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Research Article
Morphological and phylogenetic analyses reveal five new species of Porotheleaceae (Agaricales, Basidiomycota) from China
expand article infoQin Na‡, Hui Zeng§, Yaping Hu|, Hui Ding|, Binrong Ke§, Zhiheng Zeng§, Changjing Liu, Xianhao Cheng‡, Yupeng Ge‡§
‡ Ludong University, Yantai, China
§ Fujian Academy of Agricultural Sciences, Fuzhou, China
| Nanjing Institute of Environmental Sciences, Nanjing, China
¶ Nanjing Police University, Nanjing, China

Corresponding author: Yupeng Ge ( gaiyupeng@126.com )

Academic editor: Maria-Alice Neves
© 2024 Qin Na, Hui Zeng, Yaping Hu, Hui Ding, Binrong Ke, Zhiheng Zeng, Changjing Liu, Xianhao Cheng, Yupeng Ge.
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: Na Q, Zeng H, Hu Y, Ding H, Ke B, Zeng Z, Liu C, Cheng X, Ge Y (2024) Morphological and phylogenetic analyses reveal five new species of Porotheleaceae (Agaricales, Basidiomycota) from China. MycoKeys 105: 49-95. https://doi.org/10.3897/mycokeys.105.118826
Open Access

Abstract

The first occurrence of Marasmiellomycena and Pulverulina in the Chinese mycobiota are reported, M. tomentosa and P. flavoalba, two new species and M. albodescendens, a new combination, revealed by phylogenetic analyses and morphological study. These newly-recorded genera, Marasmiellomycena, which can be distinguished by their agaricoid basidiomata, dark-coloured stipe, sarcodimitic tramal structure, stipitipellis with yellow to yellowish-brown pigments and yellow-pigmented thick-walled caulocystidia and Pulverulina, which differs from other genera of Porotheleaceae by its pruinose stipe, decurrent lamellae, inamyloid basidiospores and absence of hymenial cystidia. We also formally describe three other new species of Porotheleaceae collected from Chinese temperate to subtropical zones of Fujian and Zhejiang Provinces: Clitocybula fuscostriata, Gerronema brunneosquamulosum and Leucoinocybe subglobispora. Furthermore, we include the results of a phylogenetic analysis of Porotheleaceae, based on a multi-locus (ITS, nrLSU and rpb2) dataset. According to this analysis, Chrysomycena, Clitocybula, Delicatula, Hydropodia, Hydropus, Leucoinocybe, Marasmiellomycena, Megacollybia, Pulverulina, Trogia and Vizzinia are monophyletic. However, Gerronema is identified as polyphyletic and, additionally, Porotheleum does not form a monophyletic group either because Porotheleum parvulum and Porotheleum albidum are “unassigned” in phylogenetic analysis. The results of our phylogenetic analyses, coupled with morphological observations, confirm recognition of these new taxa. Morphological descriptions, photographs, line drawings and comparisons with closely-related taxa are presented for the new species. A key to the 22 species belonging to nine genera of Porotheleaceae in China is also provided.

Key words

cyphelloid polypores, new taxon, Porotheleum, systematics, white-spored omphalinoid fungi

