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Research Article
Four new species of Mycena sect. Calodontes (Agaricales, Mycenaceae) from northeast China
expand article infoZewei Liu‡, Yupeng Ge‡, Hui Zeng§|, Xianhao Cheng‡, Qin Na‡
‡ Ludong University, Yantai, China
§ Institute of Edible Mushroom, Fujian Academy of Agricultural Sciences, Fuzhou, China
| National and Local Joint Engineering Research Center for Breeding and Cultivation of Features Edible Mushroom, Fuzhou, China

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

Academic editor: Zai-Wei Ge
© 2022 Zewei Liu, Yupeng Ge, Hui Zeng, Xianhao Cheng, 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: Liu Z, Ge Y, Zeng H, Cheng X, Na Q (2022) Four new species of Mycena sect. Calodontes (Agaricales, Mycenaceae) from northeast China. MycoKeys 93: 23-56. https://doi.org/10.3897/mycokeys.93.86580
Open Access

Abstract

Species of Mycena sect. Calodontes are representative of the Mycena genus as a whole and are easily recognised by the pinkish, reddish, purplish to brownish pileus and larger basidiomata. Furthermore, the colour of the pileus in the species of sect. Calodontes often has a transition or changes in different stages and the combination of the colour of the pileus with cystidia and basidiospores can be used to recognise taxa within this section. To date, 19 species of Mycena sect. Calodontes have been reported worldwide. Including our recent description of M. yuezhuoi, five species of sect. Calodontes have been recorded in China. During examination of specimens collected in coniferous forests or mixed broadleaf-conifer forests in temperate regions of China, additional taxa assigned to sect. Calodontes were identified. Four new species are recognised, based mostly on characters of the pileus and cystidia. Phylogenetic analysis of sequence data from multiple DNA regions (ITS + rpb1 + tef1) supported the morphological evidence. Here, we propose M. polycystidiata, M. rufobrunnea, M. shengshanensis and M. subulata as new species in Mycena sect. Calodontes. Morphological descriptions, line drawings, habitat photos and comparisons with closely-related taxa are provided. A key to the 23 known species of sect. Calodontes is presented.

Keywords

coniferous forest, new taxa, phylogeny, saprobic, taxonomy

Introduction

Mycena sect. Calodontes (Fr. ex Berk.) Quél. comprises the taxa in Mycena (Pers.) Roussel with a pinkish, reddish, purplish to brownish and mostly hygrophanous pileus, interveined lamellae, smooth cheilocystidia and pleurocystidia (if present) and mostly amyloid spores (Fries 1821; Berkeley 1836; Maas Geesteranus 1992a, 1992b; Harder et al. 2010). The group was initially proposed as Agaricus trib. Clitocybe subtrib. Calodontes Fries consisting of six species, then elevated to section rank within Agaricus subgen. Clitocybe Fr. ex Berk. and finally assigned to Mycena in 1872 (Fries 1821; Berkeley 1836; Quélet 1872). To date, 19 species are known, mostly from Europe and North America, but five species have been described from Asia (specifically, China, India and Peninsular Malaysia) (Smith 1947; Maas Geesteranus 1980; Perry 2002; Grgurinovic 2003; Robich 2003; Chew et al. 2014; Aravindakshan and Manimohan 2015; Aronsen and Læssøe 2016; Na 2019; Liu et al. 2021).

The circumscription of subsections within sect. Calodontes is problematic. Initially, Smith (1947) divided sect. Calodontes into two subsections, Granulatae and Ciliatae, according to whether the cheilocystidia were smooth or not, but this system was not widely adopted because most of the species have previously been classified in other sections of Mycena on account of the red, yellow or orange basidiomata, coloured lamellar edge and echinulate or diverticulate cheilocystidia, pleurocystidia, pileipellis or stipitipellis (Maas Geesteranus 1980, 1992a, 1992b; Perry 2002; Grgurinovic 2003; Robich 2003; Harder et al. 2010; Chew et al. 2014). Based on the lamellar edge colour, spore amyloid reaction and cystidial features, a widely accepted subsectional classification of sect. Calodontes was proposed by Maas Geesteranus (1980) and subsequent taxonomists (Grgurinovic 2003; Harder et al. 2010; Chew et al. 2014). The primary criteria for segregation were dominated by microcharacters: subsect. Purae (Konrad & Maubl.) Maas Geest. with amyloid spores and colourless pleuro- and cheilocystidia; subsect. Violacellae Sing. ex Maas Geest. with inamyloid spores and no pleurocystidia; and subsect. Marginatae J.E. Lange with amyloid spores and pleurocystidia and cheilocystidia with purplish-brown contents (Maas Geesteranus 1980; Grgurinovic 2003; Harder et al. 2010; Chew et al. 2014). Although the infrasectional classification of Maas Geesteranus (1980) is generally accepted, phylogenetic analyses have provided only weak support because subsect. Purae and subsect. Violacellae are polyphyletic (Perry 2002; Grgurinovic 2003; Robich 2003; Harder et al. 2010, 2012, 2013; Chew et al 2014; Na 2019). In studying Mycena pura (Pers.) P. Kumm., the type of subsect. Purae, Maas Geesteranus (1992a, 1992b) proposed eight forma based on pileus colour. However, 11 clades have been resolved amongst materials collected from Europe and the Americas, which suggests that there may be additional undescribed taxa in subsect. Purae (Harder et al. 2013).

Including our recent description of M. yuezhuoi Z.W. Liu, Y.P. Ge & Q. Na from Kunyushan National Nature Reserve (Yantai, Shandong Province), five species of Mycena sect. Calodontes have been previously recorded in China (Li et al. 2015; Na 2019; Liu et al. 2021). In this paper, we propose an additional four new species classified in sect. Calodontes from the temperate zone of northeast China. The four new species share a unique set of striking morphological characters and contribute to an improved understanding of the classification of sect. Calodontes.

Materials and methods

Morphological observations

Thirteen fungal specimens were examined in this study, which were mainly collected in coniferous forests and some from mixed broadleaf-conifer forests in 2021. Macrocharacters were recorded from fresh specimens. Colour codes in descriptions follow those of Kornerup and Wanscher (1978). Microcharacters were observed from tissues sampled from dried specimens and rehydrated with 5% potassium hydroxide (KOH) and stained with Congo red (1% [w/v] aqueous solution), if necessary, using a Lab A1 microscope (Carl Zeiss AG, Jena, Germany). The amyloid reaction was tested with Melzer’s Reagent (Clémençon et al. 2004; Horak 2005). Twenty basidiospores were measured per specimen. For the holotype, 40 basidiospores from different basidiomata were selected for measurement. Basidiospore statistics are expressed as (a/b/c) (d)e-f-g (h) × (i)j-k–l (m) μm [Q = (n)o-p (q), Q = r ± s], where a–c represent a basidiospores of b basidiomata from c specimens measured; d and h are the minimum and maximum length (5% extremum), respectively, e and g indicate the range of values for the remaining 90% of the spores and f is the average length; width (i–m) and Q values (n–q) are expressed in a similar manner; and r and s are the average Q value and its standard deviation, respectively (Ge et al. 2021; Liu et al. 2021; Na et al. 2021; Na et al. 2022). The measurement of basidia, cystidia and other characters were each based on 20 observations. All specimens have been deposited in the Fungarium of the Fujian Academy of Agricultural Sciences (FFAAS).

