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Research Article
Additions to the genus Mycena (Mycenaceae, Agaricales): Descriptions of five new taxa in Hunan Province, China
expand article infoYing Xin Xiao‡, Li Na Liu§, Zhu Ming Tan§, Ai Rong Shen§, Bao Ming Shen§, Yun Tan§, Sai Nan Li§, Li Guo Feng|, Jing Bo Long, Zhu Xiang Liu‡
‡ Jishou University, Jishou, China
§ Institute of Biodiversity Studies, Hunan Academy of Forestry, Changsha, China
| Hunan Edible Fungi Research Institute, Changsha, China
¶ Huangsang National Nature Reserve Management Office, Shaoyang, China

Corresponding author: Zhu Xiang Liu ( liuzhuxiang@126.com )

Academic editor: Samantha C. Karunarathna
© 2025 Ying Xin Xiao, Li Na Liu, Zhu Ming Tan, Ai Rong Shen, Bao Ming Shen, Yun Tan, Sai Nan Li, Li Guo Feng, Jing Bo Long, Zhu Xiang Liu.
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: Xiao YX, Liu LN, Tan ZM, Shen AR, Shen BM, Tan Y, Li SN, Feng LG, Long JB, Liu ZX (2025) Additions to the genus Mycena (Mycenaceae, Agaricales): Descriptions of five new taxa in Hunan Province, China. MycoKeys 115: 327-362. https://doi.org/10.3897/mycokeys.115.144137
Open Access

Abstract

Few studies have been conducted on Mycena species in Hunan Province, China. During our research on the species diversity of Mycena in Hunan Province, we identified approximately 30 Mycena species based on morphological and molecular evidence. Five species are recognized herein as new to science, namely, M. fulvomarginata, M. huangsangensis, M. hongfengensis, M. subroriduliformis, and M. roseolamellata. The phylogenetic analyses of a combined ITS and LSU sequence dataset revealed that five new species each formed an independent lineage that could separate phenotypically similar and phylogenetically related species. Descriptions, photographs, and phylogenetic analysis results are provided for the five new species, along with the comparisons with related species. A key to all Mycena species found in Hunan is also provided.

Key words:

Basidiomycota, biodiversity, five new species, phylogenetic analysis, taxonomy

Introduction

Hunan Province is located in the southern middle reaches of the Yangtze River, China, and covers an area of approximately 211,800 km2 (Gao and Dou 1981). The province is surrounded by mountains and hills in the east, west, and south, basins in the center, and plains in the north. This area has a subtropical humid monsoon climate with four distinct seasons, sufficient sunshine, and abundant rainfall, which benefits vegetation (Qing 1990). The people’s government of Hunan Province reported that there are 17 national nature reserves, including 16 focused on preserving forest ecological systems (https://www.hunan.gov.cn/). The unique topography and good forest ecological systems make Hunan home to macrofungi (Liu et al. 2024), among which the members of Mycena (Pers.) Roussel are prominent in Agaricales (Liu et al. 2022a, 2022b).

Mycena is one of the largest genera in the Mycenaceae family of Agaricales, including at least 600 species worldwide (Kirk et al. 2008; Fan et al. 2024; Zhang et al. 2024). Mycena plays an important role in ecosystems, which depend on the strong decomposition abilities of fungi to promote material circulation, forest metabolism, and natural renewal (Fukasawa et al. 2009; Guerreiro et al. 2023). Some Mycena are symbiotic fungi with Gastrodia elata seeds (Lee et al. 2017; Liu et al. 2022b, 2025). Moreover, approximately 60 Mycena species are reported to be bioluminescent worldwide (Chew et al. 2014, 2015; Cortés-Pérez et al. 2019, 2023; Lu et al. 2024).

Mycena was first moved to the genus level by Roussel in 1806, but this change was not widely accepted by the majority of mycologists until the 20th century (Roussel 1806). It is a widely studied genus despite many species in the genus having small basidiomata (Aronsen and Læssøe 2016). Many important contributions about taxonomic studies of Mycena have been made to temperate regions, and several monographs have been published (Smith 1947; Robich 2003, 2016; Aronsen and Læssøe 2016). For subtropical and tropical areas, the study of local floras has contributed to the early reporting of Mycena species (Métrod 1949; Singer and Digilio 1951; Pegler 1977, 1983, 1986, 1987), and those studies made substantial contributions to more important papers and related monographs on Mycena (Singer 1989; Corner 1994; Grgurinovic 1998, 2003; Perry 2002; Cooper et al. 2018; Bau et al. 2021).

Some classifications have been proposed based on the morphological characteristics of Mycena (Fries 1821; Lange 1914; Kühner 1926, 1931, 1938; Singer and Digilio 1951; Singer 1986; Maas Geesteranus 1992a, 1992b; Maas Geesteranus and Horak 1995; Bau et al. 2021). Currently, mycologists tend to accept, use, and update the infrageneric classification, which was proposed by Maas Geesteranus (Grgurinovic 1998, 2003; Robich 2003, 2016; Cooper et al. 2018). Species are classified into 44 sections based on a combination of macroscopic and microscopic features. With the development of molecular biology research, the phylogenetic positions of Mycena have become increasingly apparent (Moncalvo et al. 2002; Matheny et al. 2006; Wei et al. 2024). As a part of the subtropical area of China, studies on Mycena have been conducted in Hunan Province. The book Hunan Macrofungi was specially written in 1997 to document the fungi in the region and described two Mycena species (Li et al. 1993). Five Mycena species from Hupingshan, Hunan, are described in the Atlas of Macrofungi (Zhang et al. 2005). Seven Mycena species were recorded in the Atlas of Macrofungi in Hunan (Chen and Zhang 2019). A species in China, M. heteracantha (Singer) Desjardin, which was collected from Hunan Province, was described by Na Q and Bau T in 2019 (Na and Bau 2019a). Mycena subpiligera L.N. Liu, a new species collected in Hunan Province that can significantly enhance the germination efficiency of Gastrodia elata seeds, was reported in 2022 (Liu et al. 2022b). Mycena chlorocyanea L.N. Liu, another new species whose type was collected in Hunan Province, was reported in recent years (Liu et al. 2022a). As a region that is rich in natural resources, further studies on Mycena in Hunan Province are needed.

Aiming to explore the diversity of Mycena in the region, we conducted literature reviews and field investigations in Hunan Province from 2020 to 2024. Through macroscopic comparison and phylogenetic analysis, we found 30 Mycena species collected from Hunan Province, including five new species that are described in this paper.

Materials and methods

Sample collection and morphological description

Approximately 400 Mycena specimens belonging to 30 taxa were collected from Hunan Province (Table 1). During fieldwork, the collected samples were photographed, and additional information, such as elevation, habitat, and collection date, was recorded (Rathnayaka et al. 2024). Detailed morphological features, including basidiomata size, color and shape, odor, taste, and viscosity, were also documented from fresh specimens. The specimens were dried with silica gel and deposited in the Herbarium of Hunan Institute of Forestry (HUIF) and the Herbarium of Jishou University (JSU). The macromorphological characteristics of the samples were determined based on field notes and photographs. The color codes followed Kornerup & Wanscher (Kornerup and Wanscher 1978). Micromorphological characters of dry specimens were observed under light microscopy (Olympus BX51). To measure the sizes of related tissues, basidiospores, basidia, pileipellis, stipitipellis, and other tissues were mounted in pure water, 5% KOH solution, or 1% Congo Red solution. Melzer’s reagent was used to test the amyloid reaction of the spores. At least 30 basidiospores were measured in each sample. The Q value refers to the length/width ratio of the basidiospores. Q ± av represents the average Q of all basidiospores ± the sample standard deviation. The ranges of the basidiospores are presented as (a–) b–c (–d): the range ‘b–c’ represents 90% of the measured values, while ‘a’ and ‘d’ represent extreme values. The measurements of other microscopic structures were based on 20 measurements per specimen.

Table 1.
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Summary of the collection of Mycena species and their distributions and seasons in Hunan.

