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Research Article
Three new species of Mycena sect. Calodontes (Mycenaceae, Agaricales) from Northeastern China
expand article infoJingwen Guo‡, Zewei Liu‡, Hui Zeng§, Yupeng Ge§‡, Qin Na‡
‡ Ludong University, Yantai, China
§ Institute of Edible Fungi, Fujian Academy of Agricultural Sciences, Fuzhou, China

Corresponding author: Qin Na ( naqin19890317@163.com )

Academic editor: Heng Zhao
© 2025 Jingwen Guo, Zewei Liu, Hui Zeng, Yupeng Ge, Qin Na.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Guo J, Liu Z, Zeng H, Ge Y, Na Q (2025) Three new species of Mycena sect. Calodontes (Mycenaceae, Agaricales) from Northeastern China. MycoKeys 125: 51-79. https://doi.org/10.3897/mycokeys.125.169722
Open Access

Abstract

Mycena sect. Calodontes is a large section within the genus Mycena, with 44 species described worldwide. The section is well characterized by relatively large basidiomata, typically growing on the humus layer of coniferous or coniferous-broadleaved mixed forests. Only 19 species of sect. Calodontes have been previously recorded in China, more than half of sect. Calodontes species are distributed in east and north regions, but also distributed in west and south regions. Based on 8 specimens collected from Heilongjiang and Jilin Province, 3 novel species are formally described: M. brunneocystidiata sp. nov., M. roseopurpurea sp. nov., and M. rubrofusca sp. nov. Mycena brunneocystidiata is distinguished by purple-brown to brown pileus, brown lamellae margins, and cystidia with brown contents. Mycena roseopurpurea is characterized by the pileus with brownish center, white lamellae margins, and colorless, fusiform cystidia. Mycena rubrofusca is distinguished by brownish pileus, white lamellae margins, and colorless, utriform cystidia. Detailed morphological descriptions, high-resolution habitat photographs, line drawings, and comparison with closely related taxa are provided for the new species. A combined phylogenetic analysis was conducted, based on a multi-locus (ITS+RPB1+tef1-α) dataset under Bayesian Inference (BI) and Maximum Likelihood (ML) analyses. The morphological data and the results of the phylogenetic analyses support the recognition of the 3 new species. A key to the 20 species currently known species of sect. Calodontes in China is also provided.

Key words:

New taxon, systematics, taxonomy, white-spored fungi

Introduction

Mycena (Pers.) Roussel is a typical saprobic fungus which plays a pivotal role in forest ecosystems (Smith 1947; Perry 2002; Kirk et al. 2008; Na 2019). Species in this genus are known for their ability to decompose rotten branches and wood, fallen leaves, and various organic substrates, and could be facilitating nutrient cycling and sustaining energy flow (Fukasawa et al. 2009; Baldrian and Kohout 2017; Kyaschenko et al. 2017; Guerreiro et al. 2023). Within the genus, sect. Calodontes (Fr. Ex Berk.) Quél. is recognized for its large numbers of species and relatively large basidiomata, in which the pileus diameter can reach 6 cm, hygrophanous pileus, irregularly intervenose lamellae, mostly amyloid basidiospores, and smooth cystidia, pileipellis, and stipitipellis (Smith 1947; Maas Geesteranus 1992a, 1992b; Maas Geesteranus and de Meijer 1997; Grgurinovic 2003; Robich 2003; Aronsen and Læssøe 2016; Na 2019; Liu 2023). Mycena sect. Calodontes is primarily distributed in the mid- to high- latitude regions of the Northern Hemisphere, including Europe and America, with only a few species known from lower latitudes in the Southern Hemisphere (Maas Geesteranus 1992a, 1992b; Grgurinovic 2003; Chew et al. 2014; Cooper et al. 2018; Liu et al. 2021, 2022, 2024; Cortés-Pérez et al. 2023). In 1821, Fries erected subtri. Calodontes under Agaricus trib. Clitocybe, later Berkeley (1836) elevated Clitocybe (Fr.) Staude to subgenus rank, and sect. Calodontes was raised to the rank of section, and the concept of sect. Calodontes has been widely adopted (Fries 1821; Berkeley 1836; Quéltet 1875; Maas Geesteranus 1992a, 1992b; Grgurinovic 2003; Aronsen and Læssøe 2016; Na 2019; Liu 2023). By the early 21st century, 44 species of sect. Calodontes are known, including 17 newly species recorded from Asia, 5 from North America, 2 from Africa, and 1 from Oceania (Grgurinovic 2003; Chew et al. 2014; Cooper et al. 2018; Liu et al. 2021, 2022, 2024; Qiang and Bai 2022; Cortés-Pérez et al. 2023; Fan et al. 2024; Nagamune et al. 2024; Xiao et al. 2025). Species of sect. Calodontes adapt to various temperature and humidity conditions, widely growing on the humus layer of coniferous forests in North Temperate and Cold Temperate Zones, a few species also reported from tropical and subtropical climates, including India, Malaysia, São Tomé, and southern Australia (Maas Geesteranus 1992a, 1992b; Grgurinovic 2003; Chew et al. 2014; Cooper et al. 2018; Cortés-Pérez et al. 2023; Liu 2023).

Over the past three decades, the color of pileus, and the shape, contents, thickness of the walls of cheilocystidia have been confirmed to be the diagnostic characters of sect. Calodontes (Maas Geesteranus 1992a, 1992b; Grgurinovic 2003; Robich 2003; Harder et al. 2010, 2013; Liu 2023). The color of pileus in sect. Calodontes, ranging from pink, sulfur yellow, white, purple to brown, is easily changed by the temperature, humidity, and growth stage, so the characteristics of cheilocystidia show more effectiveness (Kauserud et al. 2008; Liu 2023). In sect. Calodontes, cheilocystidia are fusiform, clavate, or utriform (sometimes with tapering apices), and apart from their shape, the thickness of their walls can also be used to distinguish some species from closely related taxa (Maas Geesteranus 1992a, 1992b; Grgurinovic 2003; Robich 2003; Liu et al. 2021, 2022; Liu 2023). According to a phylogenetic reconstruction of sect. Calodontes, based on the materials from Europe and North America, derived from an internal transcribed spacer (ITS), RNA polymerase II largest subunit (RPB1), and translation elongation factor-1 alpha (tef1-α) sequence dataset, the results supported the color of pileus and the characteristics of cheilocystidia can be used to delimit species, but the varieties and forms were not supported (Harder et al. 2010, 2013). Additionally, Harder et al. (2010, 2013) also proposed that the RPB1 and tef1-α sequences improved the ability to identify phylogenetic species in the M. pura complex (Harder et al. 2010, 2013).

Materials and methods

Specimen collection and macroscopic characteristics recording

During field investigations, each specimen was assigned a unique collection number. High-resolution photos were taken using a Canon EOS 90D digital camera (Canon, Tokyo, Japan) equipped with an EF-S 60 mm f/2.8 Macro USM lens. Comprehensive ecological data were recorded, including forest type, substrate, elevation, climate, season, GPS coordinates, and macroscopic characteristics such as pileus, lamellae (intervenose), context, stipe (base), odor, and taste. Color codes and notations followed Ridgway (Ridgway 1912). A small fragment of context was excised from each specimen for molecular analyses after macroscopic documentation. Specimens were dried at 40 °C using a Stöckli dehydrator (A. & J. Stöckli AG, Netstal, Switzerland) and stored in self-sealing plastic bags containing color-changing silica gel. All voucher specimens were deposited in the Fungarium of the Fujian Academy of Agricultural Sciences (FFAAS), China.