DNA extraction, PCR, cloning and DNA sequencing

The Plant Genomic DNA Kit (CoWin Biosciences, Beijing, China) was used to isolate total genomic DNA from dried specimens in accordance with the manufacturer’s instructions. Three nuclear loci were sequenced, comprising the internal transcribed spacer (ITS), RNA polymerase II largest subunit (rpb1) and translation elongation factor-1 alpha (tef1). The primer pairs ITS1/ITS4, rpb1Mp_f1/rpb1Mp_r1 and tEFMp_f2/ tEFMp_r2 were selected to amplify ITS, rpb1 and tef1, respectively (White et al. 1990; Harder et al. 2013; Yu et al. 2020). The PCR reactions were performed in a total volume of 25 µl containing 2 µl DNA template, 1 µl for each primer, 8.5 µl nuclease-free H2O and 12.5 µl 2× Utaq PCR MasterMix (ZomanBio, Beijing, China). The PCR protocol for amplification of the ITS region was as follows: 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 (Na et al. 2022). The PCR protocol for amplification of the rpb1 and tef1 regions followed that of Harder et al. (2013): 94 °C for 60 s, then 10 cycles of 94 °C for 35 s, 53 °C for 45 s and 72 °C for 45 s; then 25 cycles of 94 °C for 35 s, 56 °C for 45 s, 72 °C for 45 s and final extension of 72 °C for 10 min. The PCR products were purified by gel electrophoresis or filter membrane and subjected to Sanger dideoxy sequencing by the Beijing Genomics Institute (Beijing, China).

Phylogenetic analysis

A combined ITS, rpb1 and tef1 dataset was analysed to infer relationships of the new taxa with other members of sect. Calodontes. We used sequences included in previous studies of sect. Calodontes and from members of the most closely-related section deposited in the GenBank database, which were mainly submitted by Harder et al. (2013), Osmundson et al. (2013), Chew et al. (2014) and Liu et al. (2021). For the analysis, representative species of Mycena sect. Supinae Konrad & Maubl., which is closely related to sect. Calodontes, were selected as the outgroup (Osmundson et al. 2013; Na and Bau 2019). Sequences for each DNA region (ITS, rpb1 and tef1) were aligned in MAFFT version 7 online and the aligned matrices were manually checked with BIOEDIT 7.2.5.0 (Hall 1999; Kuraku et al. 2013; Katoh et al. 2019). The best-fit substitution model for each gene partition was determined with MODELTEST 2.3, based on the Akaike Information Criterion (Posada and Crandall 1998). Maximum Likelihood (ML) analysis was conducted by raxmlGUI 2.09 (Edler et al. 2020). The phylogenetic analysis was performed by a single analysis with six partitions (ITS1, 5.8S, ITS2, rpb1 exons, tef1 exons, intron of rpb1 + introns of tef1), using the GTRGAMMA model and 1,000 rapid bootstrap (BS) replicates. For Bayesian Inference (BI), two runs of six chains were run for 15,000,000 generations and sampled every 10,000 generations by MrBayes 3.2.6. At the end of the run, the average deviation of split frequencies was 0.007821, ESS (effective sample size) was 1300.3 and the average Potential Scale Reduction Factor (PSRF) parameter values (excluding NA and > 10.0) = 1.000 and the “sump” and “sumt” commands were used to summarise sampled parameters with 25% burn-in (Ronquist and Huelsenbeck 2003).

Results

Phylogenetic relationships

The dataset consisted of 192 sequences, comprising 39 newly-generated sequences (13 ITS, 13 rpb1 and 13 tef1) and 153 sequences (61 ITS, 46 rpb1 and 46 tef1) downloaded from GenBank. In total, 74 accessions of 19 species were included in the dataset. Detailed information for all sequences is presented in Table 1. The aligned dataset contained 1459 nucleotide sites including gaps (229 sites for ITS1, 159 sites for 5.8S, 177 sites for ITS2, 55 sites for rpb1 exons, 295 sites for tef1 exons, 544 sites for intron of rpb1 + introns of tef1), of which 1146 were conserved, 257 were parsimony-informative and 56 were variable, but parsimony-uninformative. For Bayesian Inference (BI), the selected models for each DNA region of the concatenated dataset were as follows: HKY+G for ITS1 and intron of rpb1 + introns of tef1, JC for 5.8S and rpb1 exons, HKY+I+G for ITS2 and SYM+I+G for tef1 exons. The BI and ML analyses resulted in almost identical topologies; thus, the BI topology is presented as a master tree (Fig. 1).

Table 1.
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Specimens used in phylogenetic analysis and GenBank accession numbers.