Species name Ecology and location (in Hunan) Occurrence time (in Hunan) Geographical compositions References
М. abramsii Solitary, in small groups or subfasciculate on dead twigs or woody debris of deciduous trees; also occurs occasionally on dead twigs of coniferous trees. Widespread in Hunan. May to July Worldwide distribution (Singer and Digilio 1951; Hintikka 1963; Watling 1984; Maas Geesteranus 1988; Emmett 1992; Marra 2000; Tholl et al. 2000; Veerkamp 2001; Laganà et al. 2002; Perry 2002; Robich 2003; Afyon et al. 2005; Bronckers and Kelderman 2011; Senn-Irlet et al. 2012; Gierczyk et al. 2015; Aronsen and Læssøe 2016; Rudolf et al. 2016; Vishwakarma et al. 2017; Bau et al. 2021)
М. adnexa In small groups on dead twigs of deciduous trees. Widespread in Hunan. May to June Worldwide distribution (Bau et al. 2021)
М. algeriensis Subfasciculate on rotten stumps in deciduous forests. Collected only in western areas of Hunan. April to May Temperate-subtropical and tropical distribution (Hintikka 1963; Laganà et al. 2002; Gierczyk et al. 2015; Aronsen and Læssøe 2016; Bau et al. 2021)
М. bicystidiata Scattered on rotten wood in mixed forests. Collected in western and eastern areas of Hunan. April to June Only found in China (Bau et al. 2021)
М. castaneicola Scattered or gregarious on Castanea mollissima fruits. Collected only in western areas of Hunan. June to July Only found in China (Bau et al. 2021)
М. chlorocyanea Gregarious in leaf humus under deciduous trees. Collected only in western areas of Hunan. April to May Only found in China (Liu et al. 2022a)
М. citrinomarginata Gregarious in leaf humus under deciduous trees. Collected in western and eastern areas of Hunan. May to June Temperate-subtropical and tropical distribution (Smith 1935; Hintikka 1963; Watling 1984; Senn-Irlet 1987; Emmett 1992; Laganà et al. 2002; Perry 2002; Kalamees and Raitviir 2006; Senn-Irlet et al. 2012; Gierczyk et al. 2015; Gáperová et al. 2015; Na and Bau 2018; Lee et al. 2021)
М. corynephora Gregarious on bark of living deciduous trees. Widely distributed in Hunan. April to May Worldwide distribution (Desjardin 1995; Laganà et al. 2002; Aronsen and Læssøe 2016; Bau et al. 2021)
М. deeptha Gregarious on rotten wood of living deciduous trees. Collected in western and southern areas of Hunan. April to July Temperate-subtropical and tropical distribution (Aravindakshan et al. 2012)
М. digitifurcata Gregarious on bark of living deciduous trees. Collected only in Changsha. June Only found in China (Wei et al. 2024)
М. filopes Solitary on dead twigs of deciduous trees. Collected only in western areas of Hunan. October Worldwide distribution (Beardslee and Coker 1924; Watling 1984; Kalamees and Raitviir 2006; Bronckers and Kelderman 2011; Senn-Irlet et al. 2012; Aronsen and Læssøe 2016; Bau et al. 2021; Cho and Chung 2023)
М. fulvomarginata Gregarious on moss-covered bark of living deciduous trees. Collected only in western areas of Hunan. June to July Only found in China This study
М. galericulata Solitary to fasciculate on branches, trunks and stumps of deciduous trees. Collected only in western areas of Hunan. October to
November		 Worldwide distribution (Beardslee and Coker 1924; Swartz 1933; Imai 1938; Hintikka 1963; Ballesteros 1984; Watling 1984; Maas Geesteranus 1992c; Nauta and Vellinga 1992; Marra 2000; Tholl et al. 2000; Laganà et al. 2002; Perry 2002; Robich 2003; Senn-Irlet et al. 2012; Gáperová et al. 2015; Aronsen and Læssøe 2016; Gyosheva et al. 2016; Vishwakarma et al. 2017; Cho and Chung 2020, 2023; Bau et al. 2021; Lee et al. 2021; Łuszczyński et al. 2022)
М. haematopus In small groups or fasciculate on twigs and trunks of deciduous trees. Distributed in eastern, western and southern areas of Hunan. May, June, October,
November		 Worldwide distribution (Beardslee and Coker 1924; Swartz 1933; Imai 1938; Hintikka 1963; Watling 1984; Emmett 1992; Grgurinovic 1998; Laganà et al. 2002; Perry 2002; Robich 2003; Bronckers and Kelderman 2011; Senn-Irlet et al. 2012; Aravindakshan and Manimohan 2013b; Aronsen and Læssøe 2016; Mustafabayli and Aghayeva 2019; Bau et al. 2021; Cho and Chung 2023)
М. heteracantha Gregarious on decaying leaves and twigs of deciduous trees. Collected only in southwestern areas of Hunan. May Temperate-subtropical and tropical distribution (Desjardin 1995; Bau et al. 2021)
М. hongfengensis Gregarious on decaying leaves of deciduous trees. Collected only in western areas of Hunan. April Only found in China This study
М. huangsangensis Gregarious on decaying leaves of deciduous trees. Collected only in southwestern areas of Hunan. April to May Only found in China This study
М. laevigata In small groups or fasciculate on twigs and trunks of deciduous trees. Collected in western and southern areas of Hunan. June to September Temperate-subtropical and tropical distribution (Chen and Zhang 2019)
М. leaiana Fasciculate on rotten wood of deciduous trees. Collected in western and southern areas of Hunan. July Temperate-subtropical and tropical distribution (Chen and Zhang 2019)
М. leptocephala Gregarious on moss-covered hardwood or on branches. Collected in western and southern areas of Hunan. April to May Worldwide distribution (Beardslee and Coker 1924; Smith 1935; Hintikka 1963; Watling 1984; Senn-Irlet 1987; Laganà et al. 2002; Kalamees and Raitviir 2006; Bronckers and Kelderman 2011; Senn-Irlet et al. 2012; Gierczyk et al. 2015; Aronsen and Læssøe 2016; Baldrian et al. 2016; Bau et al. 2021; Łuszczyński et al. 2022)
М. maculata Solitary to fasciculate on branches, trunks and stumps of deciduous trees. Collected in western areas of Hunan. October to
November		 Temperate distribution (Marra 2000; Tholl et al. 2000; Laganà et al. 2002; Perry 2002; Bau et al. 2021)
М. meliigena/juniperina Gregarious on moss-covered bark of living deciduous trees. Widespread. November to June Temperate distribution (Aronsen 1996; Doğan and Karadelev 2006; Halama et al. 2014)
М. pearsoniana Scattered in leaf humus in deciduous trees. Collected only in western areas of Hunan. May to June Worldwide distribution (Hintikka 1963; Watling 1984; Moreno and Albertó 1996; Robich 2003; Senn-Irlet et al. 2012; Aronsen and Læssøe 2016; Türkekul 2017; Vishwakarma et al. 2017; Kwon et al. 2020; Bau et al. 2021)
М. picta Scattered on decaying leaves of deciduous trees. Collected only in some parks of Changsha. April to May Temperate-subtropical and tropical distribution (Miyamoto et al. 1996; Halama and Romański 2010; Shiryaeva 2018; Retnowati et al. 2020; Bau et al. 2021)
М. pluteoides Solitary or gregarious on rotten wood of deciduous trees. Collected in western and southern areas of Hunan. May, June Only found in China (Bau et al. 2021)
М. pura Scattered in leaf humus and on needles or in grasslands, on both deciduous and coniferous trees. Widely distributed in Hunan. November, March
to June		 Worldwide distribution (Beardslee and Coker 1924; Swartz 1933; Imai 1938; Hintikka 1963; Watling 1984; Senn-Irlet 1987; Emmett 1992; Maas Geesteranus 1992c; Marra 2000; Laganà et al. 2002; Perry 2002; Robich 2003; Kalamees and Raitviir 2006; Senn-Irlet et al. 2012; Casabón 2015; Gáperová et al. 2015; Aronsen and Læssøe 2016; Gyosheva et al. 2016; Mustafabayli and Aghayeva 2019; Bau et al. 2021; Lee et al. 2021)
М. roseolamellata Gregarious on decayed twigs of bamboo or woody debris of deciduous trees. Ningxiang, Hunan. November,
December and
March		 Only found in China This study
М. subpiligera Longshan and Suining Counties, Hunan. April to July Only found in China (Liu et al. 2022b)
М. subroriduliformis Gregarious on decaying leaves of deciduous trees. Suining County, Hunan. April to May Only found in China This study
М. yuezhuoi Scattered on the litter layer in Pinus, Quercus, and Robinia mixed forests. Suining County, Hunan. April to May Temperate-subtropical and tropical distribution (Liu et al. 2021; Cho et al. 2024)