Microscopic characteristics’ observation and morphological description

Dried specimens were rehydrated in 5% KOH aqueous solution and examined using a Lab A1 light microscope (Carl Zeiss AG, Jena, Germany). The tissues were stained with 1% Congo red aqueous solution when necessary. Microscopic structures were photographed and measured using ZEN 2.3 software. For each specimen, basidiospores were observed in lateral view, and 20 mature basidiospores were randomly measured. The dimensions of basidiospores and Q values are presented as [a/b/c] (d)e–f–g(h) × (i)j–k–l(m) µm [Q = (n)o–p(q), Qm = r ± s]. The notation [a/b/c] refers to the number of basidiospores, basidiomata, and specimens measured, respectively. Spore dimensions are presented as (d)e–f–g(h) × (i)j–k–l(m), where d and h denote the 5% minimum and maximum values, e–g indicate the central 90% range, and f represents the mean value. Q represents the length-to-width ratio, with Q = (n)o–p(q) indicating the range, and Qm = r ± s denoting the mean and standard deviation. For the type specimen, two basidiomata were examined, with at least 20 mature basidiospores measured per individual, following Na et al. (2021), Liu et al. (2021, 2022), and Wei et al. (2024). Amyloid reactions of basidiospores and lamellar trama were tested using Melzer’s reagent (Vizzini et al. 2020). For all other microscopic structures, at least 20 basidia were measured, as well as measurements of the shape of cheilocystidia, pleurocystidia (if present), and caulocystidia (if present), and the hyphae of the pileipellis and stipitipellis were also measured, the contents, and wall thickness of cheilocystidia, pleurocystidia, and caulocystidia were also observed (Liu et al. 2022). Line drawings were prepared based on habitat photographs, field notes, and microscopic observations, scanned using a Canon LiDE120 scanner (Canon, Tokyo, Japan), and finalized using Adobe Photoshop 2023.

Phylogenetic analyses

DNA sequence acquisition

Genomic DNA was extracted using the New Plant Genomic DNA Extraction Kit (Cowin Century, Beijing, China) and stored at -20 °C. Three nuclear loci, comprising the internal transcribed spacer (ITS), RNA polymerase II largest subunit (RPB1), and translation elongation factor-1 alpha (tef1-α), were amplified using the primer pairs ITS1/ITS4, RPB1Mp_f1/RPB1Mp_r1, tEFMp_f2/tEFMp_r2, respectively (Harder et al. 2010, 2013). PCR reactions were performed in 25 μL volumes, containing 12.5 μL of 2 × Utaq PCR MasterMix (ZomanBio, Beijing, China), 1 μL of each primer, 2 μL of DNA template, with ddH2O added to reach the final volume (Liu et al. 2022). The PCR protocol for the ITS region consisted of an initial denaturation at 94 °C for 4 min, followed by 34 cycles of 94 °C for 45 s, 52 °C for 45 s, and 72 °C for 1 min, a final extension of 72 °C for 10 min (Harder et al. 2010). The amplification protocol for the RPB1 and tef1-α regions was as follows: 94 °C for 1 min, then 10 cycles of 94 °C for 35 s, 53 °C for 45 s, 72 °C for 45 s, and 25 cycles of 94 °C for 35 s, 56 °C for 45 s, 72 °C for 45 s, ending with a final extension of 72 °C for 10 min (Harder et al. 2013). PCR products were sequenced by the Beijing Genomics Institute (Beijing, China). The PCR products were cloned using the pBLUE-T Kit (Beijing Zoman Biotechnology Co., Beijing, China) to generate high-quality sequences.

Phylogenetic analyses

All newly generated sequences from the collected specimens were compared using BLAST in the NCBI database (https://www.ncbi.nlm.nih.gov/). Homologous sequences showing nucleotide identity greater than 90% were downloaded from GenBank (https://www.ncbi.nlm.nih.gov/genbank). Mycena rubromarginata (Fr.) P. Kumm was selected as the outgroup for phylogenetic analyses (Harder et al. 2010, 2013; Liu et al. 2022). Sequence alignment was performed using MAFFT v.7.110, with gaps treated as missing data. Peak profiles of the newly generated sequences were examined in BioEdit to detect insertion and deletion sites, and regions containing multiple copies were encoded using degenerate bases (Hall 1999; Katoh et al. 2002, 2019; Alzohairy 2011). Phylogenetic analyses were conducted using both Bayesian Inference (BI) and Maximum Likelihood (ML) methods in MrBayes v3.2.6 and raxmGUI 2.0.10 (Posada and Crandall 1998; Nylander 2004; Edler et al. 2021). The sequence matrix of 3 nuclear loci were divided into 6 partitions: ITS1, 5.8S, ITS2, RPB1 exons, tef1-α exons, and the combined introns regions of RPB1 and tef1-α. MCMC was run with 6 chains, and conducted sampling at intervals of 10,000 generations until the Average Deviation of Split Frequencies was below 0.01; the first 25% of trees were discarded as burn-in, using the ‘sump’ and ‘sumt burnin’ commands to generate the results (Ronquist and Huelsenbeck 2003; Ronquist et al. 2012). Tracer v.1.7.2 was used to evaluate the Effective Sample Size (ESS) and Average Potential Scale Reduction Factor (PSRF) values as indicators of Bayesian inference (BI) analysis (Rambaut et al. 2018; Fabreti and Höhna 2022). For the Maximum Likelihood (ML) analysis, default parameters in RAxML were used with 1,000 rapid bootstrap replicates to assess branch support (Edler et al. 2021). The resulting phylogenetic trees were visualized with FigTree v.1.4.3.

Results

Phylogenetic analyses

The dataset comprised 263 sequences, containing 24 newly generated sequences (8 ITS, 8 RPB1, and 8 tef1-α) and 239 sequences downloaded from GenBank (89 ITS, 70 RPB1, and 80 tef1-α). Detailed information for all sequences was provided in Table 1. The aligned dataset contained 1,538 nucleotide sites (including gaps), with 216 bp for ITS1 region, 159 bp for 5.8S region, 247 bp for ITS2 region, 55 bp for RPB1 exons region, 307 bp for tef1-α exons region, 433 bp for RPB1 introns region, and 121 bp for tef1-α introns region. Among the 24 newly generated sequences, 29 insertion sites and 23 deletion sites were identified, along with 2 degenerate bases, specifically R (1519 bp of FFAAS3406) and W (944 bp of FFAAS3407). For BI analysis, the best-fitting models for each partition of the concatenated dataset were selected as follows: GTR+G for ITS1 and RPB1 introns+tef1-α introns, JC for 5.8S, GTR+I+G for ITS2 and tef1-α exons, and HKY for RPB1 exons. The BI analysis, after 15,000,000 generations, yielded an average deviation of split frequencies of 0.006965, an effective sample size (ESS) of 1160.2, and a potential scale reduction factor (PSRF) ranging from 1.000 to 1.002. For ML analysis, the substitution models were as follows: JC for ITS1, 5.8S, and RPB1 exons, K80+G for ITS2, TVMef+G for tef1-α exons, and HKY+G for RPB1 and tef1-α introns regions. The final log-likelihood score was -8333.401114. The BI and ML analyses tree showed similar topologies, and the ML topology was selected to present the final phylogenetic tree (Fig. 1).