No. Species Specimen voucher GenBank accession numbers Locality Reference
ITS rpb1 tef1
1. Mycena 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. arcangeliana 252b JF908401 – – Spain Osmundson et al. (2013)
6. M. arcangeliana 252f JF908402 – – Spain Osmundson et al. (2013)
7. M. cahaya ACL134 KF537248 – – Malaysia Chew et al. (2014)
8. M. cf. pura I CBH039 FN394588 KF723680 KF723634 Denmark Harder et al. (2010, 2013)
9. M. cf. pura II CBH105 FN394581 KF723671 KF723625 Denmark Harder et al. (2010, 2013)
10. M. cf. pura II CBH169 FN394579 KF723672 KF723626 Denmark Harder et al. (2010, 2013)
11. M. cf. pura II CBH366 FN394572 KF723673 KF723627 Denmark Harder et al. (2010, 2013)
12. M. cf. pura II CBH404 FN394566 KF723674 KF723628 Denmark Harder et al. (2010, 2013)
13. M. cf. pura III CBH019 FN394605 KF723675 KF723629 Denmark Harder et al. (2010, 2013)
14. M. cf. pura III CBH022 FN394574 KF723676 KF723630 Denmark Harder et al. (2010, 2013)
15. M. cf. pura III KK FN394606 KF723677 KF723631 Slovakia Harder et al. (2010, 2013)
16. M. cf. pura IV CBH410 FN394595 KF723667 KF723621 Denmark Harder et al. (2010, 2013)
17. M. cf. pura IV JV06979 FN394585 KF723668 KF723622 Denmark Harder et al. (2010, 2013)
18. M. cf. pura IV TL4571 FN394583 KF723669 KF723623 Denmark Harder et al. (2010, 2013)
19. M. cf. pura IV TL12786 FN394591 KF723670 KF723624 Sweden Harder et al. (2010, 2013)
20. M. cf. pura V CBH226 FN394604 KF723664 KF723618 Denmark Harder et al. (2010, 2013)
21. M. cf. pura V TL5614 FN394602 KF723666 KF723620 Denmark Harder et al. (2010, 2013)
22. M. cf. pura VI BAP132 FN394561 KF723660 KF723614 USA Harder et al. (2010, 2013)
23. M. cf. pura VIII CBH216 FN394598 KF723662 KF723616 Denmark Harder et al. (2010, 2013)
24. M. cf. pura VIII CBH402 FN394599 KF723663 KF723617 Denmark Harder et al. (2010, 2013)
25. M. cf. pura IX CBH166 FN394607 KF723701 KF723655 Denmark Harder et al. (2010, 2013)
26. M. cf. pura IX CBH358 FN394608 KF723702 KF723656 Denmark Harder et al. (2010, 2013)
27. M. cf. pura IX CBH367 KF913022 KF723703 KF723657 Denmark Harder et al. (2013)
28. M. cf. pura IX CBH371 KF913023 KF723704 KF723658 Denmark Harder et al. (2013)
29. M. cf. pura X BAP165A FN394563 KF723698 KF723652 USA Harder et al. (2010, 2013)
30. M. cf. pura XI CBH187 FN394564 KF723678 KF723632 Sweden Harder et al. (2010, 2013)
31. M. cf. pura XI CBH386 FN394565 KF723679 KF723633 Denmark Harder et al. (2010, 2013)
32. M. diosma CBH400 FN394617 KF723699 KF723653 Denmark Harder et al. (2010, 2013)
33. M. diosma LK1191/2000 FN394619 KF723700 KF723654 Germany Harder et al. (2010, 2013)
34. M. dura 10315 FN394560 KF723694 KF723648 Austria Harder et al. (2010, 2013)
35. M. lammiensis TUR165927 FN394552 KF723697 KF723651 Finland Harder et al. (2010, 2013)
36. M. meliigena 39 JF908423 – – Italy Osmundson et al. (2013)
37. M. meliigena 39d JF908429 – – Italy Osmundson et al. (2013)
38. M. pearsoniana CBH068 FN394614 KF723691 KF723645 Germany Harder et al. (2010, 2013)
39. M. pearsoniana JV06890 FN394612 KF723692 KF723646 Denmark Harder et al. (2010, 2013)
40. M. pearsoniana LK880/2002 FN394613 KF723693 KF723647 Germany Harder et al. (2010, 2013)
41. M. pelianthina CBH015 FN394549 KF723695 KF723649 Denmark Harder et al. (2010, 2013)
42. M. pelianthina CBH016 FN394547 KF723696 KF723650 Denmark Harder et al. (2010, 2013)
43. M. polycystidiata FFAAS0417 Holotype ON427731 ON468456 ON468469 China This study
44. M. polycystidiata FFAAS0418 ON427732 ON468457 ON468470 China This study
45. M. polycystidiata FFAAS0421 ON427733 ON468458 ON468471 China This study
46. M. polycystidiata FFAAS0422 ON427734 ON468459 ON468472 China This study
47. M. pseudocorticola 124a JF908386 – – Italy Osmundson et al. (2013)
48. M. pura IS10/11/2000 FN394611 – – USA Harder et al. (2010)
49. M. pura f. lutea DB2005/152 FN394603 – – Denmark Harder et al. (2010)
50. M. rosea UP2 FN394550 – – UK Harder et al. (2010)
51. M. rosea CBH097 FN394556 KF723681 KF723635 Denmark Harder et al. (2010, 2013)
52. M. rosea CBH383 FN394553 KF723682 KF723636 Denmark Harder et al. (2010, 2013)
53. M. rosea CBH409 FN394551 KF723683 KF723637 Germany Harder et al. (2010, 2013)
54. M. rosea TL12393 FN394555 KF723684 KF723638 Denmark Harder et al. (2010, 2013)
55. M. rosea TL12409 FN394557 KF723685 KF723639 Denmark Harder et al. (2010, 2013)
56. M. rufobrunnea FFAAS0414 ON427728 ON468453 ON468466 China This study
57. M. rufobrunnea FFAAS0415 ON427729 ON468454 ON468467 China This study
58. M. rufobrunnea FFAAS0416 Holotype ON427730 ON468455 ON468468 China This study
59. M. seminau ACL136 KF537250 – – Malaysia Chew et al. (2014)
60. M. seminau ACL308 KF537252 – – Malaysia Chew et al. (2014)
61. M. shengshanensis FFAAS0424 Holotype ON427739 ON468464 ON468477 China This study
62. M. shengshanensis FFAAS0425 ON427740 ON468465 ON468478 China This study
63. M. sinar ACL092 KF537247 – – Malaysia Chew et al. (2014)
64. M. sinar ACL135 KF537249 – – Malaysia Chew et al. (2014)
65. M. sinar var. tangkaisinar ACL307 KF537251 – – Malaysia Chew et al. (2014)
66. M. subulata FFAAS0419 ON427735 ON468460 ON468473 China This study
67. M. subulata FFAAS0420 ON427736 ON468461 ON468474 China This study
68. M. subulata FFAAS0423 Holotype ON427737 ON468462 ON468475 China This study
69. M. subulata FFAAS0426 ON427738 ON468463 ON468476 China This study
70. M. supina 128a JF908388 – – Italy Osmundson et al. (2013)
71. M. yuezhuoi FFAAS0344 MW581490 MW868166 MW882249 China Liu et al. (2021)
72. M. yuezhuoi FFAAS0345 MW581491 MW868169 MW882250 China Liu et al. (2021)
73. M. yuezhuoi FFAAS0346 MW581492 MW868168 MW882251 China Liu et al. (2021)
74. M. yuezhuoi FFAAS0347 MW581493 MW868167 MW882252 China Liu et al. (2021)
Figure 1. 

Bayesian Inference analysis of Mycena sect. Calodontes with ITS, rpb1 and tef1 sequence data. Species in Mycena sect. Supinae served as outgroup. Bootstrap values (BS) from Maximum Likelihood ≥ 75 and Bayesian posterior probabilities (BPP) ≥ 0.95 are shown on each branch (BS/BPP). The new species are marked in red.

The phylogenetic analysis revealed that sect. Calodontes was strong support (BS/Bayesian posterior probability [BPP] = 100/1.00) (Fig. 1). Fifteen species and eleven M. pura complex clades within sect. Calodontes were retrieved. Four new species were resolved as monophyletic, each with strong support: M. polycystidiata (BS/BPP = 100/1.00), M. rufobrunnea (BS/BPP = 100/1.00), M. shengshanensis (BS/BPP = 90/1.00) and M. subulata (BS/BPP = 100/1.00). A sister relationship between M. shengshanensis and M. pearsoniana Dennis ex Singer was well supported. Mycena subulata was resolved as sister, but genetically distant from M. pearsoniana and M. shengshanensis clade. In addition, the sister relationship of Mycena polycystidiata and M. rufobrunnea were unresolved.

Taxonomy

Mycena polycystidiata Z.W. Liu, Y.P. Ge, L. Zou & Q. Na, sp. nov.

MycoBank No: MycoBank No: 843977
Figs 2, 3, 4, 5

Diagnosis

Pileus greyish-rose, umbo brownish-orange, hygrophanous. Stipe pubescent. Pleurocystidia polymorphic in shape. Stipitipellis a cutis, with numerous projecting hyphae.