DNA extraction, PCR amplification, and sequencing

Genomic DNA was extracted from fresh or dried specimens using the NuClean Plant Genomic DNA kit (Kangwei Century Biotechnology Co., Beijing, China) following the manufacturer’s protocols. The ITS rDNA region (ITS1–5.8S–ITS2) was amplified using the primer pair ITS1 and ITS4 (White et al. 1990). The LSU region was amplified with the primers LR0R and LR7 (Vilgalys and Hester 1990). PCR was performed in a total volume of 25 μL containing 1 μL template DNA, 9.5 μL distilled water, 1 μL of each primer, and 12.5 μL 2x Taq PCR Master Mix with blue dye (Sangon Biotech, Shanghai, China). The PCR conditions were as follows: initial denaturation at 95 °C for 5 min, followed by 35 cycles of 94 °C for 45 s, 45 s at 52 °C, and 1 min at 72 °C for ITS (Liu et al. 2022a). For the LSU conditions: initial denaturation at 95 °C for 4 min, followed by 35 cycles of 95 °C for 1 min, 1 min at 53 °C, and 80 s at 72 °C (Zhang et al. 2024). The amplified products were determined by electrophoresis on a 1% agarose gel against a known standard DNA marker and directly sequenced at Sangon Biotech. Newly generated sequences in this study have been submitted to the NCBI GenBank database.

Molecular phylogeny

Details of the sequences used for phylogenetic analysis were obtained from this study and downloaded from GenBank (Table 2). DNA sequences were checked using Bioedit v7.0.9 to ensure sequencing quality (Hall 1999). SeqMan 7.1.0 was used for splicing and manual editing (Swindell and Plasterer 1997). The final datasets were aligned using MAFFT v.7.310 (Katoh and Standley 2013). The sequences were concatenated into one multi-loci dataset with SequenceMatrix 1.7.8 (Vaidya et al. 2011). The ALTER (Alignment Transformation EnviRonment) online tool was used for the final conversion of the FASTA format to the NEXUS format (Glez-Penñ et al. 2010). The best-fit evolutionary model was selected using MrModelTest v.2.3 under the Akaike information criterion (AIC) (Nylander 2004). A phylogenetic tree was constructed based on maximum likelihood and Bayesian inference methods. Maximum likelihood (ML) analyses were performed with RAxML-NG v.0.9.0 (Kozlov et al. 2019), and bootstrap values were calculated from 1,000 replicates. Bayesian inference analysis was performed using the Metropolis-coupled Markov chain Monte Carlo method with MrBayes v3.2.5 under the GTR +I+G model (Ronquist and Huelsenbeck 2003). Analyses were run with 4 chains of 2,000,000 generations, and trees were sampled every 100th generation. The first 25% of the sample trees were discarded as burn-in. Gaps were treated as missing data. Phylogenetic trees were visualized with FigTree v1.4.3 (http://tree.bio.ed.ac.uk/software/figtree/).

Table 2.
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Names, voucher numbers, locations, and corresponding GenBank accession numbers of the taxa used in the phylogenetic analysis. - refers to the data unavailability.

