
2021, 749 m asl, FFAAS1056 (collection no. MY0341).
Notes: M. niveipes is similar to M. limitis in pileus color and pileipellis with smooth
hyphae, but differs by rounded-apex cheilocystidia and the presence of swollen terminal
cells in the stipitipellis [
23
,
28
]. M. abramsii and M. subcana A.H. Sm also share a compa-
rable pileus color and cheilocystidia shape but are distinguished from M. limitis by the
pileipellis and stipitipellis, which are remarkedly covered with excrescences [
21
,
23
,
28
,
29
].
Similarly, M. galericulata exhibits a pileus color similar to M. limitis, but it is differentiated
by the cheilocystidia and both the hyphae of the pileipellis and stipitipellis featuring excres-
cences [
21
,
23
,
28
,
29
]. M. laevigata Gillet, while having comparable cheilocystidia, is distinct
from M. limitis due to its pale grayish-white pileus, the absence of pleurocystidia, and a
gelatinized pileipellis [28,29].
4. Discussion
Determining the identity of various brownish Mycena species based on basidiomata
color imposes certain limitations, which consequently may lead to an underestimation

J. Fungi 2024,10, 439 31 of 36
of species diversity [
11
–
14
,
21
,
23
,
28
]. Specifically, M. campanulatihemisphaerica,M. limitis,
and M. kunyuensis demonstrate close affiliations with M. abramsii,M. algeriensis Maire, M.
galericulata, and M. maculate P. Karst., all of which are endemic to China [
21
]. Notably,
M. campanulatihemisphaerica,M. kunyuensis, and M. abramsii exhibit a similar basidiomata
coloration, with M. kunyuensis frequently misidentified as M. abramsii due to the analogous
coloration of the pileus before morphological anatomy [
21
,
23
,
28
]. Similarly, M. limitis is
indistinguishable from M. algeriensis,M. galericulata, and M. maculata when identification is
based solely on the color of the basidiomata [
21
,
28
]. Moreover, species within the complex
exhibit a comparable pileus coloration and are subject to variations due to changes in growth
period and environment conditions, further complicating the accurate identification of these
brownish Mycena species [
11
–
14
,
69
–
77
]. For example, the M. filopes complex, as delineated
by Arnolds (2015) and Aronsen (2016), is a case in point. [
28
,
78
]. A parallel example is
found in the M. pura complex. While basidiomata coloration has historically served as a
criterion for subsection classification within the M. pura complex, recent insights by Liu
(2023) suggest that the characteristics of the cheilocystidia and pleurocystidia, as well as the
presence or absence of pleurocystidia, are critical for species differentiation [
11
,
12
,
14
,
79
].
Consequently, morphological anatomy is an effective method for the identification of
species within the brownish Mycena group.
Pileipellis types and cheilocystidia characteristics are integral to the delimitation of
brownish Mycena species. In the taxonomic framework proposed by Smith (1947) and Maas
Geesteranus (1992a, 1992b), pileipellis types and cheilocystidia characteristics are pivotal
for sectional categorization [
11
,
12
,
20
]. These characteristics are also crucial criteria for dis-
tinguishing various species of brownish Mycena, a delineation corroborated by molecular
systematics. The phylogenetic tree constructed in this study is divided into 13 clades, and
while some clades have lower support rates, the conclusions drawn from the phylogenetic
tree remain consistent with those from the morphological anatomy analyses. Notably, M.
campanulatihemisphaerica,M. limitis, and M. kunyuensis exhibit smooth cheilocystidia, and
cluster phylogenetically into a clade and align closely with the species of sect. Fragilipedes,
which also exhibit smooth cheilocystidia [
17
,
28
]. Moreover, phylogenetic analyses reveal
a distinct clade comprising M. digitifurcata, which aligns closely with the species of sect.