Holotype

China. Heilongjiang Province: Liangshui National Nature Reserve, Yichun City, 47°12'74"N, 128°52'86"E, 20 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0417 (collection number MY0633).

Etymology

Refers to the variable shape of pleurocystidia.

Description

Pileus 14–31 mm in diam., campanulate to hemispherical when young, plano-convex with age, with obtuse umbo at centre, margin slightly revolute, at times cracked at mature; umbo brownish-orange (7C3–7C5), disc purplish-grey (13C2, 14B2, 14C2), reddish-grey (12B2, 12C2) to greyish-rose (12B3), near margin reddish-grey (12D2), greyish-ruby (12D3) or purplish-grey (13D3), margin whitish; striate none or indistinct, greyish-ruby (12E3–12E4), towards the centre up to 1/3 diam.; surface dry and rugose, hygrophanous, generally tomentose. Context white, 2 mm thick, fragile. Lamellae emarginate, slightly decurrent when old, 20–28 reaching the stipe, 1–2 tiers of lamellulae, white, irregularly intervenose, edge concolorous, wavy. Stipe 34–73 × 3–7 mm, central, cylindrical, base occasionally compressed with age; apex violet brown (11E3–11E4, 11F4), greyish-ruby (12E4), lower part brownish-grey (11D2, 11E2) to greyish-brown (11D3, 11E3) or purplish-grey (13C2), fragile, hollow; apex to middle densely pubescent, sparser towards base; whitely villose at base. Odour strongly raphanoid, taste indistinct.

Basidiospores (130/5/4) (6.4)6.7–7.4–8.3(8.8) × (3.2)3.5–3.9–4.3(4.6) μm [Q = (1.62)1.72–2.05(2.18), Q = 1.90 ± 0.11] [holotype (70/2/1) (6.7)6.9–7.6–8.5(8.7) × (3.4)3.6–4.0–4.4(4.6) μm, Q = (1.71)1.76–2.05(2.13), Q = 1.90 ± 0.09], elongated ellipsoid to cylindrical, colourless, smooth, thin-walled, amyloid. Basidia 21–31 × 6–8 μm, 4-spored, clavate, hyaline, sterigmata approximately 4 μm in length. Cheilocystidia thin-walled, hyaline, differs in two shapes, mainly utriform, 50–65 × 20–31 μm, some subclavate, 54–78 × 14–19 μm. Pleurocystidia abundant, thin-walled, hyaline, multi-shaped: lanceolate and mostly round to blunt apices, 37–81 × 12–20 μm, lanceolate and acute apices, 51–87 × 14–22 μm, elliptical, 30–86 × 12–31 μm, ovate and acute apices, 49–71 × 15–24 μm, ovate and mostly round to blunt apices, 49–73 × 16–22 μm. Pileipellis a cutis composed of four to five layers cylindrical cells, 51–81 × 4–5 μm, smooth and thin-walled; terminal cells cylindrical or fusiform, 50–69 × 3–22 μm, thin-walled, hyaline. Hypodermium formed by fusiform to subglobose hyphae, 32–69 × 18–54 μm, thin-walled, hyaline. Lamellar trama subregular, dextrinoid. Stipitipellis a cutis composed of cylindrical hyphae 5–8 μm in diam., smooth, thin-walled, with numbers of projecting hyphae 2–6 μm in diam.; caulocystidia 29–74 × 6–19 μm, clavate or fusiform, thin-walled, smooth, hyaline. Clamps present in all tissues.

Habitat

Scattered on the litter layers in Pinus koraiensis and Larix gmelinii mixed forests.

Known distribution

Heilongjiang Province, China.

Additional material examined

China. Heilongjiang Province: Liangshui National Nature Reserve, Yichun City, 47°12'82"N, 128°52'94"E, 20 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0418 (collection number MY0634); same location, 21 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0421 (collection number MY0659); same location, 21 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0422 (collection number MY0661).

Notes

Macroscopically, Mycena luteovariegata Harder & Læssøe and M. pura resemble M. polycystidiata in pileus colour, but the latter possesses more typically utriform cheilocystidia and uncontracted pleuro- and cheilocystidia (Perry 2002; Robich 2003; Harder et al. 2013; Aronsen and Læssøe 2016; Na 2019). Mycena pearsoniana also has a rose to violaceous pileus, but differs from M. polycystidiata in having inamyloid spores and lacking pleurocystidia (Aronsen and Læssøe 2016; Na 2019). Compared with M. polycystidiata, M. sirayuktha Aravind. & Manim. has similar cheilocystidia, but has an obviously greyish-brown striate pileus, inamyloid spores and slightly glutinous pileipellis with finger-like excrescences (Aravindakshan and Manimohan 2015).

The pleurocystidia of M. polycystidiata varied in shape amongst specimens (Fig. 5). In all four specimens, most pleurocystidia were lanceolate and with round to blunt apices, but pleurocystidia with lanceolate and acute apices, elliptical and ovate and acute apices were also observed in FFAAS0417 (Holotype) and FFAAS0418, while elongated lageniform-lanceolate or round apices ovate were detected in FFAAS0421 and FFAAS0422. The multi-shaped pleurocystidia may show a morphological continuum that changes between developmental stages. Nevertheless, the multi-shaped pleurocystidia are unquestionably diagnostic for identification of this species.

Figure 2. 

Basidiomata of Mycena polycystidiata Z.W. Liu, Y.P. Ge, L. Zou & Q. Na a, b FFAAS0422 c, d FFAAS0417, holotype e–g FFAAS00421 h FFAAS0418 Scale bars: 10 mm (a–h). Photographs a–e, h by Qin Na f, g by Yupeng Ge.

Figure 3. 

Microscopic features of Mycena polycystidiata (FFAAS0417, holotype) a–f basidiospores g basidia h–l cheilocystidia m–r pleurocystidia s pileipellis and hypodermium t lamellar trama u stipitipellis and caulocystidia. Scale bars: 5 μm (a–f); 10 μm (g); 30 μm (h–r); 40 μm (s–u).

Figure 4. 

Morphological features of Mycena polycystidiata (FFAAS0417, holotype) a basidiomata b basidia c basidiospores d pleurocystidia e cheilocystidia f stipitipellis and caulocystidia g pileipellis and hypodermium. Scale bars: 10 mm (a); 10 μm (b–g). Drawings by Zewei Liu.

Figure 5. 

Pleurocystidia of Mycena polycystidiata a FFAAS0422 b FFAAS0417, holotype c FFAAS0418 d FFAAS0421. Scale bars: 25 μm (a–d).

Mycena rufobrunnea Z.W. Liu, Y.P. Ge & Q. Na, sp. nov.

MycoBank No: MycoBank No: 843978
Figs 6, 7, 8

Diagnosis

Pileus dark brown at centre, disc gradually turning paler to reddish-brown to greyish-brown, edge white. Lamellae obviously intervenose. Stipe apex to middle greyish-magenta to dull violet, lower part darker to dark purple or dark magenta. Cheilocystidia utriform, sometimes clavate. Pleurocystidia absent. Caulocystidia clavate or fusiform. Pileipellis with fusiform terminal cells.