Introduction

The family Porotheleaceae (order Agaricales), formally proposed by Murrill (1916), comprises saprotrophic, mainly wood-decaying fungi that are primarily agarics, but also include cyphelloid fungi. The type genus, Porotheleum Fr., is distinguished by fruiting in clusters of small cup-shaped to tubular cream cyphelloid basidiomes, whereas other genera are typically agaricoid (Vizzini et al. 2022). Previous taxonomic studies have included 15 genera in Porotheleaceae: Chrysomycena Vizzini, Picillo, Perrone & Dovana, Clitocybula (Singer) Singer ex Métrod, Delicatula Fayod, Gerronema Singer, Hydropodia Vizzini & Consiglio, Hydropus Kühner ex Singer, Leucoinocybe Singer ex Antonín, Borovička, Holec & Kolařík, Lignomphalia Antonín, Borovička, Holec & Kolařík, Marasmiellomycena De la Peña-Lastra, Mateos, Kolařík, Ševčíková & Antonín, Megacollybia Kotl. & Pouzar, Porotheleum, Pulverulina Matheny & K.W. Hughes, Pseudohydropus Vizzini & Consiglio, Trogia Fr. and Vizzinia Ševčíková & Kolařík (Antonín et al. 2019; Vizzini et al. 2019, 2022; Matheny et al. 2020; Senanayake et al. 2023). Most taxa, except for Porotheleum, are well characterised, based on the following features: a saprophytic habit; omphalinoid, collybioid, to clitocyboid basidiomata; partly to entirely pigmented pileus; adnexed, subdecurrent, to decurrent lamellae; smooth, thin-walled basidiospores; and the frequent presence of sarcodimitic tramal tissues (Singer 1951, 1982; Redhead 1987; Norvell et al. 1994; Hughes et al. 2007; Kumar and Manimohan 2009; Yang et al. 2012; Vizzini et al. 2019; Consiglio et al. 2022; Senanayake et al. 2023). Species of Porotheleaceae are widespread in subtropical to tropical regions and tend to be lower diversity in temperate zones (Singer 1951, 1970; Norvell et al. 1994; Antonín and Noordeloos 2004; Hughes et al. 2007; Antonín et al. 2019; Vizzini et al. 2019; Consiglio et al. 2022; Na et al. 2022a; Senanayake et al. 2023). Six new genera have recently been recognised: Chrysomycena, Hydropodia, Marasmiellomycena, Pulverulina, Pseudohydropus and Vizzinia (Vizzini et al. 2019; Matheny et al. 2020; Consiglio et al. 2022; Senanayake et al. 2023). These newly-described genera have been found in diverse regions, predominantly in Europe and North America, with some findings in Oceania, Africa and Asia, but the distribution reflects the broad yet unequal exploration of this family’s species, only one species is from Japan, in Asia and none from China (Cooper 2014; Vizzini et al. 2019; Villarreal et al. 2021; Consiglio et al. 2022; Kasuya et al. 2023; Senanayake et al. 2023). While Index Fungorum (http://www.indexfungorum.org/Names/Names.asp 2023.3.16) records 670 Porotheleaceae species, only seven species are documented from China, indicating a disparity in mycological research within the region (Liu 1995; Yang et al. 2012; Liu et al. 2019; Na et al. 2022a).