Species Voucher GenBank accession no. Location References
ITS LSU
Atheniella adonis H6036863 MW540691 - Finland Unpublished
A. aurantiidisca UBCF33062 MF908459 - Canada Unpublished
Clitocybula intervenosa BAP 588 MH414560 - Africa (Cooper et al. 2018)
C. intervenosa BAP 613 MH414561 MH385335 Africa (Cooper et al. 2018)
Hydropus murinus BAP 657 MH414565 - Africa (Cooper et al. 2018)
Mycena abramsii HUIFS50116 OP604436 OP605596 China Unpublished
M. abramsii HUIFS50074 OP604427 - China Unpublished
M. abramsii HUIF50533 PQ406957 - China This study
M. adnexa HMAJU43360 MK733290 MK722345 China Unpublished
M. adnexa HMAJU43691 MK733293 MK722346 China Unpublished
M. adnexa HUIF50339 PQ406958 - China This study
M. adnexa HUIF60005 PQ465300 - China This study
M. albiceps F27622 MZ303026 - USA Unpublished
M. albiceps RA705-6 MK234177 - USA Unpublished
M. algeriensis HMAS 291753 OR236986 - China Unpublished
M. algeriensis HUIF50368 PQ406959 - China This study
M. alniphila 904 JF908482 - Italy Unpublished
M. amicta CBS:254.53 MH857183 - France (Vu et al. 2019)
M. amicta CBS:352.50 MH856655 MH868170 France (Vu et al. 2019)
M. arcangeliana 252b JF908401 - Spain (Osmundson et al. 2013)
M. arcangeliana 252f JF908402 - Spain (Osmundson et al. 2013)
M. bicystidiata HMJAU43648 MK309773 MK629359 China (Na and Bau 2019b)
M. bicystidiata HUIF50044 PQ406952 - China This study
M. bicystidiata HUIF50583 PQ406953 - China This study
M. breviseta BAP 633 MH414551 MH385327 Africa (Cooper et al. 2018)
M. brunnescens JSU125 ON778578 OP360941 China (Zhang et al. 2024)
M. brunnescens JSU126 ON778579 OP360942 China (Zhang et al. 2024)
M. brunnescens JSU127 PP152232 - China (Zhang et al. 2024)
M. bulliformis SFSU:BAP 547 KX513844 KX513848 USA (Perry and Desjardin 2016)
M. caeruleomarginata FFAAS0358 OL711670 OL711665 China (Na et al. 2022)
M. caeruleomarginata FFAAS0357 OL711669 OL711664 China (Na et al. 2022)
M. capillaripes HRL2854 PQ811198 - USA Unpublished
M. castaneicola JSU138 PQ406949 - China This study
M. castaneicola JSU263 PQ406950 - China This study
M. castaneicola HMJAU43581 MH136827 - China (Na and Bau 2019a)
M. cf. cinerella 173 MF926553 - Russia (Malysheva et al. 2017)
M. chlorocyanea HUIF50234 OP358280 OP360937 China (Liu et al. 2022a)
M. chlorocyanea HUIF50238 OP358281 OP360938 China (Liu et al. 2022a)
M. chlorophos CT15101401 MH400938 - China Unpublished
M. cinerella Aronsen051014 KT900146 - Norway Unpublished
M. citrinomarginata SHXG OM228755 OM228763 China Unpublished
M. citrinomarginata HMJAU 43563 MG654739 - China (Na and Bau 2018)
M. confinationis MO362993 PP831662 - USA Unpublished
M. confinationis PAMP-fungi-41 MT764847 MT764850 Spain Unpublished
M. corynephora JSU145 PQ406951 - China This study
M. corynephora SJiao OP604434 - China Unpublished
M. cristinae JS347 MT921381 MT921384 Brazil (Oliveira et al. 2021)
M. cristinae JS767 MT921382 - Brazil (Oliveira et al. 2021)
M. crocea S.D. Russell iNaturalist #16588497 OM473679 - USA Unpublished
M. crocea OMDL K. Canan iNaturalist 182892200 PP436589 - USA Unpublished
M. cyanorhiza J24082010 MW540696 - Finland Unpublished
M. cyanorhiza 120b JF908385 - Italy (Osmundson et al. 2013)
M. deeptha DM334g (K(M)178333) JX481737 - India (Aravindakshan et al. 2012)
M. deeptha HUIF50518 PQ406962 - China This study
M. digitifurcata HUIF60006 PQ406940 - China (Wei et al. 2024)
M. digitifurcata FFAAS1055 PP706100 PP704700 China (Wei et al. 2024)
M. entolomoides HMJAU 43126 MG654738 - China (Na and Bau 2018)
M. entolomoides HMJAU 43052 MG654737 - China (Na and Bau 2018)
M. entolomoides HMJAU 43048 MG654736 - China (Na and Bau 2018)
M. filopes HUIF50198 OP604441 OP605599 China Unpublished
M. filopes HMAS 291835 OR236988 - China Unpublished
M. flosoides HUIF50128 OP358282 OP360939 China (Liu et al. 2022a)
M. flosoides HUIF50129 OP358283 OP360940 China (Liu et al. 2022a)
M. flosoides HUIF50128-R OP745013 - China (Liu et al. 2022a)
M. fulgoris ACP1690 MG926694 - Mexico (Cortés-Pérez et al. 2019)
M. fulgoris ACP1785 MG926693 - Mexico (Cortés-Pérez et al. 2019)
M. fulvomarginata HUIF50088 Holotype PQ406943 - China This study
M. fulvomarginata HUIF50089 PQ406944 PQ406964 China This study
M. galericulata TFB14675 MN088380 - USA (Hughes et al. 2020)
M. galericulata TFB14649 MN088382 - USA (Hughes et al. 2020)
M. galericulata HUIF50196 OP604439 - China Unpublished
M. haematopus HUIF50203 OP604443 OP605601 China Unpublished
M. haematopus HMJAU43662 MK733299 MK722353 China Unpublished
M. huangsangensis HUIF50526 Holotype PQ406935 - China This study
M. huangsangensis HUIF50528 PQ406936 PQ406965 China This study
M. interrupta HMJAU43849 MK733301 - China Unpublished
M. interrupta HMJAU43791 MK733300 - China Unpublished
M. juniperina 869 JF908478 - Italy (Osmundson et al. 2013)
M. laevigata HMJAU43618 MK733304 MK722355 China Unpublished
M. laevigata MHHNU 8626 MK453048 - China Unpublished
M. leaiana MHHNU 30544 MK250916 - China Unpublished
M. leaiana HKAS126400 OQ025147 China Unpublished
M. leptocephala HUIF50005 PQ406956 - China This study
M. leptocephala CA FUNDIS iNaturalist #160824125 OR778420 - USA Unpublished
M. longinqua BAP 648 MH414552 MH385328 Africa (Cooper et al. 2018)
M. maculata HUIFS50209 OP604446 - China Unpublished
M. maculata HMJAU43009 MK309791 MK629347 China Unpublished
M. meliigena 39c JF908428 - Italy (Osmundson et al. 2013)
M. meliigena 39 JF908423 - Italy (Osmundson et al. 2013)
M. meliigena/ juniperina HUIF60003 PQ406954 - China This study
M. meliigena/ juniperina HUIF60004 PQ406955 - China This study
M. metata 313b JF908412 - Italy (Osmundson et al. 2013)
M. metata HMJAU43625 MH396636 - China Unpublished
M. hongfengensis JSU114 Holotype PQ406945 PQ406967 China This study
M. hongfengensis JSU121 PQ406946 PQ406968 China This study
M. oryzifluens FFAAS1051 PP706096 PP704696 China (Wei et al. 2024)
M. pasvikensis AAronsen50-13 KU861558 - Norway Unpublished
M. pasvikensis AAronsen86-12 KU861556 - Norway Unpublished
M. pearsoniana TENN61384 JN182200 - USA (Harder et al. 2012)
M. pearsoniana TENN61544 JN182199 - USA (Harder et al. 2012)
M. pearsoniana HUIF50392 PQ406948 - China This study
M. picta CA FUNDIS iNaturalist 171114596 OR858681 - USA Unpublished
M. picta TUR194167 MW540717 - Finland Unpublished
M. pluteoides HMJAU43771 MK733307 MK722357 China Unpublished
M. pluteoides HMJAU43765 MK733306 - China Unpublished
M. pluteoides HUIF50584 PQ406961 - China This study
M. pluteoides HUIF50591 PQ406960 - China This study
M. polygramma CBS:240.47 MH856235 MH867764 France (Vu et al. 2019)
M. polygramma 439b JF908433 - Italy (Osmundson et al. 2013)
M. pura HUIF50006 OP604419 OP605597 China Unpublished
M. pura TENN60139 EU517505 - Russia (Petersen et al. 2008)
M. purpureofusca HMJAU 43554 MG654740 - China (Na and Bau 2018)
M. purpureofusca HMJAU 43624 MG654741 - China (Na and Bau 2018)
M. rosella 73h JF908471 - Italy (Osmundson et al. 2013)
M. rosella 53 MW576937 - Norway Unpublished
M. roseolamellata HUIF60001 Holotype PQ406941 PQ406969 China This study
M. roseolamellata HUIF60002 PQ406942 - China This study
M. rubromarginata CBS:265.48 MH856335 MH867890 France (Vu et al. 2019)
M. rubromarginata CBS:268.48 MH856338 MH867891 France (Vu et al. 2019)
M. sanguinolenta TENN59879 FJ596764 - USA (Hughes et al. 2009)
M. seynesii 71h JF908470 - Italy (Osmundson et al. 2013)
M. seynesii 71I JF908469 - Italy (Osmundson et al. 2013)
Mycena sp. JSU008 PQ465299 - China Unpublished
Mycena sp. JSU132 PQ406963 - China This stufy
Mycena sp. 080108 LC504829 - Japan This study
M. silvaenigrae HMJAU43815 MK733310 MK722359 China Unpublished
M. subcaerulea TENN-F-051121 OL711671 OL711666 USA (Na et al. 2022)
M. subcaerulea TENN-F-057919 OL711672 OL711667 USA (Na et al. 2022)
M. subpiligera HUIF50036 OM228758 - China (Liu et al. 2022b)
M. subpiligera HUIFS50007 OM228759 - China (Liu et al. 2022b)
M. subroriduliformis HUIF50540 Holotype PQ406937 PQ406970 China This study
M. subroriduliformis HUIF50546 PQ406938 - China This study
M. substylobates HMJAU43444 MH216190 - China (Na and Bau 2019a)
M. substylobates HMJAU43418 MH216189 - China (Na and Bau 2019a)
M. tenax OSC 113746 EU846251 - USA Unpublished
M. tenax OSC 113728 EU669224 - USA Unpublished
M. vulgaris 447h JF908435 - Italy Unpublished
M. vulgaris CBS:248.47 MH856240 MH867770 France (Vu et al. 2019)
M. xantholeuca CBS370.50 MH856663 MH868180 France (Vu et al. 2019)
M. xantholeuca CBS371.50 MH856664 MH868181 France (Vu et al. 2019)
M. yuezhuoi FFAAS0346 MW581492 - China (Liu et al. 2021a)
M. yuezhuoi HUIF50535 PQ406947 - China (Liu et al. 2021)
M. zephirus CBS:270.48 MH856339 MH867892 France (Liu et al. 2021)
M. zephirus AH60146 PP868143 - Spain (Villarreal et al. 2024)
Phloeomana minutula H6036841 MW540684 - Finland Unpublished
P. speirea iNAT: 100003394 ON206666 - USA Unpublished

Results

Phylogenetic analysis

The two-locus dataset (ITS + LSU) consisted of 191 sequences and 1,680 nucleotide sites in total, which are shown in Table 2. It includes sequences of 28 Mycena taxa except for M. picta (Fr.) Harmaja and M. heteracantha, which are present in Hunan. Sequences of closely related species with high homology and morphologically similar species were also downloaded from GenBank. Atheniella adonis (Bull.) Redhead, Moncalvo, Vilgalys, Desjardin, & B.A. Perry, A. aurantiidisca (Murrill) Redhead, Moncalvo, Vilgalys, Desjardin, & B.A. Perry; Clitocybula intervenosa A.C. Cooper, Desjardin, & B.A. Perry (BAP 588, BAP 613), Phloeomana minutula (Sacc.) Redhead; P. speirea (Fr.) Redhead, and Hydropus murinus A.C. Cooper, Desjardin, & B.A. Perry were chosen as the outgroup (Liu et al. 2022a). The topologies generated from maximum likelihood (ML) and Bayesian inference (BI) analyses were identical, although statistical support for some branches showed slight differences. The BI tree with branch lengths inferred from the ITS and LSU datasets is shown in Fig. 1.

Figure 1. 

Phylogenetic relationships of Mycena species in Hunan Province inferred from the combined dataset (ITS and nrLSU) using Bayesian posterior probabilities (BP) ≥ 0.95; Bootstrap support (BS) ≥ 70% are reported on the branches. Red text represents new taxa. The black dots indicate the Mycena species collected from Hunan Province.