Holotype

China. Jilin Province: Dayangcha, Erdaobaihe Town, Antu County, Yanbian Korean Autonomous Prefecture, 42°20'73"N, 127°56'06"E, 16 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0416 (collection number MY0581).

Etymology

Refers to reddish-brown pileus.

Description

Pileus 12–34 mm in diam., hemispherical to convex when young, then plano-convex, sometimes an unclear umbo at centre, margin slightly revolute, acute to subacute, at times cracked at mature; dark brown (8F6–8F8) at centre, disc gradually turning paler to reddish-brown (8D4–8D5, 8E6–8E8) to greyish-brown (8D3) and turning to whitish at margin; striate reddish-brown (8D4–8D5, 8E6–8E8), towards the centre up to 1/3–1/2 diam.; surface humidus when wet. Context white, 1.5 mm thick, fragile. Lamellae adnexed to emarginate, 20–23 reaching the stipe, 1–3 tiers of lamellulae, white, irregularly intervenose, edge concolorous, slightly serrulate. Stipe 19–62 × 2–6 mm, central, cylindrical, apex to middle greyish-magenta (14E4–14E5) to dull violet (16E3–16E4), lower part darker to dark purple (14F4–14F5) or dark magenta (13F3), fragile, hollow, base slightly swollen with whitish villose. Odour raphanoid, taste indistinct.

Basidiospores (80/4/3) (7.1)7.6–8.4–9.2(9.6) × (3.8)4.0–4.5–5.0 μm [Q = (1.73)1.77–1.98(2.05), Q = 1.88 ± 0.07] [holotype (40/2/1) (7.9)8.1–8.6–9.2(9.4) × 4.2–4.6–5.0 μm, Q = (1.73)1.77–1.96(1.98), Q = 1.87 ± 0.06], elongated ellipsoid to cylindrical, colourless, smooth, thin-walled, amyloid. Basidia 24–34 × 7–10 μm, 4-spored, clavate, hyaline, sterigmata 2–3 μm in length. Cheilocystidia thin-walled, hyaline, utriform, sometimes clavate, 23–44 × 7–17 μm, abundant. Pleurocystidia absent. Pileipellis a cutis composed of four to five slightly interwoven layers of cylindrical cells, 44–70 × 4–7 μm, smooth, thin-walled; terminal cells cylindrical or fusiform, 34–65 × 4–17 μm, thin-walled, hyaline. Hypodermium formed by fusiform, subcylindrical to subglobose hyphae, 15–50 × 12–37 μm, thin-walled, hyaline. Lamellar trama subregular, dextrinoid. Stipitipellis a cutis composed of hyphae 3–9 μm in diam., smooth, thin-walled; caulocystidia common in the apex, sparse in the middle and base, 23–76 × 6–14 μm, clavate and fusiform, thin-walled, hyaline, smooth. Clamps present in all tissues.

Habitat

Scattered on the decayed logs of Acer, Larix, Pinus, Populus, Quercus and Ulmus mixed forests.

Known distribution

Jilin Province, China.

Additional material examined

China. Jilin Province: Dayangcha, Erdaobaihe Town, Antu County, Yanbian Korean Autonomous Prefecture, 42°20'72"N, 127°56'08"E, 16 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0414 (collection number MY0579); same location, 16 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0415 (collection number MY0580).

Notes

Species of sect. Calodontes that are macroscopically similar to Mycena rufobrunnea have been recorded in many regions of the world. Most taxa resemble M. rufobrunnea in pileus colour (Smith 1947; Maas Geesteranus 1992a, 1992b; Grgurinovic 2003; Robich 2003; Chew et al. 2014; Aronsen and Læssøe 2016). Mycena dura Maas Geest. & Hauskn., recorded in Europe, also has a dark brown to greyish-brown pileus, but can be distinguished from M. rufobrunnea in having a white stipe and having pleurocystidia (Robich 2003; Aronsen and Læssøe 2016). Mycena kuehneriana A.H. Sm., which is recorded from the United States and Canada, is distinguished from M. rufobrunnea in that its pileus is pale avellaneous with rose and lilac, almost white when faded and the spores are obviously smaller (5–6 × 2–3 μm) (Smith 1947; Maas Geesteranus 1992a, 1992b). Mycena clarkeana Grgur. and M. nullawarrensis Grgur., described from Australia, are similar to M. rufobrunnea in having a reddish-brown pileus, but both species have broader spores and possess pleurocystidia (Grgurinovic 2003). Mycena cahaya A.L.C. Chew & Desjardin, M. seminau A.L.C. Chew & Desjardin and M. sinar A.L.C. Chew & Desjardin, known from Malaysia, resemble M. rufobrunnea owing to the brown pileus, but differ in having adnate to subdecurrent lamellae, a yellowish-grey or brownish-orange stem, mucronate cheilocystidia and lack caulocystidia (Chew et al. 2014). Microscopically, utriform or clavate cheilocystidia and absence of pleurocystidia are key characteristics of M. rufobrunnea. Mycena diosma Krieglst. & Schwöbel has similar cheilocystidia and pleurocystidia are absent or rare, but it has a strongly hygrophanous pileus and a remarkable change in colour (Robich 2003; Aronsen and Læssøe 2016). Mycena pura, M. sirayuktha and M. vinacea Cleland have similar cheilocystidia, but are easily distinguished from M. rufobrunnea by the presence of pleurocystidia (Perry 2002; Grgurinovic 2003; Robich 2003; Aravindakshan and Manimohan 2015; Aronsen and Læssøe 2016; Na 2019).

Figure 6. 

Basidiomata of Mycena rufobrunnea Z.W. Liu, Y.P. Ge & Q. Na a, b FFAAS0414 c, d FFAAS0415 e–h FFAAS0416, holotype. Scale bars: 10 mm (a–h). Photographs a–h by Qin Na.

Figure 7. 

Microscopic features of Mycena rufobrunnea (FFAAS0416, holotype) a–d basidiospores e basidia f–l cheilocystidia m lamellar trama n pileipellis and hypodermium o stipitipellis and caulocystidia. Scale bars: 5 μm (a–d); 20 μm (e–o).

Figure 8. 

Morphological features of Mycena rufobrunnea (FFAAS0416, holotype) a basidiomata b basidia c cheilocystidia d basidiospores e stipitipellis and caulocystidia f pileipellis and hypodermium. Scale bars: 10 mm (a); 10 μm (b–f). Drawings by Zewei Liu.

Mycena shengshanensis Z.W. Liu, Y.P. Ge & Q. Na, sp. nov.

MycoBank No: MycoBank No: 843979
Figs 9, 10, 11, 12

Diagnosis

Pileus brown to violet-brown at centre, disc light brown to dull red. Cheilocystidia clavate with slightly inflated apex, thick-walled. Pleurocystidia absent. Caulocystidia clavate with tapered apices, apex to middle thick-walled. Scattered to gregarious under Larix gmelinii.