Table 1.
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Sequences of Mycena sect. Calodontes used in the phylogenetic analyses.

No. Species Voucher/Strain No. GenBank NO. Locality Reference
ITS RPB1 tef1-α
1 M. aff. pura TL8052 FN394623 KF723687 KF723641 Ecuador Harder et al. (2010, 2013)
2 M. aff. pura TL9433 FN394622 KF723688 KF723642 Ecuador Harder et al. (2010, 2013)
3 M. aff. pura TL9450 KJ144653 KF723689 KF723643 Ecuador Harder et al. (2010, 2013)
4 M. aff. pura TL9678 FN394621 KF723690 KF723644 Ecuador Harder et al. (2010, 2013)
5 M. brunnea (M. pura XI) CBH187 FN394564 KF723678 KF723632 Denmark Harder et al. (2010, 2013)
6 M. brunnea (M. pura XI) CBH386 FN394565 KF723679 KF723633 Denmark Harder et al. (2010, 2013)
7 M. brunneocystidiata FFAAS3400 Holotype PV939239 PV952260 PV952232 China This study
8 M. brunneocystidiata FFAAS3401 PV939240 PV952261 PV952233 China This study
9 M. cahaya ACL134 KF537248 – – Malaysia Chew et al. (2014)
10 M. densilamellata TUFC 101999 LC777686 LC777726 LC777734 Japan Nagamune et al. (2024)
11 M. densilamellata TUMH 65486 LC777687 LC777727 LC777735 Japan Nagamune et al. (2024)
12 M. densilamellata TUMH 65482 LC777688 LC777728 LC777736 Japan Nagamune et al. (2024)
13 M. densilamellata TNS-F-75029 LC777689 LC777729 LC777737 Japan Nagamune et al. (2024)
14 M. dura 10315 FN394560 KF723694 KF723648 Austria Harder et al. (2010, 2013)
15 M. lammiensis TUR165927 FN394552 KF723697 KF723651 Finland Harder et al. (2010, 2013)
16 M. luceata ACP2116 OR233614 OR233746 OR233755 Mexico Cortés-Pérez et al. (2023)
17 M. luceata ACP2126 OR233613 OR233745 OR233754 Mexico Cortés-Pérez et al. (2023)
18 M. lucisnieblae ACP2139 OR233611 OR233743 OR233753 Mexico Cortés-Pérez et al. (2023)
19 M. lucisnieblae ACP2166 OR233607 OR233740 OR233750 Mexico Cortés-Pérez et al. (2023)
20 M. lucisnieblae ACP2352-B OR233608 – OR233756 Mexico Cortés-Pérez et al. (2023)
21 M. luteovariegata (M. pura V) CBH226 FN394604 KF723664 KF723618 Denmark Harder et al. (2010, 2013)
22 M. luteovariegata (M. pura f. lutea) DB2005/152 FN394603 – – Denmark Harder et al. (2010)
23 M. luteovariegata (M. pura V) TL5614 FN394602 KF723666 KF723620 Denmark Harder et al. (2010, 2013)
24 M. luxmanantlanensis ACP2160 OR233603 OR233737 OR233747 Mexico Cortés-Pérez et al. (2023)
25 M. luxmanantlanensis ACP2159 OR233604 OR233738 OR233748 Mexico Cortés-Pérez et al. (2023)
26 M. pearsoniana LK880/2002 FN394613 KF723693 KF723647 Germany Harder et al. (2010, 2013)
27 M. pearsoniana CBH068 FN394614 KF723691 KF723645 Germany Harder et al. (2010, 2013)
28 M. pearsoniana JV06890 FN394612 KF723692 KF723646 Denmark Harder et al. (2010, 2013)
29 M. pelianthina CBH015 FN394549 KF723695 KF723649 Denmark Harder et al. (2010, 2013)
30 M. pelianthina CBH016 FN394547 KF723696 KF723650 Denmark Harder et al. (2010, 2013)
31 M. polycystidiata FFAAS0417 ON427731 ON468456 ON468469 China Liu et al. (2022)
32 M. polycystidiata FFAAS0418 ON427732 ON468457 ON468470 China Liu et al. (2022)
33 M. polycystidiata FFAAS0421 ON427733 ON468458 ON468471 China Liu et al. (2022)
34 M. polycystidiata FFAAS0422 ON427734 ON468459 ON468472 China Liu et al. (2022)
35 M. pura I CBH039 FN394588 KF723680 KF723634 Denmark Harder et al. (2010, 2013)
36 M. pura II CBH105 FN394581 KF723671 KF723625 Denmark Harder et al. (2010, 2013)
37 M. pura II CBH169 FN394579 KF723672 KF723626 Denmark Harder et al. (2010, 2013)
38 M. pura II CBH366 FN394572 KF723673 KF723627 Denmark Harder et al. (2010, 2013)
39 M. pura II CBH404 FN394566 KF723674 KF723628 Denmark Harder et al. (2010, 2013)
40 M. pura III CBH019 FN394605 KF723675 KF723629 Denmark Harder et al. (2010, 2013)
41 M. pura III CBH022 FN394574 KF723676 KF723630 Denmark Harder et al. (2010, 2013)
42 M. pura III KK FN394606 KF723677 KF723631 Slovakia Harder et al. (2010, 2013)
43 M. pura IV CBH410 FN394595 KF723667 KF723621 Denmark Harder et al. (2010, 2013)
44 M. pura IV JV06979 FN394585 KF723668 KF723622 Denmark Harder et al. (2010, 2013)
45 M. pura IV TL4571 FN394583 KF723669 KF723623 Denmark Harder et al. (2010, 2013)
46 M. pura IV TL12786 FN394591 KF723670 KF723624 Sweden Harder et al. (2010, 2013)
47 M. pura VI BAP132 FN394561 KF723660 KF723614 USA Harder et al. (2010, 2013)
48 M. pura VII IS10/11/2000 FN394611 – – USA Harder et al. (2010)
49 M. pura VIII CBH216 FN394598 KF723662 KF723616 Denmark Harder et al. (2010, 2013)
50 M. pura VIII CBH402 FN394599 KF723663 KF723617 Denmark Harder et al. (2010, 2013)
51 M. pura IX CBH166 FN394607 KF723701 KF723655 Denmark Harder et al. (2010), 2013
52 M. pura IX CBH358 FN394608 KF723702 KF723656 Denmark Harder et al. (2010, 2013)
53 M. pura IX CBH367 KF913022 KF723703 KF723657 Denmark Harder et al. (2013)
54 M. pura IX CBH371 KF913023 KF723704 KF723658 Denmark Harder et al. (2013)
55 M. rosea UP2 FN394550 – – UK Harder et al. (2010)
56 M. rosea CBH097 FN394556 KF723681 KF723635 Denmark Harder et al. (2010, 2013)
57 M. rosea CBH383 FN394553 KF723682 KF723636 Denmark Harder et al. (2010, 2013)
58 M. rosea CBH409 FN394551 KF723683 KF723637 Germany Harder et al. (2010, 2013)
59 M. rosea TL12393 FN394555 KF723684 KF723638 Denmark Harder et al. (2010, 2013)
60 M. rosea TL12409 FN394557 KF723685 KF723639 Denmark Harder et al. (2010, 2013)
61 M. roseopurpurea FFAAS3402 PV939241 PV952262 PV952228 China This study
62 M. roseopurpurea FFAAS3403 PV939242 PV952263 PV952229 China This study
63 M. roseopurpurea FFAAS3404 Holotype PV939243 PV952264 PV952230 China This study
64 M. roseopurpurea FFAAS3405 PV939244 PV952265 PV952231 China This study
65 M. rubrofusca FFAAS3406 Holotype PV939245 PV952266 PV952235 China This study
66 M. rubrofusca FFAAS3407 PV939246 PV952267 PV952234 China This study
67 M. rubromarginata JV09362 FN394624 KF723705 KF723659 Denmark Harder et al. (2010, 2013)
68 M. rufobrunnea FFAAS0414 ON427728 ON468453 ON468466 China Liu et al. (2022)
69 M. rufobrunnea FFAAS0415 ON427729 ON468454 ON468467 China Liu et al. (2022)
70 M. rufobrunnea FFAAS0416 ON427730 ON468455 ON468468 China Liu et al. (2022)
71 M. seminau ACL308 KF537252 – – Malaysia Chew et al. (2014)
72 M. seminau ACL136 KF537250 – – Malaysia Chew et al. (2014)
73 M. shengshanensis FFAAS0424 ON427739 ON468464 ON468477 China Liu et al. (2022)
74 M. shengshanensis FFAAS0425 ON427740 ON468465 ON468478 China Liu et al. (2022)
75 M. sinar ACL092 KF537247 – – Malaysia Chew et al. (2014)
76 M. sinar ACL135 KF537249 – – Malaysia Chew et al. (2014)
77 M. sinar var. tangkaisinar ACL307 KF537251 – – Malaysia Chew et al. (2014)
78 M. sophiae ACP2161 OR233605 – OR233757 Mexico Cortés-Pérez et al. (2023)
79 M. subbrunnea Liu 59 PP037944 PP034075 PP034079 China Liu et al. (2024)
80 M. subbrunnea Liu 265 PP037946 PP034076 PP034081 China Liu et al. (2024)
81 M. subbrunnea Liu 315 PP037948 – PP034083 China Liu et al. (2024)
82 M. subbrunnea Liu 453 PP037951 PP034077 PP034086 China Liu et al. (2024)
83 M. subpura Liu 10 PP037943 – PP034078 China Liu et al. (2024)
84 M. subpura Liu 489 PP037954 – PP034089 China Liu et al. (2024)
85 M. subulata FFAAS0419 ON427735 ON468460 ON468473 China Liu et al. (2022)
86 M. subulata FFAAS0420 ON427736 ON468461 ON468474 China Liu et al. (2022)
87 M. subulata FFAAS0423 ON427737 ON468462 ON468475 China Liu et al. (2022)
88 M. subulata FFAAS0426 ON427738 ON468463 ON468476 China Liu et al. (2022)
89 M. variispora Liu 129 PP037945 – PP034080 China Liu et al. (2024)
90 M. variispora Liu 369 PP037949 – PP034084 China Liu et al. (2024)
91 M. variispora Liu 370 PP037950 – PP034085 China Liu et al. (2024)
92 M. violocea-ardesiaca Liu 475 PP037952 – PP034087 China Liu et al. (2024)
93 M. violocea-ardesiaca Liu 477 PP037953 – PP034088 China Liu et al. (2024)
94 M. yuezhuoi FFAAS0344 MW581490 MW868166 MW882249 China Liu et al. (2021)
95 M. yuezhuoi FFAAS0345 MW581491 MW868169 MW882250 China Liu et al. (2021)
96 M. yuezhuoi FFAAS0346 MW581492 MW868168 MW882251 China Liu et al. (2021)
97 M. yuezhuoi FFAAS0347 MW581493 MW868167 MW882252 China Liu et al. (2021)