The phylogeny inferred from the combined dataset revealed that the Mycena split into two well-supported clades, and all new taxa formed a well-supported monophyletic lineage. Mycena hongfengensis formed a small branch and grouped with an unidentified Mycena sp. in clade 1 (BS/BP = 100/1.00). Mycena roseolamellata, M. fulvomarginata, M. huangsangensis, and M. subroriduliformis were members of clade 2. Mycena roseolamellata and M. entolomoides T. Bau formed a supported branch in the tree (BS/BP = 100/1.00), and their genetic distance is substantial enough to distinguish between the two species. Mycena fulvomarginata is most related to M. capillaripes Peck. They grouped together with BS/BP = 83/1.00 statistical support, and they were distinct. Mycena albiceps (Peck) Gilliam, M. flosoide L.N. Liu, M. brunnescens L.N. Liu, and our specimens (M. huangsangensis and M. subroriduliformis) formed a separate branch with strong statistical support (BS/BP = 86/1.00).

Taxonomy

Mycena huangsangensis L.N. Liu, sp. nov.

MycoBank No: 856016
Figs 2, 3

Diagnosis.

Differs from the most similar species, M. alniphila, by its decurrent lamellae and longer basidiospores.

Holotype.

China • Hunan Province, Shaoyang City, Suining County, Hunan Huangsang National Nature Reserve, 26°24'18"N, 110°05'37"E, elev. 644 m, 24 April 2024, LiNa Liu, HUIF50526 (collection number NN526).

Etymology.

Refers to the Huangsang National Nature Reserve, from where the holotype was collected.

Description.

Pileus 1–5 mm diam., hemispherical to obtusely conical, expanding with age, umbilicate or depressed center, sulcate, translucent-striate, pruinose and pubescent, light brown (6B4–6B7) to dark brown (6E7–6E8), or brownish-pink (7A4–7B4), paler brown towards margin. Context white, thin, fragile. Lamellae 9–11 reach the stipe, with 1–2 tiers of lamellulae, decurrent, white (4A1) to brown (7D7), edge concolorous with face. Stipe 6–25 × 1–2 mm, cylindrical, hollow, fragile, light brown (6B4–6B7) to brown (6E5–6E8) at the base, gradually becoming paler to white (4A1) towards the apex. Base covered with white fibrils. Odor and taste indistinctive.

Figure 2. 

Basidiomata and microscopic features of Mycena huangsangensis a–f basidiomata g basidiospores h basidia i cheilocystidia j pileipellis k stipitipellis. Structures (g–i, k) were stained in 1% Congo red solution and j were rehydrated in 5% KOH solution. Scale bars: 5 mm (a–f); 5 μm (g); 10 μm (h–k).

Basidiospores 6.0–7.4 (7.8) × (3.2) 3.4–4.2 (4.3) μm, Q = 1.4–2.1, Q = 1.8 ± 0.2, pip-shaped, elongated, hyaline, smooth, thin-walled, amyloid. Basidia 15.4–20.9 × 6.2–9.0 μm, 4-spored, clavate. Cheilocystidia 13.2–25.0 × 8.5–19.9 μm, abundant, clavate to obpyriform, covered with fairly numerous, simple to furcate, cylindrical excrescences. 1.2–9.5 × 0.5–1.4 μm. Pleurocystidia absent. Hyphae of the pileipellis 12–27 μm wide, densely covered with warts or short cylindrical excrescences. Hyphae of the stipitipellis 1.0–3.0 μm wide, densely covered with simple, cylindrical excrescences 1.0–3.2 × 0.8–1.5 μm. Clamp connections are present in the basidia, pileipellis, and stipitipellis hyphae.

Figure 3. 

Morphological features of Mycena huangsangensis a basidiomata b basidia c basidiospores d cheilocystidia e pileipellis f hyphae of stipitipellis. Scale bars: 1 cm (a); 5 μm (c); 10 μm (b, d–f).

Habitat.

Gregarious on decaying leaves of deciduous trees.

Known distribution.

Shaoyang City, Hunan Province.

Additional materials examined.

China• Hunan Province, Shaoyang City, Suining County, Hunan Huangsang National Nature Reserve, 26°24'21"N, 110°05'36"E, elev. 675 m, 24 April 2024, LiNa Liu, HUIF50528 (collection number NN528).

Notes.

Mycena huangsangensis can be considered to be a member of sect. Polyadelphia owing to very small basidiomata, a small number of lamellae, and a slender stipe and hyphae of the pileipellis, which are ornamented with short warts. Mycena huangsangensis belongs to the section with a brownish pileus, while M. alniphila Robich shows the most significant morphological similarity to M. huangsangensis. They have similar basidiomata color and shape of cheilocystidia, pileipellis hyphae densely covered with cylindrical excrescences, and diverticulate stipitipellis hyphae. However, M. alniphila differs in having adnate lamellae, slightly longer spores measuring 8.5–11.0 × (3.5) 4.0–5.5 μm, simple cheilocystidia without branching, and caulocystidia present (Robich 2003). Mycena albiceps and M. catalaunica Robich are somewhat similar to the new species; in particular, they share the same basidiomata shape and similar habitats. Mycena albiceps differs in the white colors of the pileus and black stipe (Gilliam 1976; Maas Geesteranus 1986). The latter, M. catalaunica, has a pale violaceous pink to pale vinaceous pink pileus, subglobose spores, and caulocystidia; the clamp connection is absent in all tissues, and cheilocystidia are subglobose (Robich 2003).

Mycena fulvomarginata L.N. Liu, sp. nov.

MycoBank No: 856027
Figs 4, 5

Diagnosis.

Differs from the closest species, M. rubromarginata, in having yellow lamellae edges and light-yellow contents in cheilocystidia, hyphae of the pileipellis, and stipitipellis.

Holotype.

China• Hunan Province, Suining County, Hunan Huangsang National Nature Reserve, Shaoyang City, 26°25'41"N, 110°03'27"E, elev. 1075 m, 26 June 2021, LiNa Liu, HUIF50088 (collection number NN88).

Etymology.

Refers to the yellow color of the lamellae edges.

Description.

Pileus 4–8 mm diam., hemispherical when young, paraboloid or campanulate with age, sulcate, pellucid-striate, pruinose, apex with obtuse umbo, the margin infrequently out of flatness, dark reddish brown (7C8) at center, gradually becoming paler towards the margin to light brown (7A6), turning purple (12B5) with age. Context white, thin, fragile. Lamellae 10–12 reach the stipe, with 1 tier of lamellulae, adnexed, white (4A1), edge yellow (5B7), stipe 6.0–12.0 × 0.5–1.0 mm, central, cylindrical, hollow, fragile, finely white pruinose and pubescent, pale brown (6D7) to brown (6F7), fading to purple (12B5). Base slightly bulbous, covered with white fibrils. Odor and taste indistinctive.

Figure 4. 

Basidiomata and microscopic features of Mycena fulvomarginata a–d basidiomata e basidiospores f basidia and basidioles g cheilocystidia h pileipellis i stipitipellis. Structures (e, f, i) were stained in a 1% Congo red solution, and g, h were rehydrated in a 5% KOH solution. Scale bars: 5 mm (a–d); 10 μm (e–i).

Basidiospores (7.8) 7.9–9.9 (10.0) × (4.5) 5.7–8.0 (8.2) μm, Q = 1.2–1.7, Q = 1.3 ± 0.1, subglobose to broadly ellipsoid, hyaline, smooth, thin-walled, amyloid. Basidia 24.8–33.5 × 8.2–13.5 μm, 4-spored, clavate. Cheilocystidia 22.6–45.8 × 7.7–17.8 μm, abundant, fusiform to ventricose, clavate, subcylindrical, or somewhat irregularly shaped, smooth or covered with one or more apical simple to furcate excrescences, with light yellow contents. Pleurocystidia absent. Hyphae of the pileipellis 3.0–9.0 μm wide, covered with simple to much-branched excrescences, 2.0–6.0 × 2.0–4.0 μm, with light yellow contents. Hyphae of the stipitipellis 2.0–5.0 μm wide, covered with numerous simple to furcate cylindrical excrescences, 2.0–11.0 × 2.0–8.0 μm, with light yellow contents. Clamp connections are present in the basidia, pileipellis, and stipitipellis hyphae.

Figure 5. 