Rubromarginatae [
26
,
57
]. Sect. Rubromarginatae is characterized by marginta lamellae, rang-
ing from deep yellow to dark greenish [
11
,
12
]. If the sectional division relies solely on this
feature, M. digitifurcata would ostensibly not qualify for inclusion within the sect. Rubro-
marginatae due to its lamellae not being marginated [
11
,
12
]. However, its pileipellis types
and cheilocystidia characteristics are congruent with those of the sect. Rubromarginatae,
suggesting the need for a broader consideration of morphological characters. Accordingly,
M. digitifurcata is classified within the sect. Rubromarginatae. This classification, proposed by
Jadson (2021), identifies M. cristinae as a non-marginate lamellae member of the sect. Rubro-
marginatae [
26
]. Our research aligns with Maas Geesteranus’ (1992a, 1992b) interpretation
of the sect. Rubromarginatae, which is further supported by the perspectives advanced by
Na (2019) [
11
,
12
,
21
]. Specifically, it emphasizes that the delimitation of sections exhibiting
colored lamellae edges should predominantly consider variations in pileipellis types and
cheilocystidia characteristics. Furthermore, pileipellis and cheilocystidia are present in
almost all brownish Mycena, reinforcing their utility as key diagnostic features [
23
,
28
]. For
example, M. galericulata can be distinguished from its morphologic relative, M. algeriensis,
by its tuberculated cheilocystidia, and M. leptocephala is distinguished from its close rela-
tive M. polygramma (Bull.) Gray by smooth pileipellis [
11
,
12
,
21
,
23
,
28
]. Consequently, we
reaffirm the role of pileipellis types and cheilocystidia characteristics as primary taxonomic
criteria within the brownish Mycena group, corroborating the classifications of Smith (1947)
and Maas Geesteranus (1992a, 1992b) and substantiated by the molecular systematics of
Na (2019) [11,12,20,21].
Additionally, this study suggests that M. oryzifluens may represent a prospective novel
section. Phylogenetically, M. oryzifluens is closely related to three established sections,
namely: sect. Exornatae, sect. Cyanocephalae, and sect. Amictae. Despite these affiliations,

J. Fungi 2024,10, 439 32 of 36
M. oryzifluens exhibits distinct morphological characteristics that preclude its classification
within these existing sections. These distinguishing features include a dry pileus, the
absence of a blue disc at the stipe base, branched cheilocystidia apex, and non-gelatinized
pileipellis [
11
,
12
,
21
,
28
,
57
]. Consequently, M. oryzifluens does not align with any current
sectional definitions and suggests the potential for defining a new section. However, due
to the lack of enough specimen samples at present, it is prudent not to propose it as a new
section within this publication. Future research efforts will collect and examine additional
specimens to substantiate the distinctiveness of this taxonomic group.
At present, there are 24 brownish Mycena species in China. This study introduces
5 new species, increasing the total to 29. Our analysis of morphological characteristics has
identified the types of pileipellis and cheilocystidia as critical distinguishing features for
brownish Mycena. To facilitate future research and better species distinction, we provide an
identification key for the brownish Mycena species in China.
Key to the known species of brownish Mycena from China
1. Basidiospores inamyloid 2
1. Basidiospores amyloid 3
2. Basidiospore broadly ellipsoid, Q = 1.1–1.5 Mycena hiemalis
2. Basidiospore narrowly ellipsoid, Q = 1.7–1.9 Mycena speirea
3. Pileipellis gelatinized 4
3. Pileipellis not gelatinized 10
4. Stipitipellis hyphae ornamented 5
4. Stipitipellis hyphae smooth 6
5. Pleurocystidia present Mycena clavicularis
5. Pleurocystidia absent Mycena polygramma
6. Caulocystidia absent Mycena semivestipes
6. Caulocystidia present 7
7. Cheilocystidia densely covered with tuberculate excrescences Mycena pluteoides
7. Cheilocystidia smooth 8
8. Pileus with conical spines, basal disc of the stipe present, caulocystidia
with outgrowths Mycena stylobates
8. Pileus without any spines, stipe base without disc, caulocystidia without
outgrowths 9
9. Pileus convex, depressed at center, lamellae stained with yellow-brown to
orange-brown spots Mycena subpiligera
9. Pileus conical to campanulate, umbonate at center, lamellae without
any spots Mycena amicta
10. Pileipellis hyphae smooth 11
10. Pileipellis hyphae ornamented 12
11. Caulocystidia present Mycena algeriensis
11. Caulocystidia absent Mycena limitis
12. Lamellae flesh pink, pleurocystidia with flesh-pink contents Mycena entolomoides
12. Lamellae white to grayish white, pleurocystidia hyaline 13
13. Caulocystidia present 14
13. Caulocystidia absent 18