Remarks: New generated sequences are emphasized in bold; “-” show missing sequence.
Figure 1. 

Bayesian Inference tree based on concatenated ITS+RPB1+tef1-α dataset. Only branch nodes with both Maximum Likelihood bootstrap support values (BS) above 75% and Bayesian posterior probabilities (BPP) exceeding 0.95 are indicated. New taxonomic groups are marked in red.

According to the phylogenetic tree in Fig. 1 10 well-supported clades were identified. The three new species were located in Clade 3, Clade 5, and Clade 10, respectively, each forming an independent lineage with strong statistical support (BS/BPP = 100/1.00). Among the three clades, Clade 3 was morphologically characterized by the absence of pleurocystidia, and 6 species included in the clade: M. rubrofusca, M. shengshanensis Z.W. Liu, Y.P. Ge & Q. Na, M. pearsoniana Dennis ex Singer, M. violocea-ardesiaca Shun Liu & Biao Zhu, M. subulata Z.W. Liu, Y.P. Ge & Q. Na, and M. lucisnieblae Cortés-Pérez, Racm.-Cruz & Guzm.-Dáv. In Clade 3, M. rubrofusca and M. violocea-ardesiaca were identified as the two most closely related taxa but with low support (BS/BPP = 81/--). There were 3 species in Clade 5, M. brunneocystidiata, M. pelianthina (Fr.) Quél., and M. lammiensis Harmaja, the species in the clade have colored lamellae margins and cystidia with colored contents, M. brunneocystidiata showed a close phylogenetic relationship with M. pelianthina (BS/BPP = 93/0.98) than M. lammiensis. Clade 10 merely contained 2 species, M. roseopurpurea and M. yuezhuoi Z.W. Liu, Y.P. Ge & Q. Na, each formed a distinct lineage with statistical support (BS/BPP = 100/1.00). Furthermore, Clades 1, 2, 4, 6, 7, 8, and 9 also each formed separate lineages with high bootstrap and posterior probability values.

Taxonomy

Mycena brunneocystidiata J.W. Guo, Z.W. Liu, Y.P. Ge & Q. Na, sp. nov.

MycoBank No: 860072
Figs 2, 3, 4, 5

Diagnosis.

Pileus brown. Lamellae densely covered with dark brown dots, margin brown. Cheilocystidia, pleurocystidia, caulocystidia and terminal cells of stipitipellis with brownish contents. Differ from M. lammiensis by wider basidiospores (width > 4 μm) and fusiform caulocystidia.

Figure 2. 

Basidiomata of Mycena brunneocystidiata. A–D. Collection FFAAS3400, holotype; E–I. Collection FFAAS3401. Scale bars: 20 mm (A–B, E, G–H); 1 mm (C, F); 5 mm (D, I). Photographs (A–I) by Jingwen Guo and Yupeng Ge.

Holotype.

China • Heilongjiang Province, Mudanjiang City, Mudanfeng National Forest Park, 42°45'74"N, 128°14'41"E, 22 August 2024, Jingwen Guo, Tian Wang, Qin Na, Zengcai Liu, Ruipeng Liu, Pengyu Du, Ying Yu, and Yupeng Ge leg., FFAAS3400 (collection no. NJ 6538).

Etymology.

Name refers to the cheilocystidia, pleurocystidia, and caulocystidia with brown contents.

Description.