Holotype

China. Heilongjiang Province: Shengshan National Nature Reserve, Heihe City, 49°37'45"N, 126°47'39"E, 23 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0424 (collection number MY0686).

Etymology

Refers to the type locality.

Description

Pileus 13–26 mm in diam., when young parabolic to convex, with obtuse umbo at centre, then plano-convex, margin wavy and revolute, at times cracked at mature; centre light brown (7D5–7D6), brown (7E4–7E8), dark brown (8F5–8F6), violet brown (11F4–11F6), disc paler to light brown (7D4–7D5), brown (7E5–7E6), greyish-brown (8D3), reddish-brown (8D4, 8E4), brownish-grey (11C2), dull red (11C3), margin whitish; striate indistinct, brownish-orange (7C3), greyish-brown (7D3), reddish-grey (12D2), greyish-ruby (12E3–12E5), towards the centre up to 1/3–1/2 diam.; surface slightly moist, smooth. Context white, 1–2 mm thick, fragile. Lamellae sinuate to subdecurrent, 19–25 reaching the stipe, 1–3 tiers of lamellulae, white, irregularly intervenose, edge concolorous, wavy and slightly serrulate. Stipe 26–42 × 2–4 mm, central, cylindrical, apex reddish-brown (8E4–8E5), greyish-ruby (12E4–12E5), greyish-brown (11F3), violet brown (11F4–11F5), dark ruby (12F4–12F5), lower part paler to brownish-grey (7C2), brownish-orange (7C3), greyish-brown (11E3), greyish-ruby (12E3), base darker to brown (7E5), reddish-brown (8E4–8E5), greyish-brown (11F3), violet brown (11F4–11F5), greyish-ruby (12E3), dark ruby (12F6–12F8), fragile, hollow, base swollen with white fibrils. Odour raphanoid, taste indistinct.

Basidiospores (60/3/2) (5.9)6.1–6.9–8.1(8.7) × 3.4–4.0–4.4(4.7) μm [Q = 1.62–1.93 (1.97), Q = 1.75 ± 0.09] [holotype (40/2/1) 6.1–7.0–8.2(8.7) × 3.4–4.0–4.4(4.7) μm, Q = (1.62)1.65–1.93(1.97), Q = 1.77 ± 0.08], elongated ellipsoid, colourless, smooth, thin-walled, amyloid. Basidia 22–32 × 6–8 μm, 4-spored, clavate, hyaline, sterigmata 3–4 μm in length. Cheilocystidia moderately thick-walled (0.5–0.6 μm), clavate with slightly inflated apex, 25–63 × 6–12 μm, hyaline. Pleurocystidia absent. Pileipellis a cutis composed of three to four layers of cylindrical cells, 24–57 × 3–5 μm, smooth and thin-walled; terminal cells cylindrical, apically narrow, 28–49 μm in length, apex 1–3 μm and base 2–5 μm in diam., thin-walled, hyaline. Hypodermium formed by fusiform to subglobose hyphae, 19–53 × 13–30 μm, thin-walled, hyaline. Lamellar trama subregular, weakly dextrinoid to dextrinoid. Stipitipellis a cutis composed of hyphae 4–9 μm in diam., smooth, thin-walled; caulocystidia 22–61 × 5–20 μm, clavate with tapered apices, apex to middle thick-walled, smooth, hyaline. Clamps present in all tissues.

Habitat

Scattered to gregarious on the litter layer in Larix gmelinii.

Known distribution

Heilongjiang Province, China.

Additional material examined

China. Heilongjiang Province: Shengshan National Nature Reserve, Heihe City, 23 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0425 (collection number MY0687).

Notes

In sect. Calodontes, M. diosma, M. pearsoniana and M. yuezhuoi also have clavate cheilocystidia with a slightly inflated apex and lack pleurocystidia, similar to M. shengshanensis, but M. diosma differs in pileus characters, M. pearsoniana has inamyloid spores and M. yuezhuoi has a more purple pileus and subcellular lamellar trama (Robich 2003; Aronsen and Læssøe 2016; Na 2019; Liu et al. 2021). Clavate cheilocystidia are also present in M. luteovariegata and M. pura, but these species differ in having pleurocystidia (Perry 2002; Robich 2003; Harder et al. 2013; Aronsen and Læssøe 2016; Na 2019). Macroscopically, several species in sect. Calodontes also have a brown with reddish or violet pileus or stipe (Smith 1947; Maas Geesteranus 1992a, 1992b; Grgurinovic 2003; Robich 2003; Aronsen and Læssøe 2016). Mycena dura, M. kuehneriana and M. nullawarrensis are distinguished by basidiospore size (Smith 1947; Maas Geesteranus 1992a, 1992b; Grgurinovic 2003; Robich 2003; Aronsen and Læssøe 2016). Two species, M. seminau and M. sirayuktha, reported from Southeast Asia, are similar to M. shengshanensis owing to the brown pileus, but they differ in having gelatinised or sometimes mucronate cheilocystidia and caulocystidia have not been observed (Chew et al. 2014; Aravindakshan and Manimohan 2015). Fusiform, obclavate, ovate and clavate cheilocystidia with a subcapitate protuberance were observed occasionally, but clavate cheilocystidia with a slightly inflated apex represented the predominant morphological type in M. shengshanensis (Fig. 12).

Figure 9. 

Basidiomata of Mycena shengshanensis Z.W. Liu, Y.P. Ge & Q. Na a–d FFAAS0424, holotype e–h FFAAS0425 Scale bars: 10 mm (a–h). Photographs a–c, e–g by Yupeng Ge d, h by Zewei Liu.

Figure 10. 

Microscopic features of Mycena shengshanensis (FFAAS0424, holotype) a–e basidiospores f basidia g–r cheilocystidia s lamellar trama t pileipellis and hypodermium u stipitipellis and caulocystidia. Scale bars: 5 μm (a–e); 20 μm (g–r); 40 μm (s–u).

Figure 11. 

Morphological features of Mycena shengshanensis (FFAAS0424, holotype) a basidiomata b basidia c basidiospores d cheilocystidia e stipitipellis and caulocystidia f pileipellis and hypodermium. Scale bars: 10 mm (a); 10 μm (b–f). Drawings by Zewei Liu.

Figure 12. 

Cheilocystidia of Mycena shengshanensis FFAAS0424, holotype. Scale bars: 10 μm (a).

Mycena subulata Z.W. Liu, Y.P. Ge & Q. Na, sp. nov.

MycoBank No: MycoBank No: 843980
Figs 13, 14, 15

Diagnosis

Pileus reddish-grey to dull red, slightly hygrophanous. Cheilocystidia thick-walled, slenderly fusiform with distinctly long and narrow protuberance. Stipitipellis a cutis, with projecting hyphae, caulocystidia thick-walled.

Holotype

China. Heilongjiang Province: Liangshui National Nature Reserve, Yichun City, 47°13'13"N, 128°53'21"E, 21 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0423 (collection number MY0671).

Etymology

Refers to cheilocystidia with distinctly long and narrow protuberance.