Morphological features of Mycena fulvomarginata a basidiomata b basidia c basidiospores d cheilocystidia e pileipellis f hyphae of stipitipellis. Scale bars: 5 mm (a); 10 μm (b–f).

Habitat.

Gregarious on moss-covered bark of living deciduous trees.

Known distribution.

Hunan Province and Guangxi Zhuang Autonomous Region, China.

Additional materials examined.

China • Guangxi Zhuang Autonomous Region, Baise City, Leye County, Yachang Orchid National Nature Reserve, 24°46'29"N, 106°20'09"E, elev. 1080 m, 24 April 2024, LiNa Liu, HUIF50089 (collection number NN89).

Notes.

Mycena fulvomarginata belongs to sect. Rubromarginatae Singer ex. Maas Geest. due to the very small basidiomata, yellow lamellar edges, cheilocystidia with colored contents, diverticular pileipellis, and stipitipellis hyphae, along with exhibiting a growth habit on decaying wood. Mycena fulvomarginata is similar to the species described in this section: M. rubromarginata (Fr.) P. Kumm., M. seynii Quél., and M. bulliformis B.A. Perry & Desjardin. They resemble M. fulvomarginata in the shape of their cheilocystidia, are covered with excrescences on the pileipellis and stipe cortical hyphae, and have a similar brown pileus. However, M. rubromarginata differs in that it has longer pileipellis excrescences, up to 36 μm, and cheilocystidia that are 20–65 (up to 90) μm long, with reddish-brown contents (Robich 2003; Aronsen and Læssøe 2016). Mycena seynii should be easy to separate from the new species by its medium basidiomata, larger basidiospores measuring 10.5–15.0 × 6.0–7.5 μm, reddish-purple edge, larger cheilocystidia measuring 30–85 × 8–16 μm, pileipellis, and stipitipellis hyphae with brown to dark red granular contents (Robich 2003; Aronsen and Læssøe 2016). In addition, M. bulliformis differs from M. fulvomarginata by its violet to violet-brown edges, ellipsoid to broadly ellipsoid spores, and some smooth pileipellis hyphae (Perry and Desjardin 2016).

Mycena hongfengensis L.N. Liu, sp. nov.

MycoBank No: 856029
Figs 6, 7

Diagnosis.

Differs from M. castaneicola in having smooth cheilocystidia, dermatocysts present in the pileipellis, and stipitipellis hyphae.

Holotype.

China • Hunan Province, Xiangxi Tujia-Miao Autonomous Prefecture, Jishou City, Hongfeng Forest Park, 28°16'23"N, 109°40'45"E, elev. 230 m, 22 April 2024, ZhuXiang Liu, JSU114 (collection number JD114).

Etymology.

Refers to the Hongfeng Forest Park, from where the holotype was collected.

Description.

Pileus 2–5 mm diam., hemispherical when young, becoming nearly campanulate or plano-convex with age, with a centrally flattened depression, margin smooth, sulcate, translucently striate, pure white (4A1), white pubescent. Context pure white, thin, fragile. Lamellae 16–18 reach the stipe, with 1–2 tiers of lamellulae, narrowly free, pure white (4A1), concolorous with the sides. Stipe 15–40 × 0.1–0.5 mm, almost equal or slightly expanding towards the base, hollow, white (4A1) to greyish-white (4B1), pubescent or puberulous, base swollen. With a not well-developed basal disc, covered with white hirsute. Odor and taste not distinctive.

Figure 6. 

Basidiomata and microscopic features of Mycena hongfengensis a–e basidiomata f basidiospores g, h basidia i cheilocystidia j, k dermatocysts in the pileipellis l dermatocysts in the stipitipellis. Structures (f–j, l) were stained in a 1% Congo red solution and k were rehydrated in a 5% KOH solution. Scale bars: 5 mm (a–c, e); 1 mm (d); 5 μm (f); 10 μm (g–l).

Basidiospores (6.2) 6.3–7.6 (7.7) × (3.4) 3.5–4.9 (5.2) μm, Q = 1.4–2.0, Qm = 1.7 ± 0.2, oblong or pip-shaped, hyaline, thin-walled, amyloid. Basidia 8–16 × 4–8 μm, two- and four-spored, clavate, hyaline. Cheilocystidia 11–43 × 5–9 μm, obpyriform, fusiform, ventricose, filiform, with a long neck, up to 25 μm, with an acute and occasionally branched apex, hyaline. Pleurocystidia absent. Pileipellis hyphae 2–13 μm wide, hyphae cylindrical, densely covered with warts and cylindrical excrescences, 1.0–6.0 × 1.0–2.0 μm, with irregularly cylindric to strangulated dermatocysts, 63–200 × 7–20 μm, walls 1.0–2.0 μm, greenish grey (1C2). Hyphae of the stipitipellis 1.0–6.0 μm wide, smooth, dermatocysts numerous, clavate to pyriform, 50–320 × 5–20 μm, long, flexuous, filiform, simple, and tapering towards the apex. Clamp connections are absent in the basidia, pileipellis, and stipitipellis hyphae.

Figure 7. 

Morphological features of Mycena hongfengensis a basidiomata b basidia c basidiospores d cheilocystidia e dermatocysts in the pileipellis f dermatocysts in the stipitipellis. Scale bars: 1 cm (a); 10 μm (b–f).

Habitat.

Gregarious on decaying leaves of deciduous trees.

Known distribution.

Xiangxi Tujia-Miao Autonomous Prefecture, Hunan Province.

Additional materials examined.

China• Hunan Province, Xiangxi Tujia-Miao Autonomous Prefecture, Jishou City, Hongfeng Forest Park, 28°16'26"N, 109°40'48"E, elev. 255 m, 22 April 2024, ZhuXiang Liu, JSU121 (collection number JD121).

Notes.

Mycena hongfengensis is characterized by its pure white basidiomata, free lamellae, oblong to pip-shaped spores, and the presence of dermatocysts in the pileipellis and stipitipellis. According to the Maas Geesteranus classification, the new species could belong to an uncertain position. When we first found this specimen in the field, we thought it might be a member of either sect. Saccharifera or sect. Amparoina. All species in two sections have a white basidiomata, pubescent pileus and stipe, and stipe with a basic disc (Maas Geesteranus 1983; Desjardin 1995; Na and Bau 2019b). However, M. hongfengensis can be clearly distinguished from other species through microscopic characteristics. The presence of dermatocysts in the pileipellis and stipitipellis is the most important characteristic for separating M. hongfengensis from species of sect. Sacchariferae and sect. Amparoina. Mycena castaneicola T. Bau & Q. Na is the most similar to the new species in the macroscopic characteristics, but the microscopic features differ significantly. Sequences labeled as Mycena sp., which were extracted from the host Myrica rubra (Lour.) Siebold & Zucc. and originated from Japan, along with the sequence of M. hongfengensis, formed a well-supported lineage in the phylogenetic analysis (BS/BP = 100/1.00). We are very fortunate to have collected Mycena sp. in the field. However, we only collected one tiny basidioma; the microscopic structures we observed are not comprehensive. Both taxa have white pileus and pubescent pileus and stipe. The uncertain Mycena species differs in pileipellis, which has “acanthocysts” that are spherical or vesicular and covered with spines. These features can be used to easily differentiate the two species.

Mycena subroriduliformis L.N. Liu, sp. nov.

MycoBank No: 856045
Figs 8, 9

Diagnosis.

Differs from M. surculosa in having a viscid pileus and stipe.

Holotype.

China • Hunan Province, Suining County, Hunan Huangsang National Nature Reserve, Shaoyang City, 26°24'47"N, 110°05'20"E, elev. 542 m, 25 April 2024, LiNa Liu, HUIF50540 (collection number NN540).

Etymology.

Refers to the viscid pileus and stipe of the new species.

Description.

Pileus 2–8 mm diam., hemispherical when young, campanulate with age, with obvious depression at the center, sulcate, translucent-striate, surface wet, glabrous, brownish white (7A1), light brown (7C3), brownish grey (7D1–6D2) when old. Context white, thin, and fragile. Lamellae 18–20 reach the stipe, with 1–2 tiers of lamellulae, decurrent, white (4A1), concolorous with the sides. Stipe 2–45 × 1–2 mm, cylindrical, hollow, fragile, surface glutinous, white (4A1) to brownish grey (5A1–5D3) towards the apex, light brown to brown (5D6–6D6) towards the base, base swollen. Odor and taste indistinctive.