Pileus 10–36 mm in diam., plano-convex, with slightly umbo at center, margin revolute, wavy, cracked at mature; *Drab (XLVI17′′′′) at center, gradually towards margin to PaMid Vinaceous-Drab (XLV5′′′′f), Pale Drab-Gray (XLVI17′′′′f) to *Drab-Gray (XLVI17′′′′d), margin *Drab (XLVI17′′′′); striate *Hair Brown (XLVI17′′′′i), towards the center up to 1/2–2/3 diam.. Context White (LIII), 1.0 mm thick, fragile. Lamellae subdecurrent, 23–27 reaching the stipe, 1–3 tiers of lamellulae, White (LIII), densely covered with Deep Brownish Drab (XLV9′′′′i) dots, irregularly intervenose, stretching downward to 2/3–3/4 of the width of lamellae, edge entirely Deep Brownish Drab (XLV9′′′′i), wavy. Stipe 24–44 × 2–4 mm, central, cylindrical; apex to middle *Smoke Gray (XLVI21′′′′d) to Pale Smoke Gray (XLVI21′′′′f), base *Ecru-Drab (XLVI13′′′′d) to *Drab-Gray (XLVI17′′′′d), hollow, fragile, apex with Light Drab (XLVI17′′′′b) to *Drab (XLVI17′′′′) striates, sparse White (LIII) pubescent at base. Odor and taste not distinctive.

Basidiospores (60/3/2) (5.4)5.6–6.3–6.9(7.1) × (2.6)2.9–3.2–3.5(4.0) μm [Q = (1.72)1.77–2.16(2.19), Qm = 1.97 ± 0.09] [holotype (40/2/1) 6.0–6.5–6.9(7.1) × 3.0–3.2–3.5 μm [Q = (1.83)1.89–2.15(2.18), Qm = 2.00 ± 0.08], narrowly ellipsoid to cylindrical, colorless, smooth (1000×), thin-walled, amyloid. Basidia clavate, 16–21 × 5–7 μm, hyaline, thin-walled, 4-spored, sterigmata 2–3 μm in length. Cheilocystidia fusiform with tapered apices, 36–62 × 9–14 μm (Fig. 3G–J, Fig. 4D1), acicular to lanceolate, 46–84 × 10–17 μm (Fig. 3M–P, Fig. 4D2), with brownish contents, thin-walled, smooth. Pleurocystidia similar to cheilocystidia, 39–72 × 8–15 μm, with brownish contents, thin-walled, smooth. Pileipellis a cutis composed of cylindrical cells, 32–121 × 3–6 μm, smooth, thin-walled; terminal cells cylindrical, apex tapering, 34–87 μm in length, apex 1–3 μm, base 3–7 μm, thin-walled, hyaline. Hypodermium formed by fusiform to subglobose hyphae, 20–72 × 12–39 μm, thin-walled, hyaline. Lamellar trama subregular, dextrinoid. Stipitipellis a cutis composed of cylindrical hyphae, 9–18 μm in diam., smooth, thin-walled; terminal cells fusiform or cylindrical, apex tapering, 31–72 × 4–8 μm, with pale brownish contents, thin-walled, smooth; caulocystidia fusiform, 33–70 × 7–13 μm, with pale brownish contents, thin-walled, smooth. Clamps present in all tissues.

Figure 3. 

Microscopic features of Mycena brunneocystidiata (FFAAS3400, holotype). A–E. Basidiospores; F. Basidia; G–J. Fusiform cheilocystidia; K–L. Acicular to lanceolate cheilocystidia; M–P. Fusiform pleurocystidia; Q–R. Acicular to lanceolate pleurocystidia; S. Pileipellis and hypodermium; T. Lamellar trama; U. Stipitipellis and caulocystidia. Scale bars: 5 μm (A–E); 10 μm (F); 25 μm (G–R); 20 μm (S–U). Structures (A–E) were rehydrated in 5% KOH aqueous solution, (G–R, U) were rehydrated in sterile water and (F, S–T) were stained in 1% Congo red aqueous solution.

Figure 4. 

Morphological features of Mycena brunneocystidiata (FFAAS3400, holotype). A. Basidiomata; B. Basidiospores; C. Basidia; D1. Fusiform cheilocystidia; D2. Acicular to lanceolate cheilocystidia; E1. Fusiform pleurocystidia; E2. Acicular to lanceolate pleurocystidia; F. Stipitipellis and caulocystidia; G. Pileipellis and hypodermium. Scale bars: 20 mm (A); 5 μm (B); 10 μm (C); 25 μm (D1–G). Drawings by Jingwen Guo.

Habit and habitat.

Scattered on the litter layer in Acer mono Maxim., Larix gmelinii (Ruprecht) Kuzeneva, Pinus koraiensis Siebold et Zuccarini, and Quercus mongolica Fischer ex Ledebour mixed forests during summer and autumn.

Known distribution.

Heilongjiang Province, Jilin Province, China.

Additional material examined.

China • Jilin Province, Yanbian Korean Autonomous Prefecture, Antu County, Erdaobaihe Town, Back Mountain of Changbai Mountain Natural History Museum, 42°46'43"N, 128°14'49"E, 17 August 2021, Zewei Liu, Qin Na, Shixin Wang, and Yupeng Ge leg., FFAAS3401 (collection no. MY 0611).

Notes.

Mycena brunneocystidiata is considered to be a distinct species of sect. Calodontes subsect. Marginatae J. E. Lange on account of its lamellae margins brown and cheilocystidia, pleurocystidia, and caulocystidia with brown contents (Maas Geesteranus 1992a, 1992b). In the subsection, M. lammiensis and M. pelianthina share the same colored lamellae margins, cheilocystidia, and pleurocystidia with purple-brown contents, but M. lammiensis differs in having larger basidiospores (7.5–9.0 × 4.0–5.0 μm) and cylindrical caulocystidia, while M. pelianthina is identified by purple-brown pileus and lacking caulocystidia (Harmaja 1985; Robich 2003; Harder et al. 2010; Aronsen and Læssøe 2016). Mycena shengshanensis resembles M. brunneocystidiata in growing on the humus layer of Larix gmelinii and having light purple-brown to brown pileus, but differs in colorless, clavate, and thick-walled cheilocystidia (Liu et al. 2022).

The main shape of cheilocystidia and pleurocystidia in FFAAS3401 is fusiform with tapered apices (Fig. 5A–E, K–O), but in specimen FFAAS3400, acicular to lanceolate cheilocystidia and pleurocystidia can be observed occasionally, which are larger than the fusiform ones (Fig. 5F–J, P–T).

Figure 5. 

Morphological features of the Cheilocystidia and pleurocystidia of Mycena brunneocystidiata. A–E. Cheilocystidia of FFAAS3401; F–J. Cheilocystidia of FFAAS3400, holotype; K–O. Pleurocystidia of FFAAS3401; P–T. Pleurocystidia of FFAAS3400, holotype. Scale bars: 25 μm (A–E, K–O); 30 μm (F–J, P–T). Drawings by Jingwen Guo.

Mycena roseopurpurea J.W. Guo, Z.W. Liu, Y.P. Ge & Q. Na, sp. nov.

MycoBank No: 860104
Figs 6, 7, 8

Diagnosis.

Pileus light pinkish-purple, light brown at center, hygrophanous when old. Cheilocystidia and pleurocystidia fusiform with tapered apices, thin-walled. Differ from M. subulata by lacking pleurocystidia and having acicular to lanceolate, thick-walled cheilocystidia.

Figure 6. 