A comprehensive phylogenetic analysis of Porotheleaceae has not been performed because few sequences are available. Prior to 2012, the family was informally cited in literature as the ‘hydropoid’ clade within the ‘marasmioid’ clade (Moncalvo et al. 2002; Matheny et al. 2006; Antonín et al. 2019). Many authors have since suggested that members of the hydropoid clade should be placed in the phylogenetically defined Porotheleaceae clade (Henrici 2012; Redhead 2013; Cooper 2016; Vizzini et al. 2019, 2022; Kalichman et al. 2020; Matheny et al. 2020; Senanayake et al. 2023). According to a study based on the large subunit of nuclear ribosomal DNA (nrLSU) sequences (Moncalvo et al. 2002), eight species in five genera (Clitocybula, Gerronema, Hydropus, Megacollybia and Porotheleum) constitute this hydropoid (/hydropoid) clade. The results of that study also support the placement of Megacollybia and Clitocybula as close relatives of Hydropus. Moncalvo et al. (2002) also proposed that Gerronema sensu Singer (1986) was polyphyletic (Lutzoni 1997; Moncalvo et al. 2000), whereas this genus as delineated by Norvell et al. (1994) was monophyletic. However, the type species of Gerronema was not included in the molecular phylogeny of Moncalvo et al. (2002). The delimitation of Gerronema by Norvell et al. (1994) was based solely on morphology in comparison to an epitype, with emphasis on the presence of sarcodimitic tissue. The hydropoid clade configuration defined by Moncalvo et al. (2002) based on ribosomal LSU is also presented in Bodensteiner et al. (2004). In a multigenic analysis (18S, 5.8S, 25S, rpb1 and rpb2) performed by Matheny et al. (2006), the hydropoid clade included Clitocybula atrialba (Murrill) Singer [currently Gerronema atrialbum (Murrill) Borovička & Kolařík], Clitocybula oculus (Peck) Singer, Henningsomyces candidus (Pers.) Kuntze, Hydnopolyporus fimbriatus (Cooke) DA Reid (currently Irpex rosettiformis C.C. Chen & Sheng H. Wu), Hydropus marginellus (Pers.) Singer, Hydropus cf. scabripes (Murrill) Singer [currently Mycopan scabripes (Murrill) Redhead, Moncalvo & Vilgalys], Megacollybia platyphylla (Pers.) Kotl. & Pouzar and several species formerly placed in Mycena (Pers.) Roussel [i.e. Mycena auricoma Har. Takah. (currently Leucoinocybe auricoma (Har. Takah.) Matheny), Mycena amabilissima (Peck) Sacc. (currently Atheniella amabilissima (Peck) Redhead, Moncalvo, Vilgalys, Desjardin & B.A. Perry) and Mycena aurantiidisca (Murrill) Murrill (currently Atheniella aurantiidisca (Murrill) Redhead, Moncalvo, Vilgalys, Desjardin & B.A. Perry)]. Henrici (2012) combined Megacollybia, Clitocybula and Hydropus, along with other genera, into the family Porotheleaceae, comprising a total of 19 genera. Redhead (2012, 2013) expanded the ‘hydropoid’ clade by introducing Atheniella Redhead, Moncalvo, Vilgalys, Desjardin & B.A. Perry and established the genus Phloeomana Redhead within the family Porotheleaceae. Cooper (2016) also believes that Porotheleum should belong to the Porotheleaceae family, despite the possibility of misidentification in the sequenced material of Porotheleum fimbriatum (generic type). Finally, Antonín et al. (2019) introduced the new genera Leucoinocybe and Lignomphalia, which were separated from Clitocybula. However, it should be noted that Singer (1943) originally proposed Leucoinocybe as a provisional name, rendering the use by Antonín et al. (2019) as a validation rather than the establishment of a completely new genus. In an analysis by Vizzini et al. (2019), Porotheleaceae was statistically well supported (MLB = 100%) when only Hydropus, Clitocybula, Leucoinocybe, Megacollybia, Porotheleum, Trogia and some species of Gerronema were included. In addition, Chrysomycena formed a distinct monophyletic lineage corresponding to a separate genus, sister to a clade formed by Megacollybia, Trogia and some species of Gerronema (Vizzini et al. 2019). Matheny et al. (2020) performed a phylogenetic analysis of a combined ITS–28S dataset of 73 taxa and found that Delicatula and Pulverulina (representing a new genus) are members of Porotheleaceae sensu Vizzini et al. (2019); this was in agreement with the concept of Porotheleaceae s.l. of Kalichman et al. (2020), which comprises Porotheleaceae sensu Vizzini et al. (2019), Actiniceps Berk. & Broome, Atheniella, Calyptella Quél., Chaetotyphula Corner, Hemimycena Singer, Lignomphalia, Phloeomana and Scytinotus P. Karst. Vizzini et al. (2022) considered the family Porotheleaceae to be equivalent to Porotheleaceae sensu Vizzini et al. (2019) and included the other taxa in Porotheleaceae s.l. Kalichman et al. (2020) in Cyphellaceae Burnett, a sister family to Porotheleaceae. Senanayake et al. (2023) agree with the concept and composition of Porotheleaceae as defined by Vizzini et al. (2019, 2022) and proposed two new genera of the family, Marasmiellomycena and Vizzinia. Finally, Hydropus subalpinus (Höhn.) Singer, which was not aggregated into clade Hydropus with high statistical support, was recently treated as Hydropodia subalpina (Höhn.) Vizzini, Consiglio & M. Marchetti by Consiglio et al. (2022). In the same study, Pseudohydropus Vizzini & Consiglio was established, with Pseudohydropus floccipes (Fr.) Vizzini & Consiglio designated as the type species, comprising a total of four species.

Seventeen species belonging to seven genera of Porotheleaceae, namely, one species of Clitocybula (Singer) Singer ex Métrod, one species of Delicatula, seven species of Gerronema, four species of Hydropus, one species of Leucoinocybe, two species of Megacollybia and one species of Trogia, have been recognised in China as of 2023 (Liu 1995; Dai et al. 2010; Yang et al. 2012; Liu et al. 2019; Wang et al. 2021; Na et al. 2022a). Progress has recently been made in clarifying the status of mycenoid and omphalinoid fungi in China, including the discovery of four new taxa from Anhui, Fujian and Zhejiang Provinces: Gerronema baishanzuense Q. Na, H. Zeng & Y.P. Ge; G. microcarpum Q. Na, H. Zeng & Y.P. Ge; G. zhujian Q. Na, H. Zeng & Y.P. Ge; and Leucoinocybe lishuiensis Q. Na, H. Zeng & Y.P. Ge (Na et al. 2021, 2022a). As part of our ongoing research on omphalinoid fungi, we uncovered the first occurrence of two newly-recorded genera, Marasmiellomycena and Pulverulina, including two new species and we incorporated one species from Porotheleum into Marasmiellomycena. We also discovered three new species belonging to Clitocybula, Gerronema and Leucoinocybe in temperate and subtropical China. We accordingly present a morphological description of the new species and provide an identification key to the 22 species of Porotheleaceae currently known from China.