Figure 8. 

Basidiomata and microscopic features of Mycena subroriduliformis a–f basidiomata g basidiospores h basidia i cheilocystidia j, k pileipellis l stipitipellis. Structures (g–i, k, l) were stained in a 1% Congo red solution and j were rehydrated in a 5% KOH solution. Scale bars: 1 cm (a–f); 10 μm (g–l).

Basidiospores (6.2) 6.6–8.5 (9.0) × (3.6) 3.8–5.2 (5.3) μm, Q = 1.5–2.1, Q = 1.7 ± 0.1, pip-shaped, cylindrical, hyaline, amyloid, smooth. Basidia 19.3–26.9 × 5.6–8.0 μm, 4-spored, clavate, hyaline. Cheilocystidia 16.8–26.9 × 6.3–17.1 μm, abundant and variably shaped, clavate to cylindrical with short, more or less branched projections, 1.0–6.0 × 1.0–2.0 μm, thin-walled, hyaline. Pleurocystidia absent. Pileipellis hyphae somewhat gelatinized, 2.0–5.0 μm wide, with irregular simple to branched warts or cylindrical excrescences, 1.0–4.0 × 1.0–2.0 μm. Hyphae of the stipitipellis 1.0–6.0 μm wide, covered with cylindrical excrescences. 1.0–4.0 × 1.0–2.0 μm. Clamp connections are present in the basidia, pileipellis, and stipitipellis hyphae.

Figure 9. 

Morphological features of Mycena subroriduliformis a basidiomata b basidia c basidiospores d cheilocystidia e pileipellis f hyphae of stipitipellis. Scale bars: 5 mm (a); 10 μm (b–f).

Habitat.

Gregarious on decaying leaves of deciduous trees.

Known distribution.

Shaoyang City, Hunan Province.

Additional materials examined.

China• Hunan Province, Suining County, Hunan Huangsang National Nature Reserve, Shaoyang City, 26°24'39"N, 110°05'25"E, elev. 588 m, 25 April 2024, LiNa Liu, HUIF50546 (collection number NN546).

Notes.

The following characteristics placed this new species in the sect. Insignes Maas G. due to the viscid pileus and stipe, decurrent lamellae, pip-shaped spores, clavate cheilocystidia with coarse excrescences, hyphae of the pileipellis embedded in gelatinous matter, and diverticulate (Maas Geesteranus 1989). The three other similar species in sect. Insignes are M. surculosa Maas G. & de Meijer, M. odorifera (Peck) Sacc., and M. calceata Robich. All are differentiated from M. subroriduliformis as follows: M. surculosa has a dry pileus, with only the stipe being viscid. The apical side branches of the pileipellis are densely covered and simple to forked, with cylindrical excrescences. The stipitipellis is somewhat gelatinized (Maas Geesteranus and De Meijer 1997). Mycena odorifera has a distinctive alkaline-like odor, a pruinose stipe, pubescence, and cheilocystidia occasionally with forked apices (Smith 1935). Mycena calceata has dark brown basidiomata and relatively large spores (11.0–13.5 × 5.5–8.0 μm), smooth cheilocystidia, or one or two branches at the apex (Robich 2003).

Mycena roseolamellata L.N. Liu, sp. nov.

MycoBank No: 856026
Figs 10, 11

Diagnosis.

Differs from M. pura in having a brown pileus and pink lamellae.

Holotype.

China• Hunan Province, Ningxiang City, Biandan’ao, Lijingpu Subdistrict, 28°12'07"N, 112°32'43"E, elev. 110 m, 28 November 2023, ShengQiang Liu, HUIF60001 (collection number NN601).

Etymology.

Refers to the pink colors of lamellae.

Description.

Pileus 7–17 mm diam., parabolic when young, then campanulate or broadly conical with age, apex with an obtuse umbo, sulcate, translucent-striate, glabrous, dark brown (6F5–6F8) at first, then turning pale brownish yellow (6A4) to pale brown (6E4) with age, margin brownish white (6A2) to pale brown (6D4–6D6). Context white, fragile, thin. Lamellae 24–26 reach the stipe, with 1–2 tiers of lamellulae, adnate or slightly adnex, white (4A1) when young, pinkish to light pink (9A3–9A2) at maturity, concolorous with faces. Stipe 29–30 × 2.0–3.0 mm, central, cylindrical, hollow, dark brown (6F5-6F8), pale brownish yellow (6A4) to pale brown (6D7) with age, base covered with long, dense, white fibrils. Odor and taste indistinctive.

Figure 10. 

Basidiomata and microscopic features of Mycena roseolamellata a–g basidiomata h basidiospores i, j basidia k, l cheilocystidia m, n pleurocystidia o pileipellis p stipitipellis. Structures (j, p) were stained in 1% Congo red solution, and h, i, k–o were rehydrated in 5% KOH solution. Scale bars: 1 cm (a–g); 10 μm (h–p).

Basidiospores (8.3) 8.6–10.8 (11.5) × (5.3) 5.4–6.3 (6.4) μm, Q = 1.5–2.0, Q = 1.7 ± 0.1, ellipsoid to elongated, hyaline, smooth, thin-walled, amyloid. Basidia 21.7–30.8 × 6.8–8.9 μm, 2-spored, clavate. Cheilocystidia 26.7–84.9 × 8.6–18.7 μm, abundant, fusiform, ventricose-rostrate, obtuse apex, base tapered, with short to long stalk, smooth, hyaline, amyloid, thin-walled. Pleurocystidia similar to cheilocystidia, 52.3–105.7 × 12.3–20.9 μm. Pileipellis 1.0–6.0 μm wide, smooth, terminal hyphae sometimes diverticulate, 1.0–6.0 × 1.0–2.0 μm. Stipitipellis 2.0–5.0 μm, smooth, terminal hyphae sometimes diverticulate, 1.0–4.0 × 1.0–2.0 μm. Clamp connections are absent in the basidia, pileipellis, and stipitipellis hyphae.

Figure 11. 

Morphological features of Mycena roseolamellata a basidiomata b basidia c basidiospores d pleurocystidia e cheilocystidia f pileipellis g hyphae of stipitipellis. Scale bars: 1 cm (a); 10 μm (b–g).

Habitat.

Gregarious on decayed twigs of bamboo or woody debris of deciduous trees.

Known distribution.

Ningxiang City, Hunan Province.

Additional materials examined.

China • Hunan Province, Ningxiang City, Biandan’ao, Lijingpu Subdistrict, 28°12'07"N, 112°32'43"E, elev. 110 m, 22 December 2023, ShengQiang Liu, HUIF60002 (collection number NN602).

Notes.

Mycena roseolamellata is classified into sect. Calodontes based on the smooth cheilocystidia and stipitipellis. Microscopically, M. pura (Pers.) P. Kumm. is the most similar to M. roseolamellata; however, M. pura is distinguished from M. roseolamellata by its purple pileus with pinkish or brown tints, lamellae interveined with age, the presence of clamp connections in all tissues, and the absence of a root-like, pruinose stipe (Robich 2003; Aronsen and Læssøe 2016). Mycena rosea Gramberg is somewhat similar to M. roseolamellata; they have pink lamellae, smooth cheilocystidia, and pleurocystidia. Mycena rosea can be distinguished from M. roseolamellata by having a pink pileus with a dull yellow center, a white or pink stipe, and the presence of clamp connections in all tissues (Robich 2003; Aronsen and Læssøe 2016). Mycena roseolamellata is very different from any species of sect. Calodontes, owing to its brown pileus. Mycena galericulata (Scop.) Gray shares some similarities with M. roseolamellata in terms of pileus color, but M. galericulata is differentiated by the presence of non-smooth cheilocystidia, pileipellis, and stipitipellis (Maas Geesteranus 1992a, 1992b).

Discussion

The combination of morphological features and phylogenetic analyses revealed the presence of 30 species in Hunan Province, including five new species. In the Maas Geesteranus classification, the new species M. fulvomarginata, M. huangsangensis, M. subroriduliformis, and M. roseolamellata are classified into sect. Rubromarginatae, sect. Polyadelphia, sect. Insignes, and sect. Calodontes, respectively (Maas Geesteranus 1980, 1992a, 1992b; Maas Geesteranus and De Meijer 1997; Robich 2003, 2016). However, we could not place the new species M. hongfengensis in any section at present due to its special characteristics. The taxonomy of Mycena is overly complex, and infrageneric classification generally relies on the reported morphology of Mycena, and the characteristics of some species are not consistent with the common features shared by species of the numerous sections. Therefore, we need to increase the number of related species to identify the common characteristics of those species and further improve the taxonomy of Mycena.