14. Cheilocystidia present 15
14. Cheilocystidia absent 16
15. Cheilocystidia fusoid to lageniform, with long tapered necks, smooth Mycena subcana
15. Cheilocystidia clavate to obpyriform, densely covered with cylindrical
excrescences Mycena mirata
16. Pileus with brown spots in age, stipitipellis hyphae ornamented Mycena zephirus
16. Pileus without any spots in age, stipitipellis hyphae smooth 17
17. Basidiospore subglobose to broadly ellipsoid, Q = 1.1–1.4 Mycena oryzifluens
17. Basidiospore narrowly ellipsoid to cylindrical, Q = 1.6–2.1 Mycena leptocephala
18. Pleurocystidia present 19
18. Pleurocystidia absent 23

J. Fungi 2024,10, 439 33 of 36
19. Cheilocystidia apically rounded or narrowed into one to several
cylindrical or furcate necks 20
19. Cheilocystidia covered with cylindrical excrescences 21
20. Stipe grayish brown, basidiospore broadly ellipsoid to ellipsoid,
Q = 1.4–1.7 Mycena silvae-nigrae
20. Stipe dark brown, basidiospore narrowly ellipsoid to cylindrical,
Q = 1.7–2.1 Mycena abramsii
21. Pileus convex, basidiospore narrower than 4 µmMycena luguensis
21. Pileus conical to campanulate, basidiospore broader than 4 µm 22
22. Pileipellis with clavate to subglobose terminal cell Mycena filopes
22. Pileipellis without clavate to subglobose terminal cell Mycena metata
23. Cheilocystidia apex narrowed into a cylindrical or furcate neck, rarely
with short apical outgrowths 24
23. Cheilocystidia densely covered with tuberculate or furcate excrescences,
or apex with several branches 25
24. Basidiospore ellipsoid to narrowly ellipsoid, Q = 1.4–1.7 Mycena kunyuensis
24. Basidiospore narrowly ellipsoid to cylindrical, Q = 1.7–2.0 Mycena campanulati-
hemisphaerica
25. Cheilocystidia apex with several branches 26
25. Cheilocystidia densely covered with tuberculate or furcate excrescences
27
26. Pileus bell-shaped, brown sulcate present, not depressed at center,
stipe brown Mycena venus
26. Pileus oblate hemispherical to convex, brown sulcate absent, depressed
at center, stipe gray Mycena digitifurcata
27. Pileus and lamellae with red-brown spots when old, cheilocystidia
sparsely covered with excrescences Mycena maculata
27. Pileus and lamellae without any spots when old, cheilocystidia densely
covered with excrescences 28
28. Basidiospore narrowly ellipsoid to cylindrical, Q = 1.9–2.2 Mycena flos-nivium
28. Basidiospore broadly ellipsoid to ellipsoid, Q = 1.1–1.7 Mycena galericulata
Author Contributions: Conceptualization and methodology: Q.N.; validation, investigation, data
curation, and writing—original draft preparation: R.W., Y.G., L.Q., M.H., H.Z., Y.H., L.Z., X.C. and
X.W.; writing—review and editing: R.W. and Y.G. All authors have read and agreed to the published
version of the manuscript.
Funding: This study was supported by the National Natural Science Foundation of China (grant no.
32200008), the Natural Science Foundation of Shandong Province (grant no. ZR2020QC001), the 5511
Collaborative Innovation Project of Fujian Province (grant no. XTCXGC2021007), the Central Public
Interest Scientific Institution Basal Research Fund (grant no. GYZX200203), the east-west cooperation
project, FAAS (grant no. DKBF-2022-12), the Natural Science Foundation of Fujian Province (grant
no. 2023J01379), the Project of Biological Resources Survey in Wuyishan National Park (grant no.
HXQT2020120701), the Project of Biodiversity Conservation in Lishui, Zhejiang Province (grant no.
HXYJCP2021110648), the Shandong Agricultural Industry Technology System (2021 grant no. 26,
SDAIT-07-03), the China Agriculture Research System (CARS20), the Technology Development Fund
of Guangxi Academy of Agricultural Sciences (Guinongke 2021JM19), and the China Agriculture
Research System, Guangxi Edible Fungi Innovation Team (nycytxgxcxtd-2021-07-01).
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: Data are contained within the article.
Acknowledgments: Thanks to Junqing Yan from the Jiangxi Agriculture University; Chi Yang and
Binrong Ke from the Institute of Edible Fungi, Fujian Academy of Agricultural Sciences; Tingting Sun
Zengcai Liu, Nannan Geng, Xinyu Tong, and Yawei Li from the College of Forestry, Northeast Forestry
University; Liying Li and Yongqiang Hu from the Microbiology Research Institute, Guangxi Academy
of Agricultural Sciences; Zewei Liu from the Kunming Institute of Botany, Chinese Academy of
Sciences; and Yulan Sun, Liming Xue, and Ruichen Liu from Ludong University, for collecting

J. Fungi 2024,10, 439 34 of 36
samples jointly for this study. We sincerely thank the reviewers for their corrections and suggestions
to improve our work.
Conflicts of Interest: The authors declare no conflicts of interest.
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