Basidiomata of Mycena roseopurpurea. A–C. Collection FFAAS3405; D–G. Collection FFAAS3404, holotype; H–J. Collection FFAAS3403; K–L. Collection FFAAS3402. Scale bars: 10 mm (A–E, G, I–J); 5 mm (F, H, K–L). Photographs (A–L) by Qin Na.

Holotype.

China • Heilongjiang Province, Yichun City, Liangshui National Nature Reserve, 47°12'74"N, 128°52'86"E, 21 August 2021, Zewei Liu, Qin Na, Shixin Wang, and Yupeng Ge leg., FFAAS3404 (collection no. MY 0660).

Figure 7. 

Microscopic features of Mycena roseopurpurea (FFAAS3404, holotype). A–E. Basidiospores; F. Basidia; G–L. Cheilocystidia; M–R. Pleurocystidia; S. Pileipellis and hypodermium; T. Lamellar trama; U. Stipitipellis and caulocystidia. Scale bars: 5 μm (A–E); 15 μm (F); 30 μm (G–U). Structures (A–E) were rehydrated in 5% KOH aqueous solution and (F–U) were stained in 1% Congo red aqueous solution.

Etymology.

Name refers to light pinkish-purple pileus.

Figure 8. 

Morphological features of Mycena roseopurpurea (FFAAS3404, holotype). A. Basidiomata; B. Basidia; C. Basidiospores; D. Cheilocystidia; E. Pleurocystidia; F. Stipitipellis and caulocystidia; G. Pileipellis and hypodermium. Scale bars: 10 mm (A); 15 μm (B); 5 μm (C); 30 μm (D–G). Drawings by Jingwen Guo.

Description.

Pileus 13–22 mm in diam., oblate hemispherical to plano-convex, with slightly depressed at center, margin wavy, occasionally cracked at mature; *Ecru-Drab (XLVI13′′′′d), Light Drab (XLVI17′′′′b) to *Drab-Gray (XLVI17′′′′d) at center, gradually towards margin to Pale Verbena Violet (XXXVI55′′f), Light Pinkish Lilac (XXXVII65′′f), Pale Grayish Vinaceous (XXXIX9′′′f), margin White (LIII); striate Pale Smoke Gray (XLVI21′′′′f), Pale Ecru-Drab (XLVI13′′′′f) to *Ecru-Drab (XLVI13′′′′d), towards the center up to 1/3 diam.; surface dry, hygrophanous when old. Context White (LIII), 1.0 mm thick, fragile. Lamellae sinuate to subdecurrent, 21–28 reaching the stipe, 1–3 tiers of lamellulae, White (LIII), irregularly intervenose, stretching downward to 1/3–3/4 of the width of lamellae, edge concolorous, wavy, serrated. Stipe 32–57 × 1–4 mm, central, cylindrical; apex Pale Drab-Gray (XLVI17′′′′f), Pale Vinaceous-Drab (XLV5′′′′d), Pale Brownish Drab (XLV9′′′′d), lower part Pale Purple-Drab (XLV1′′′′d), Light Purple-Drab (XLV1′′′′b) to *Ecru-Drab (XLVI13′′′′d), hollow, fragile; sparse White (LIII) pubescent at base. Odor and taste raphanoid.

Basidiospores (100/5/4) (5.6)5.9–6.7–7.4(7.8) × 3.0–3.5–3.9(4.1) μm [Q = (1.70)1.72–2.10, Qm = 1.90 ± 0.09] [holotype (40/2/1) (6.0)6.2–6.7–7.4(7.7) × 3.0–3.5–3.9(4.1) μm, Q = (1.71)1.77–2.12, Qm = 1.93 ± 0.11], narrowly ellipsoid to cylindrical, colourless, smooth (1000×), thin-walled, amyloid. Basidia clavate, 18–31 × 4–7 μm, hyaline, thin-walled, 4-spored, sterigmata 2–4 μm in length. Cheilocystidia fusiform, subfusiform, with apices tapered, 35–97 × 7–18 μm, thin-walled, smooth. Pleurocystidia similar to cheilocystidia, 30–80 × 6–18 μm, thin-walled, smooth. Pileipellis a cutis composed of cylindrical cells, 29–81 × 3–11 μm, smooth, thin-walled; terminal cells cylindrical, apex tapering, 20–99 μm in length, apex 2–4 μm, base 2–9 μm, thin-walled, hyaline. Hypodermium formed by fusiform to subglobose hyphae, 20–76 × 6–37 μm, thin-walled, hyaline. Lamellar trama subregular, dextrinoid. Stipitipellis a cutis composed of cylindrical hyphae, 4–17 μm in diam., smooth, thin-walled; caulocystidia fusiform, clavate, sometimes utriform, 23–66 × 6–20 μm, thin-walled, smooth. Clamps present in all tissues.

Habit and habitat.

Scattered on the litter layers in Betula platyphylla Suk., Larix gmelinii, Pinus koraiensis, and P. syluestriformis (Takenouchi) T.Wang ex Cheng mixed forests during summer and autumn.

Known distribution.

Heilongjiang Province, Jilin Province, China.

Additional material examined.

China • Heilongjiang Province, Yichun City, Liangshui National Nature Reserve, 47°12'74"N, 128°52'56"E, 20 August 2021, Zewei Liu, Qin Na, Shixin Wang, and Yupeng Ge leg., FFAAS3403 (collection no. MY 0635); • same location, 21 August 2021, Zewei Liu, Qin Na, Shixin Wang, and Yupeng Ge leg., FFAAS3405 (collection no. MY 0668). • Jilin Province, Yanbian Korean Autonomous Prefecture, Antu County, Erdaobaihe Town Beauty Pine Forest, 42°45'74"N, 128°14'41"E, 18 August 2021, Zewei Liu, Qin Na, Shixin Wang, and Yupeng Ge leg., FFAAS3402 (collection no. MY 0625).

Notes.

Mycena subulata, originally described from Heilongjiang province in China, can be easily mistaken for M. roseopurpurea due to morphological similarity, and growing on the humus layer of mixed forests of Larix gmelinii and Pinus koraiensis, but M. subulata is distinguished by acicular to lanceolate and thick-walled cheilocystidia (Liu et al. 2022). Mycena pearsoniana, while having similar stipe color and lacking pleurocystidia, differs by subfusiform or clavate cheilocystidia and inamyloid basidiospores (Smith 1947; Dennis 1970; Kost 2002; Harder et al. 2012; Aronsen and Læssøe 2016; Na 2019; Kwon et al. 2020). Mycena dura Maas Geesteranus & Hauskn. and M. subcorticalis (Cooke & Massee) Sacc., reported from Europe and southern Australia, resemble M. roseopurpurea; however, M. dura is distinctively characterized by its growth in grasslands and white stipe, and M. subcorticalis is distinguished by inamyloid basidiospores and gelatinized pileipellis (Saccardo 1891; Maas Geesteranus and Hausknecht 1994; Grgurinovic 2003; Olariaga et al. 2015). Mycena pura differs by having light purple to purplish-red pileus and pinkish-purple to purple stipe (Maas Geesteranus 1992a, 1992b; Perry 2002; Robich 2003; Thormann et al. 2006; Aronsen and Læssøe 2016; Na 2019; Okon et al. 2022; Liu 2023).

Mycena rubrofusca J.W. Guo, Z.W. Liu, Y.P. Ge & Q. Na, sp. nov.,

MycoBank No: 860105
Figs 9, 10, 11

Diagnosis.