Among the 30 Mycena species collected in Hunan, we found M. picta (Fr.) Harmaja on Yuelu Mountain, Hunan. As we obtained only one basidiomata in the field, sequences of M. picta could not be obtained for phylogenetic analysis. However, we can confirm that the specimen is M. picta based on its distinct macroscopic and microscopic characteristics (Aronsen and Læssøe 2016; Bau et al. 2021). And M. heteracantha was also collected in Yuelu Mountain (Na and Bau 2019a). We are uncertain of the classification of M. juniperina Aronsen and M. meliigena (Berk. & Cooke) Sacc. Distinguishing the 8 related specimens we collected in Hunan was difficult because of their similar microcharacteristics and the limited number of sequences available for downloading from GenBank, although the color of their basidiomata varies. Therefore, we identified all the related specimens we collected as M. juniperina/meliigena, as we did not obtain type specimens. Additional studies are needed.

Among the phylogenetic trees, M. hongfengensis, M. deeptha Aravind. & Manim. and M. pluteoides T. Bau & Q. Na were grouped into clade 1. Interestingly, all species have a non-smooth pileipellis hyphae. Most species collected from Hunan are mainly concentrated in clade 2. Four new species are grouped in clade 2; all Mycena species in this clade have a stipe or a stipe base that is covered with white fibrils.

The distinct topography, climate, and flora of Hunan Province have produced abundant and distinctive Mycena specimens. We conducted a comparative analysis of the geographical compositions of the 30 Mycena species we collected and preliminarily divided the distribution of the genus Mycena in Hunan Province into the following four types (Table 1). The analysis of the floristic components of the species reveals that Mycena in Hunan Province is cosmopolitan, exhibiting tropical-subtropical floristic characteristics as well as a certain proportion of northern temperate characteristics, indicating that Mycena species in Hunan possess diverse and transitional features.

Although Mycena was widely distributed in the world, the earliest and most detailed research was conducted in Europe and North America (Singer and Digilio 1951; Robich 2003, 2016). The morphological and microscopic characteristics of the Mycena species collected in Hunan Province are basically consistent with those of materials from Europe. However, the sizes of the basidiomata, basidiospores, basidia, cheilocystidia, and caulocystidia occasionally vary subtly. This may be due to the geographical location and environmental changes in the species at that time. From a geographical point of view, the Mycena species we collected show some differences compared with those documented in monographs from Europe and North America. Free lamellae are important for diagnosing these species in Mycena. We found that Mycena with free lamellae are more commonly collected in Hunan Province than in temperate regions in China; for example, M. deeptha and M. pluteoides, belonging to sect. Exornatae Maas Geest, are widely distributed in the western and southern areas of Hunan. And members of the sect. Exornatae are most commonly found in subtropical regions of Asia (Desjardin et al. 2010; Aravindakshan et al. 2012; Aravindakshan and Manimohan 2013a; Bau et al. 2021). These findings also indirectly suggest that sect. Exornatae has a certain preference for geographical location. Our research may contribute to the exploration of the origin and evolution of Mycena.

Key to the known species of Mycena in Hunan Province

1 Stipe arising from a basal disc 2
– Stipe not arising from a basal disc 3
2 Dermatocysts present in the pileipellis and stipitipellis M. hongfengensis
– Dermatocysts absent in the pileipellis and stipitipellis 4
3 Fresh and young basidiomata exude colored fluid when damaged M. haematopus
– Basidiomata do not exude colored fluid when damaged 7
4 Pileus glabrous, viscid, white, with a pale brown center, depressed at the center 5
– Pileus dry, pubescent, pure white, not depressed at center 6
5 Cheilocystidia vesiculose, smooth M. deeptha
– Cheilocystidia densely covered with projections M. pluteoides
6 Basidiomata growing on Castanea burs, pileus slightly pubescent M. castaneicola
– Basidiomata growing on dead wood or humus layer, pileus with bran-like covering M. heteracantha
7 Lamellae not white, or occasionally white when young 8
– Lamellae white 9
8 Lamellae faces pink, occasionally white when young, cheilocystidia hyaline M. roseolamellata
– Lamellae faces orange-yellow, cheilocystidia with yellow contents M. leaiana
9 Cheilocystidia smooth 10
– Cheilocystidia with simple to branched excrescences 21
10 Lamellae faces not concolorous with the sides 11
– Lamellae faces concolorous with the sides 12
11 Lamellae edges light yellow M. citrinomarginata
– Lamellae edges light brown to yellowish-brown M. fulvomarginata
12 Hyphae of the pileipellis smooth 13
– Hyphae of the pileipellis diverticulate 16
13 Pileus brown M. algeriensis
– Pileus violet 14
14 Pleurocystidia absent 15
– Pleurocystidia present M. pura
15 Pileipellis not gelatinized M. yuezhuoi
– Pileipellis gelatinized M. pearsoniana
16 Lamellae adnate or adnexed 17
– Lamellae decurrent 18
17 Pileus grey brown 19
– Pilues white 20
18 Cheilocystidia thick-walled M. subpiligera
– Cheilocystidia thin-walled M. digitifurcata
19 Hyphae of the stipitipellis smooth, caulocystidia present M. leptocephala
– Hyphae of the stipitipellis covered with warty or diverticulae, caulocystidia absent M. abramsii
20 Pileipellis and stipitipellis gelatinized M. laevigata
– Pileipellis and stipitipellis not gelatinized M. adnexa
21 Basidiomata sticky M. subroriduliformis
– Basidiomata dry 22
22 Pileus with yellow tone 23
– Pileus without yellow tone 24
23 Pileus bucket-shaped, lamellae broader than the length M. picta
– Pileus hemispherical, parabolical to convex, lamellae broad M. meliigena / M. juniperina
24 Basidiospores globose M. corynephora
– Basidiospores broadly ellipsoid to ellipsoid 25
25 Pileus and lamella with red spots when old M. maculata
– Pileus and lamella without red spots when old 26
26 Rhizomorphs present M. chlorocyanea
– Rhizomorphs absent 27
27 Clamp connections absent in all tissues M. galericulata
– Clamp connections present in all tissues 28
28 Acanthocysts present, acanthocysts of two types, pyriform to vesicular M. bicystidiata
– Acanthocysts absent 29
29 Pileus and stipe pruninose, iodoform when dry M. filopes
– Pileus and stipe glabrous, odor indistinctive M. huangsangensis

Acknowledgements

Thanks to Prof. Zuo Hong Chen of Hunan Normal University for providing help in fieldwork. Thanks to Prof. Xun Lin Yu from the Central South University of Forestry and Technology for his help in plant identification. Thanks to Ms. Shuang Qu and Ms. Chang Zhuo Liu for helping with the article layout design and specimen collection and to Mr. Sheng Qiang Liu and Mr. Wu Ping Luo for providing the specimens in this study.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was supported by the Hunan Natural Science Foundation (2023JJ50073) and the Graduate Scientific Research Innovation Project of Jishou University (Jdy22072).

Author contributions

Data curation: YT. Investigation: SNL, YT. Resources: JBL, LGF. Validation: SNL, ARS, BMS. Writing - original draft: YXX. Writing - review and editing: ZXL, ZMT, LNL.

Author ORCIDs

Ying Xin Xiao https://orcid.org/0009-0008-7041-503X

Li Na Liu https://orcid.org/0000-0003-3241-4016

Zhu Ming Tan https://orcid.org/0000-0001-6696-1179

Ai Rong Shen https://orcid.org/0000-0003-4543-0250

Bao Ming Shen https://orcid.org/0000-0003-0110-3370

Yun Tan https://orcid.org/0000-0002-8129-0163

Sai Nan Li https://orcid.org/0000-0002-8818-446X

Zhu Xiang Liu https://orcid.org/0000-0001-6014-3554

Data availability

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

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Ying Xin Xiao and Li Na Liu contributed equally to this work.
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