Materials and methods

Specimens and morphology

Macroscopic descriptions were based on the study of fresh specimens, whereas micromorphological descriptions relied on dried materials. In our descriptions, colour abbreviations follow the colour standards and colour nomenclature of Ridgway (1912). Microscopic observations were made on dried specimens mounted in 5% potassium hydroxide (KOH) and stained with Congo red when necessary. The prepared specimens were observed under a Lab A1 microscope (Carl Zeiss AG, Jena, Germany) and photographed and recorded using ZEN 2.3 software (Carl Zeiss AG). Melzer’s reagent was used to test whether spores and tissues were amyloid (Horak 2005). Twenty mature basidiospores from each basidiomata (two basidiomata per holotype) were measured in side view. Sizes of basidiospores were recorded, with the notation [a/b/c] used at the beginning of each entry in the description to indicate a basidiospores from b basidiomata of c specimens were measured. Measured sizes (including Q values) are given in the description as (d)e–f–g(h) × (i)j–k–l(m), where d is the smallest length, e–g represents the range of at least 90% of values, f is the average length and h is the largest value; width (i–m) is expressed in the same way. In addition, Q stands for the length-width ratio of a spore and Q ± av is the average Q of all basidiospores ± the sample standard deviation (Ge et al. 2021; Liu et al. 2021, 2022; Na et al. 2021, 2022a, 2022b). Hyphae of the pileipellis and stipitipellis and a total of 20 basidia, cheilocystidia and caulocystidia were measured from each collection. The examined collections have been deposited in the fungarium of the Fujian Academy of Agricultural Sciences (FFAAS), China. Author abbreviations follow Index Fungorum (http://www.indexfungorum.org).

DNA extraction, polymerase chain reaction (PCR) amplification and sequencing

Genomic DNAs of the putative new species were extracted from dried materials using a NuClean PlantGen DNA kit (Kangwei Century Biotechnology Co., Beijing, China). Gene regions were amplified using the following primer pairs: ITS1/ITS4 (White et al. 1990) for 5.8S and internal transcribed spacer ITS1 and ITS2 regions (ITS), LR0R/LR7 (Hopple and Vilgalys 1999) for the large subunit of nuclear ribosomal DNA (nrLSU) and bRPB2-6f/bRPB2-7.1R (Matheny 2005) for the second largest subunit of RNA polymerase II (rpb2). Amplifications were performed in 25 µl reaction mixtures consisting of 9.5 μl ddH2O, 12.5 µl 2× UTaq PCR Master Mix (Zoman Biotechnology Co., Beijing, China), 1 µl of each primer (10 mM) and 1 μl DNA template. PCR amplification of the ITS region used the following protocol: initial denaturation at 95 °C for 4 min, followed by 34 cycles of 94 °C for 45 s, 52 °C for 45 s and 72 °C for 1 min and a final extension at 72 °C for 10 min. Cycling conditions used for amplification of the nrLSU were as follows: initial denaturation at 93 °C for 2 min, followed by 20 cycles of 93 °C for 1 min, 50 °C for 1 min and 72 °C for 1 min and a final extension at 72 °C for 10 min. The PCR protocol for rpb2 amplification was as follows: initial denaturation at 93 °C for 2 min, 20 cycles of 93 °C for 1 min, 50 °C for 1 min and 72 °C for 1 min, 20 cycles of 93 °C for 1 min, 53 °C for 1 min and 72 °C for 1 min and a final extension at 72 °C for 10 min. The PCR products were subjected to Sanger dideoxy sequencing at the Beijing Genomics Institute (Beijing, China).