Description

Pileus 9–32 mm in diam., convex to campanulate when young, hemispherical to applanate with age, margin sometimes wavy, slightly deflexed; at centre dull red (8C3), brownish-grey (8D2), greyish-brown (8D3), reddish-brown (8D4, 8E4–8E5) and dark brown (8F5), disc paler to reddish-grey (8B2, 9B2), brownish-grey (9C2), dull red (9B3, 9C3), greyish-magenta (13D3), margin light brown (7D4), brown (7E4) or dull red (9C3); striate none or indistinct, reddish-brown (8E4–8E5), towards the centre up to 1/5 diam.; surface dry, unclearly rugose or none, margin slightly hygrophanous. Context white, 1 mm thick, fragile. Lamellae sinuate to subdecurrent, 31–33 reaching the stipe, 1–3 tiers of lamellulae, white, irregularly intervenose, edge concolorous, wavy and slightly serrulate. Stipe 27–75 × 2–5 mm, central, cylindrical, apex to middle brownish-orange (7C3), dull red (8C3), brownish-grey (7D2), greyish-magenta (14D3), lower part brownish-grey (8C2), greyish-brown (8D3), reddish-brown (8D4), fragile, hollow, white granular near apex, base slightly swollen with white fibrils. Odour raphanoid, taste indistinct.

Basidiospores (100/5/4) 6.0–6.7–7.3(7.9) × 3.3–3.8–4.3(4.6) μm [Q = (1.61)1.65–1.87(1.90), Q = 1.76 ± 0.07] [holotype (40/2/1) (6.0)6.2–6.6–7.1 × 3.4–3.7–4.0(4.2) μm, Q = 1.65–1.87(1.90), Q = 1.78 ± 0.07], elongated ellipsoid, colourless, smooth, thin-walled, amyloid. Basidia 23–34 × 5–6 μm, 4-spored, clavate, hyaline, sterigmata 2–3 μm in length. Cheilocystidia moderately thick-walled (0.5–0.6 μm), hyaline, narrowly fusiform with long and narrow protuberance, 43–82 × 4–11 μm, protuberance 14–36 × 1–2 μm. Pleurocystidia absent. Pileipellis a cutis composed of three to four layers of cylindrical cells, 20–89 × 3–7 μm, smooth and thin-walled; terminal cells cylindrical, apically narrow, 24–61 μm in length, the apex 3–4 μm and base 4–7 μm in diam., thin-walled, hyaline. Hypodermium formed by fusiform to subglobose hyphae, 21–41 × 17–25 μm, thin-walled, hyaline. Lamellar trama subregular, dextrinoid. Stipitipellis a cutis composed of hyphae 3–8 μm in diam., smooth, thin-walled, with projecting hyphae 3–8 μm in diam.; caulocystidia 29–47 × 6–12 μm, clavate and apices tapered, thick-walled (0.5–0.6 μm), smooth. Clamps present in all tissues.

Habitat

Scattered on the litter layer in Pinus koraiensis, Larix gmelinii and Tilia sp. mixed forests.

Known distribution

Heilongjiang Province, China.

Additional material examined

China. Heilongjiang Province: Liangshui National Nature Reserve, Yichun City, 21 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0419 (collection number MY0654); same location, 21 August 2021, Zewei Liu, Yupeng Ge, Qin Na and Shixin Wang, FFAAS0420 (collection number MY0657); Heilongjiang Province: Taipinggou National Nature Reserve, Hegang City, 3 September 2021, Shixin Wang, FFAAS0426 (collection number MY0795).

Notes

Cheilocystidia with a long and narrow protuberance is the key microscopic character that distinguishes M. subulata and is uncommon in sect. Calodontes (Smith 1947; Maas Geesteranus 1992a, 1992b; Grgurinovic 2003; Robich 2003; Chew et al. 2014; Aravindakshan and Manimohan 2015; Aronsen and Læssøe 2016; Na 2019; Liu et al. 2021). Mycena lammiensis Harmaja and M. pelianthina (Fr.) Quél. have similar cheilocystidia, but differ from M. subulata by their broader cheilocystidia with purplish-brown contents and having pleurocystidia (Smith 1947; Robich 2003; Aronsen and Læssøe 2016). The cheilocystidia of M. subcorticalis (Cooke & Massee) Sacc. with a protuberance are similar to those of M. subulata. However, M. subcorticalis has larger and inamyloid spores, a gelatinised pileipellis and a stipitipellis with sparse excrescences (Grgurinovic 2003). More rarely, mucronate cheilocystidia and absence of pleurocystidia have been described for M. pearsoniana and its clay pink pileus is similar to that of M. subulata, but M. pearsoniana differs in having a slightly glutinous pileus when wet and inamyloid spores (Aronsen and Læssøe 2016; Na 2019). Other species that are macroscopically similar to M. subulata, namely M. luteovariegata, M. nullawarrensis and M. pura, can be distinguished by cheilocystidia shape and presence of pleurocystidia (Perry 2002; Robich 2003; Grgurinovic 2003; Harder et al. 2013; Aronsen and Læssøe 2016; Na 2019).

Figure 13. 

Basidiomata of Mycena subulata Z.W. Liu, Y.P. Ge & Q. Na a, b FFAAS0419 c, d FFAAS0420 e–g FFAAS0423, holotype h FFAAS0426 Scale bars: 10 mm (a–h). Photographs a, b, e, f by Yupeng Ge c, d by Qin Na g by Zewei Liu h by Shixin Wang.

Figure 14. 

Microscopic features of Mycena subulata (FFAAS0423, holotype) a–e basidiospores f basidia g–m cheilocystidia n lamellar trama o pileipellis and hypodermium p stipitipellis and caulocystidia. Scale bars: 5 μm (a–e); 10 μm (f); 20 μm (g–p).

Figure 15. 

Morphological features of Mycena subulata (FFAAS0423, holotype) a basidiomata b basidia c basidiospores d cheilocystidia e stipitipellis and caulocystidia f pileipellis and hypodermium. Scale bars: 10 mm (a); 5 μm (c); 10 μm (b, d–f). Drawings by Zewei Liu.