Pileus light reddish-brown to light grayish-brown, near margin hygrophanous. Pleurocystidia and caulocystidia absent. Differ from M. polycystidiata Z.W. Liu, Y.P. Ge, L. Zou & Q. Na by having pleurocystidia and caulocystidia.

Figure 9. 

Basidiomata of Mycena rubrofusca. A–E. Collection FFAAS3406, holotype; F–J. Collection FFAAS3407. Scale bars: 15 mm (A–B, F–G); 1 mm (C–D); 5 mm (E, H–J). Photographs (A–J) by Jingwen Guo and Yupeng Ge.

Holotype.

China • Heilongjiang Province, Mudanjiang City, Mudanfeng National Forest Park, 42°45'74"N, 128°14'41"E, 22 August 2024, Jingwen Guo, Tian Wang, Qin Na, Zengcai Liu, Ruipeng Liu, Pengyu Du, Ying Yu, and Yupeng Ge leg., FFAAS3406 (collection no. NJ 6508).

Figure 10. 

Microscopic features of Mycena rubrofusca (FFAAS3406, holotype). A–E. Basidiospores; F. Basidia; G–R. Cheilocystidia; S. Pileipellis and hypodermium; T. Lamellar trama; U. Stipitipellis. Bars: 5 μm (A–E); 15 μm (F, T, U); 25 μm (G–R); 30 μm (S). Structures (A–E) were rehydrated in 5% KOH aqueous solution and (F–U) were stained in 1% Congo red aqueous solution.

Etymology.

Name refers to the light reddish-brown, light grayish brown to brown pileus.

Figure 11. 

Morphological features of Mycena rubrofusca (FFAAS3406, holotype). A. Basidiomata; B. Basidiospores; C. Basidia; D. Cheilocystidia; E. Stipitipellis; F. Pileipellis and hypodermium. Scale bars: 15 mm (A, C, E–F); 5 μm (B); 25 μm (D). Drawings by Jingwen Guo.

Description.

Pileus 17–26 mm in diam., hemispherical, plano-convex at mature, margin wavy; Light Drab (XLVI17′′′′b) at center, gradually towards margin to Light Vinaceous-Fawn (XL13′′′d), Pale Ecru-Drab (XLVI13′′′′f) to *Drab-Gray (XLVI17′′′′d), margin Light Drab (XLVI17′′′′b) to *Drab (XLVI17′′′′); striate *Drab (XLVI17′′′′), towards the center up to 1/8–1/7 diam.; surface moist, near margin hygrophanous. Context White (LIII), 1.0 mm thick, fragile. Lamellae adnexed or subdecurrent, approximately 38 reaching the stipe, 1–3 tiers of lamellulae, White (LIII), irregularly intervenose, stretching downward to 2/3–3/4 of the width of lamellae, edge concolorous, wavy. Stipe 44–45 × 2–4 mm, central, cylindrical; apex to lower part Vinaceous-Drab (XLV5′′′′), Light Brownish Drab (XLV9′′′′b) to *Hair brown (XLVI17′′′′i), base Light Grayish Olive (XLVI21′′′′b), hollow, fragile; base swollen, sparse White (LIII) pubescent at base. Odor and taste faintly raphanoid.

Basidiospores (40/2/2) (5.1)6.2–7.4–8.5(8.8) × (3.8)4.0–4.6–5.5(5.7) μm [Q = (1.52)1.55–1.84(1.89), Qm = 1.69 ± 0.10] [holotype (20/1/1) (6.8)7.2–7.6–8.3(8.5) × (3.8)4.1–4.6–4.8(5.0) μm [Q = 1.60–1.81, Qm = 1.70 ± 0.08], ellipsoid to narrowly ellipsoid, colourless, smooth (1000×), thin-walled, amyloid. Basidia clavate, 16–28 × 5–8 μm, hyaline, thin-walled, 4-spored, sterigmata 2–4 μm in length. Cheilocystidia utriform, clavate, 24–67 × 8–23 μm, thin-walled, smooth. Pleurocystidia absent. Pileipellis a cutis composed of cylindrical cells, 28–76 × 5–15 μm, smooth, thin-walled; terminal cells cylindrical, apex tapering, 33–75 μm in length, apex 2–9 μm, base 3–11 μm, thin-walled, hyaline. Hypodermium formed by fusiform to subglobose hyphae, 26–84 × 11–34 μm, thin-walled, hyaline. Lamellar trama subregular, dextrinoid. Stipitipellis a cutis composed of cylindrical hyphae, 6–16 μm in diam, smooth, thin-walled; projecting terminal cells cylindrical or fusiform, 31–73 × 4–9 μm, thin-walled, smooth, hyaline; caulocystidia absent. Clamps present in all tissues.

Habit and habitat.

Scattered on the litter layers in Fraxinus mandschurica Rupr., Pinus koraiensis, and Tilia amurensis Rupr. mixed forests during summer and autumn.

Known distribution.

Heilongjiang Province, China.

Additional material examined.

China • Heilongjiang Province, Hegang City, Taipinggou National Nature Reserve, 48°12'43"N, 130°66'04"E, 3 September 2021, Zewei Liu, Qin Na, Shixin Wang, and Yupeng Ge leg., FFAAS3407 (collection no. MY 0793).

Notes.

Mycena polycystidiata is the closest species to M. rubrofusca because it has a light grayish-brown to brown pileus and stipe, and utriform cheilocystidia, but M. polycystidiata has pleurocystidia and caulocystidia (Liu et al. 2022). Mycena pura, a widely distributed species in the North temperate zone, differs from M. rubrofusca by its fusiform, clavate, or subglobose cheilocystidia and pleurocystidia, and clavate or conical caulocystidia (Maas Geesteranus 1992a, 1992b; Perry 2002; Robich 2003; Thormann et al. 2006; Aronsen and Læssøe 2016; Na 2019; Okon et al. 2022; Liu 2023). Due to the light gray-brown to light brown pileus, M. variispora Shun Liu & Biao Zhu and M. subbrunnea Shun Liu & Biao Zhu are difficult to distinguish from M. rubrofusca, but they can be distinguished by the occurrence in mixed forests of Larix gmelinii and Betula platyphylla, purple lamellae, and clavate, thick-walled cheilocystidia (Liu et al. 2024). Mycena pearsoniana differs from M. rubrofusca in having purple pileus and clavate or fusiform caulocystidia (Smith 1947; Dennis 1970; Kost 2002; Harder et al. 2012; Aronsen and Læssøe 2016; Na 2019; Kwon et al. 2020). Mycena rufobrunnea Z.W. Liu, Y.P. Ge & Q. Na and M. vinacea Cleland, which were reported from northeastern China and southern Australia, respectively, both resemble M. rubrofusca in having utriform cheilocystidia (Cleland 1931; Liu et al. 2022). However, both of them also have clavate cheilocystidia; M. rufobrunnea is distinguished by a light brown to reddish-brown pileus, whereas M. vinacea is characterized by cylindrical pleurocystidia and basidiospores (Cleland 1931; Liu et al. 2022).