Phylogenetic analysis

For phylogenetic analysis, we constructed a concatenated dataset of 168 ITS, 87 nrLSU and 14 rpb2 sequences from 58 taxa of 14 genera of Porotheleaceae. In addition, six sequences (three ITS and three nrLSU) of Mycena purpureofusca (Peck) Sacc. were included as outgroups according to the results of Na et al. (2022a). Sequences retrieved from GenBank and those obtained in this study are listed in Table 1. Alignments were performed in Mafft 7.376 (Katoh and Standley 2013). Sequence editing and necessary adjustments were carried out in BioEdit 7.0.4.1 and Clustal X 1.81 (Thompson et al. 1997; Hall 1999). Bayesian Inference (BI) and Maximum Likelihood (ML) bootstrap analyses were performed using the best-fit substitution models identified in ModelTest 3.7 (Posada and Crandall 1998). The BI analysis was carried out in MrBayes 3.2.6 (Ronquist and Huelsenbeck 2003). Runs of 1,000,000 generations, with trees sampled every 100th generation, were initiated for eight heated and one cold Markov chain(s). Analyses were automatically terminated when the average standard deviation of split frequencies reached a value below 0.01 and the first 25% of trees were discarded as burn-in (Ronquist and Huelsenbeck 2003). The ML analysis was performed in RAxML GUI 2.0 using a rapid bootstrapping algorithm involving 1,000 replicates (Edler et al. 2021). The aligned datasets for Bayesian and ML analyses have been deposited in TreeBASE (submission ID 31062; study accession URL: http://purl.org/phylo/treebase/phylows/study/TB2:S31062). Phylogenetic trees were displayed using FigTree v.1.4.3.

Table 1.
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Specimens used in phylogenetic analysis, with geographic origin and GenBank accession numbers.