Key to species of sect. Calodontes known worldwide

1 Stipe white 2
– Stipe coloured 4
2 Pileus white Mycena subaquosa
– Pileus coloure 3
3 Pileus pink and lamellae emarginate, pileipellis without inflated terminal cells Mycena rosea
– Pileus brown and lamellae adnate, pileipellis with fusiform, subcylindrical to lageniform terminal cells Mycena dura
4 Lamellae edge with coloured dots 5
– Lamellae edge white or without dots 6
5 Caulocystidia present, spores almost broader than 4 μm Mycena lammiensis
– Caulocystidia absent, spores almost narrower than 4 μm Mycena pelianthina
6 Basidiospores inamyloid 7
– Basidiospores amyloid 10
7 Pleurocystidia absent 8
– Pleurocystidia present 9
8 Stipitipellis and caulocystidia smooth Mycena pearsoniana
– Stipitipellis and caulocystidia with nodulose excrescences Mycena subcorticalis
9 Pileipellis gelatinised, caulocystidia absent, cheilo- and pleurocytsidia base uncontracted, disc greyish-red or orange white in pileus Mycena sirayuktha
– Pileipellis not gelatinised, caulocystidia present, cheilo- and pleurocytsidia base contracted, disc wood brown or reddish-brown in pileus Mycena vinacea
10 Pleurocystidia present 11
– Pleurocystidia absent 16
11 Caulocystidia absent, almost cheilocystidia apically mucronate or subcapitate Mycena cahaya
– Caulocystidia present, almost cheilocystidia apically broadly rounded 12
12 Caulocystidia with apical excrescences, spores more than 5.6 μm width Mycena clarkeana
– Caulocystidia without apical excrescences, spores less than 5.6 μm width 13
13 Cheilocystidia base uncontracted Mycena polycystidiata
– Cheilocystidia base contracted 14
14 Stipe brown to dark brown, Qav = 1.5 Mycena nullawarrensis
– Stipe not brown to dark brown, Qav > 1.5 15
15 Pileus sulphur yellow to reddish-grey, stipe reddish-grey Mycena luteovariegata
– Pileus generally pinkish or purplish, stipe whitish to pinkish-purple Mycena pura
16 Caulocystidia absent 17
– Caulocystidia present 18
17 Pileus brown to dark brown, spores weakly amyloid Mycena seminau
– Pileus brownish-orange to greyish-yellow, spores amyloid Mycena sinar
18 Cheilocystidia slender fusiform, with distinctly long and narrow protuberance Mycena subulata
– Cheilocystidia clavate, utriform, subfusiform, or subcylindrical, with short mucronate or none 19
19 Spores less than 6 μm length Mycena kuehneriana
– Spores more than 6 μm length 20
20 Pileus more than 35 mm in diam., lamellae dark brownish-violet to reddish-violet Mycena diosma
– Pileus less than 35 mm in diam., lamellae white 21
21 Lamellar trama subcellular, pileus lilac to purple Mycena yuezhuoi
– Lamellar trama subregular, pileus brownish 22
22 Cheilocystidia utriform, sometimes clavate, thin-walled, lamellae adnexed to emarginate Mycena rufobrunnea
– Cheilocystidia clavate with slightly inflated apex, thick-walled, lamellae sinuate to subdecurrent Mycena shengshanensis

Discussion

Maas Geesteranus (1980) proposed that Mycena sect. Calodontes could be divided into three subsections based on the colour of the lamellar edge and the amyloid reaction of the basidiospores. Subsequently, taxonomists have followed this division, but opinions have differed on the diagnostic characters that support this classification (Grgurinovic 2003; Robich 2003; Harder et al. 2010; Chew et al. 2014). Some taxonomists classified the subsections according to the amyloid reaction of basidiospores, cheilocystidia and pleurocystidia contents and presence or absence of pleurocystidia, but the shapes of cheilocystidia and pleurocystidia were not considered (Grgurinovic 2003; Harder et al. 2010; Chew et al. 2014). Robich (2003) also did not consider the shapes of cheilocystidia and pleurocystidia, but the colour of the lamellar edge and cheilocystidia contents were emphasised to distinguish subsections. According to the historical infrasectional classification of sect. Calodontes, M. polycystidiata could be classified in subsect. Purae, whereas M. rufobrunnea, M. shengshanensis and M. subulata cannot be assigned to a subsection owing to their having amyloid spores and lacking pleurocystidia (Maas Geesteranus 1980; Harder et al. 2010).

Phylogenetic reconstructions do not fully support recognition of three subsections defined by morphological characters; notably, subsect. Violacellae and subsect. Purae are polyphyletic in the phylogenies (Harder et al. 2010, 2012, 2013). Chew et al. (2014) supported the views of Harder et al. (2010, 2012) and the new taxa proposed by the former authors were not assigned to a subsection. Additionally, subsect. Purae was proved to be polyphyletic in our combined analysis of ITS, rpb1 and tef1 dataset, which also supported analysis, based on single gene region (Harder et al. 2013).

The five taxa of Mycena sect. Calodontes recorded from China show obvious differences in pileus colour and in the shapes of cheilocystidia and pleurocystidia (if present) (Liu et al. 2021). The colour of the pileus includes greyish-rose, reddish-grey, purple, reddish-brown and violet-brown and most show a gradual transition with age. Clavate, obclavate, utriform and fusiform cheilocystidia with a long, narrow protuberance are observed, but pleurocystidia are present only in M. polycystidiata. Forms and variations within M. pura complex had a wide range of pileus colour, but the shape of cheilocystidia was highly similar and could be clearly distinguished from the four new taxa (Robich 2003).

In our phylogenetic analysis, four new species all formed separate clades with high support and had obvious genetic distance from other species in sect. Calodontes. Mycena rufobrunnea is more closely related to the phylogenetic species within M. pura complex by Harder et al. (2013). While the other three new species are significantly more distant from M. pura complex genetically, M. shengshanensis and M. subulata, formed a sister relationship with high support from M. pearsoniana; M. polycystidiata clustered with M. diosma, but is poorly supported.

Based on extensive field work in China, most specimens of sect. Calodontes have been observed in coniferous forests or mixed coniferous-broadleaved forests in early autumn (Na 2019; Liu et al. 2021). Specimens of the four new taxa described in the present study were collected from Changbai Mountain and the Lesser Khinggan Mountains in northeast China from mixed broadleaf-Korean pine (Pinus koraiensis) forests (Zhao et al. 2004; Wang and Guo 2016). In particular, M. polycystidiata and M. subulata were both distributed in the Liangshui National Nature Reserve on the Lesser Khinggan Mountains, where the dominant forest species is P. koraiensis, mixed with fewer Betula, Tilia, Quercus and Picea individuals (She et al. 2022). Moreover, more specimens were located in the northern region of China with an average temperature not more than 20 °C in August. For example, the average temperature is 16.4 °C in Liangshui National Nature Reserve and 16.3 °C in Shengshan National Nature Reserve (Liu 2017). Therefore, we speculate that members of this section in China prefer the climate types Dwa, Dwb and Dwc according to the Köppen climate classification (Kottek et al. 2006; Wang et al. 2020).

Acknowledgements

This study was supported by the Natural Science Foundation of Shandong Province (grant No. ZR2020QC001), the National Natural Science Foundation of China (grant No. 3190012), the Natural Science Foundation of Shandong Province (grant No. ZR2019PC028) and the Shandong Agricultural Industry Technology System (grant No. SDAIT-07-03). We sincerely thank Prof. Li Zou and Mr. Shixin Wang (Northeast Forestry University, Harbin, Heilongjiang Province), Mr. Jingyun Song and Mr. Yong Li (Yanhexiang Township Government, Sunwu County, Heilongjiang Province), Ms Yaping Hu (Nanjing Institute of Environmental Sciences, Ministry of Ecology and Environment, Nanjing, Jiangsu Province) and Mr. Bai Wang (Changbai Mountain National Nature Reserve, Jilin Province) for their help during fieldwork.

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