Key to 20 species belonging to Mycena sect. Calodontes in China

1 Lamellae edge purplish-brown or brown 2
– Lamellae edge white 3
2 Caulocystidia present M. brunneocystidiata
– Caulocystidia absent M. pelianthina
3 Pileus pink M. rosea
– Pileus not pink 4
4 Cheilocystidia thick-walled 5
– Cheilocystidia thin-walled 10
5 Pileus lilac M. variispora
– Pileus not lilac 6
6 Lamellae surfaces white 7
– Lamellae surfaces lilac 9
7 Cheilocystidia acicular to lanceolate M. subulata
– Cheilocystidia clavate 8
8 Stipe light grayish-brown to purplish-brown M. shengshanensis
– Stipe white to pinkish-yellow M. subbrunnea
9 Lamellae adnexed M. subpura
– Lamellae subfree M. violocea-ardesiaca
10 Basidiospores inamyloid 11
– Basidiospores amyloid 12
11 Pileus white to cream-colored M. glabera
– Pileus pinkish-purple to purple M. pearsoniana
12 Cheilocystidia utriform 13
– Cheilocystidia fusiform or clavate 15
13 Pleurocystidia present M. polycystidiata
– Pleurocystidia absent 14
14 Caulocystidia present M. rufobrunnea
– Caulocystidia absent M. rubrofusca
15 Pileus white 16
– Pileus not white 17
16 Lamellae adnate M. cinereoalba
– Lamellae adnexed M. subaquosa
17 Pleurocystidia present 18
– Pleurocystidia absent M. yuezhuoi
18 Stipe purple M. pura
– Stipe not purple 19
19 Basidiospores (5.6)5.9–7.4(7.8) × 3.0–3.9(4.1) μm, elongated ellipsoid to cylindrical M. roseopurpurea
– Basidiospores (8.3)8.6–10.8(11.5) × (5.3)5.4–6.3(6.4) μm, ellipsoid to elongated ellipsoid M. roseolamellata

Discussion

The comprehensive morphological characteristics of sect. Calodontes cannot be currently shown by the taxonomic systems (Harder et al. 2010, 2013; Liu 2023). Three subsections were proposed based on the amyloid reaction of basidiospores, the presence of colored contents in cheilocystidia and pleurocystidia, and the presence of pleurocystidia, namely subsect. Purae Konrad & Maubl., subsect. Marginatae, and subsect. Violacellae Singer ex Maas Geest. (Maas Geesteranus 1992a, 1992b). In this study, M. brunneocystidiata was assigned to subsect. Marginatae due to its amyloid basidiospores and cheilocystidia, pleurocystidia, and caulocystidia with brown contents; M. roseopurpurea was assigned to subsect. Purae owing to its amyloid basidiospores and having colorless cheilocystidia and pleurocystidia. However, M. rubrofusca, which was assigned to subsect. Violacellae due to its lack of pleurocystidia, cannot be placed in this subsection on account of its amyloid basidiospores. Notably, Harder et al. (2012) and Pérez-De-Gregorio (2024) reported that M. pearsoniana was assigned to subsect. Violacellae, which was defined by Maas Geesteranus based on inamyloid basidiospores, but its basidiospores showed inamyloid to weakly amyloid basidiospores after 40 minutes (Maas Geesteranus 1992a, 1992b; Harder et al. 2012; Pérez-De-Gregorio 2024). The subsections and species were also not supported by the multi-locus (ITS+RPB1+tef1-α) analyses, subsect. Marginatae was monophyletic, and subsect. Purae and subsect. Violacellae were polyphyletic (Harder et al. 2010, 2012, 2013; Chew et al. 2014; Liu 2023). Species clustered into a single clade with high support in phylogenetic analyses showed significant morphological differences. Mycena subbrunnea, M. variispora, M. subpura Shun Liu & Biao Zhu, and 7 phylogenetic species from the M. pura complex were gathered in the same clade, but the shape and wall thickness of cheilocystidia, and the presence or absence of pleurocystidia were different (Maas Geesteranus 1992a, 1992b; Perry 2002; Robich 2003; Thormann et al. 2006; Harder et al. 2010, 2013; Liu 2023; Liu et al. 2024).

According to previous literature, species of sect. Calodontes predominantly inhabit coniferous forests or coniferous-broadleaved mixed forests in Europe, North America, Asia, and Northern Africa, mainly within the Northern Hemisphere (Maas Geesteranus 1992a, 1992b; Robich 2003; Harder et al. 2010, 2013; Chew et al. 2014; Aronsen and Læssøe 2016; Cooper et al. 2018; Cortés-Pérez et al. 2023). Only a few occur in grasslands or in broadleaved forests of the Southern Hemisphere, southern Oceania (Grgurinovic 2003). In China, species of sect. Calodontes are predominantly found in the litter layer of coniferous-broadleaved mixed forests in Northeast China, and in coniferous forests in East, Southwest, Northwest, and Southeast China (Bau et al. 2019; Na 2019; Liu et al. 2021, 2022, 2024; Liu 2023; Fan et al. 2024; Xiao et al. 2025). Most species of sect. Calodontes, including the three new species in this study, are distributed in low-altitude regions (< 1000 m), with a few recorded in mid-altitude areas (1000–3500 m), and no species have been reported from high-altitude regions (3500–5000 m) (Maas Geesteranus 1992a, 1992b; Maas Geesteranus and de Meijer 1997; Grgurinovic 2003; Cooper et al. 2018; Bau et al. 2019; Na 2019; Liu 2023; Gao et al. 2024).

Acknowledgements

We thank Prof. Li Zou (Northeast Forest University), Dr Zengcai Liu (Northeast Forest University), Dr Shixin Wang (Northeast Forest University), Mr Ruipeng Liu (Northeast Forest University), Mr Pengyu Du (Northeast Forest University), Mrs Ying Yu (Northeast Forest University), Mrs Tian Wang (Ludong University), and Mrs Guanyu Qiu (Ludong University) for the kind help during field work. We sincerely thank the reviewers for their corrections and suggestions on how to improve our work.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of AI

No use of AI was reported.

Funding

This study was supported by the National Natural Science Foundation of China (grant no. 32200008), the Census and Collection of Edible Mushroom Germplasm Resources in Fujian Province (grant no. fjmacro-001), the 5511 Collaborative innovation project of Fujian Province (grant no. XTCXGC2021007), the Central Public-Interest Scientific Institution Basal Research Fund (grant no. GYZX200203), the Natural Science Foundation of Fujian Province (grant no. 2023J01379), and the Biodiversity investigation, observation and assessment program of Ministry of Ecology and Environment of China (grant no. 2110404 and 2019-2023).

Author contributions

Conceptualization, J.W.G., Z.W.L., H.Z., Y.P.G. and Q.N.; molecular experiments and data analysis, J.W.G. and Z.W.L.; field investigation, H.Z. and Y.P.G.; writing—original draft preparation, J.W.G. and Q.N.; writing—review and editing, Q.N. and Y.P.G; funding acquisition, H.Z. and Y.P.G. All authors have read and agreed to the published version of the manuscript.

Author ORCIDs

Jingwen Guo https://orcid.org/0009-0005-2779-7456

Zewei Liu https://orcid.org/0000-0002-2700-0831

Hui Zeng https://orcid.org/0000-0003-2025-844X

Yupeng Ge https://orcid.org/0000-0001-5754-201X

Qin Na https://orcid.org/0000-0001-8406-6389

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.

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Supplementary material

Supplementary material 1 

Fasta of Phylogentic tree

Jingwen Guo, Zewei Liu, Hui Zeng, Yupeng Ge, Qin Na

Data type: fas

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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