No. Taxa Voucher Locality ITS Sequence ID LSU Sequence ID rpb2 Sequence ID Reference
1 Chrysomycena perplexa MCVE:30184 TYPE Italy NR172974 NG071251 – Vizzini et al. (2019)
2 Clitocybula albida CUH AM064 India MG250188 – – Dutta et al. (2018)
3 Clitocybula albida CUH AM065 India MG250189 – – Dutta et al. (2018)
4 Clitocybula abundans STU:SMNS-B-FU-2017/00898 Germany MF627833 – – Unpublished
5 Clitocybula familia 2319-QFB-25741 Canada KM406970 – – Unpublished
6 Clitocybula familia PRM 921866 Czech Republic JF730327 JF730320 – Antonín et al. (2011)
7 Clitocybula familia BRNM 736053 Slovakia JF730328 JF730323 – Antonín et al. (2011)
8 Clitocybula familia STU:SMNS-B-FU-2017/00926 Germany MF627834 – – Unpublished
9 Clitocybula familia NAMA 2017-349 USA MH979253 – – Unpublished
10 Clitocybula fuscostriata FFAAS1029 China OR238881 OR238893 OR258374 This study
11 Clitocybula fuscostriata FFAAS1030 Holotype China OR238882 OR238894 OR258375 This study
12 Clitocybula fuscostriata FFAAS1031 China OR238883 OR238895 OR258376 This study
13 Clitocybula lacerata LE 6639 Russia HM191746 – – Malysheva and Morozova (2011)
14 Clitocybula lacerata LE 262744 Russia HM191747 – – Malysheva and Morozova (2011)
15 Clitocybula lacerata LE 262743 Russia HM191748 – – Malysheva and Morozova (2011)
16 Clitocybula lacerata PRM 915404 Czech Republic LT854054 LT854030 – Antonín et al. (2019)
17 Clitocybula lacerata WU 19575 Austria LT854053 LT854031 – Antonín et al. (2019)
18 Clitocybula oculus 3512 Canada KM406971 – – Unpublished
19 Clitocybula oculus WU 20008 Canada LT854017 LT854017 – Antonín et al. (2019)
20 Clitocybula oculus S.D. Russell iNaturalist # 8606755 India MN906165 – – Unpublished
21 Clitocybula oculus S.D. Russell iNaturalist # 8591258 India MN906164 – – Unpublished
22 Clitocybula oculus BIOUG24046-B03 Canada KT695321 – – Telfer et al. (2015)
23 Clitocybula oculus AFTOL-ID 1554 USA DQ192178 DQ192178 – Matheny et al. (2006)
24 Delicatula integrella KA12-1305 Korea KR673538 – – Kim et al. (2015)
25 Delicatula integrella S.D. Russell MycoMap # 6067 USA MN906231 – – Unpublished
26 Delicatula integrella G0060 USA – MK277924 – Varga et al. (2019)
27 Gerronema baishanzuense FFAAS0359 Holotype China OL985962 OL985984 – Na et al. (2022a)
28 Gerronema baishanzuense FFAAS0360 China OL985963 – – Na et al. (2022a)
29 Gerronema baishanzuense FFAAS0361 China OL985964 OL985985 – Na et al. (2022a)
30 Gerronema baishanzuense FFAAS0362 China OL985965 OL985986 – Na et al. (2022a)
31 Gerronema baishanzuense FFAAS0363 China OL985966 OL985987 – Na et al. (2022a)
32 Gerronema baishanzuense FFAAS0366 China OL985967 OL985988 – Na et al. (2022a)
33 Gerronema brunneosquamulosum FFAAS1032 Holotype China OR238884 OR238896 OR258377 This study
34 Gerronema brunneosquamulosum FFAAS1033 China OR238885 OR238897 OR258378 This study
35 Gerronema indigoticum HMJAU 47636 China MK693727 MK693732 – Liu et al. (2019)
36 Gerronema indigoticum HMJAU 47942 China MK693728 MK693733 – Liu et al. (2019)
37 Gerronema indigoticum HMJAU 47943 China MK693729 MK693734 – Liu et al. (2019)
38 Gerronema keralense 2 India MH156555 NG_064531 – Latha et al. (2018)
39 Gerronema keralense BKF10263 Thailand MZ452107 MZ452144 Direct Submission
40 Gerronema kuruvense CAL 1665 India NG_159831 NG_064530 – Latha et al. (2018)
41 Gerronema kuruvense BKF10266 Thailand MZ452090 MZ452669 – Direct Submission
42 Gerronema kuruvense DCY3362(HGASMF01-15010) Chian MZ951144 – – Direct Submission
43 Gerronema microcarpum FFAAS0365 China – OL985989 – Na et al. (2022a)
44 Gerronema microcarpum FFAAS0371 China OL985968 OL985990 – Na et al. (2022a)
45 Gerronema microcarpum FFAAS0372 China OL985969 OL985991 – Na et al. (2022a)
46 Gerronema microcarpum FFAAS0373 Holotype China OL985970 OL985992 – Na et al. (2022a)
47 Gerronema microcarpum FFAAS0374 China OL985971 – – Na et al. (2022a)
48 Gerronema microcarpum FFAAS0375 China OL985972 OL985993 – Na et al. (2022a)
49 Gerronema nemorale KACC 43599 Korea EU883592 – – Unpublished
50 Gerronema nemorale KACC 43600 Korea EU883593 – – Unpublished
51 Gerronema nemorale not indicated Korea EU883594 – – Unpublished
52 Gerronema nemorale FA249 Pakistan MN744686 – – Aqdus and Khalid (2021)
53 Gerronema nemorale FA236 Pakistan MN744687 – – Aqdus and Khalid (2021)
54 Gerronema nemorale FA239 Pakistan MN744688 – – Aqdus and Khalid (2021)
55 Gerronema strombodes DJL05NC72 USA EU623639 – – Hughes et al. (2007)
56 Gerronema strombodes TFB12519/TENN60718 USA EU623640 – – Hughes et al. (2007)
57 Gerronema strombodes TFB12783/TENN61350 USA EU623641 – – Hughes et al. (2007)
58 Gerronema strombodes TFB11947 clone C2 USA KY242503 – – Hughes et al. (2007)
59 Gerronema strombodes TFB11947 clone C3 USA KY242504 – – Hughes et al. (2007)
60 Gerronema strombodes TFB11947 clone C5 USA KY242506 – – Hughes et al. (2007)
61 Gerronema strombodes TFB14234 USA KY242507 – – Hughes et al. (2007)
62 Gerronema strombodes TFB14514 USA KY242509 – – Hughes et al. (2007)
63 Gerronema strombodes TFB11947 USA KY271083 – – from GenBank
64 Gerronema subclavatum Redhead 5175, DAOM not indicated U66434 – – Lutzoni (1997)
65 Gerronema subclavatum FLAS-F-60986 USA MH016932 – – from GenBank
66 Gerronema subclavatum FLAS-F-61518 USA MH211945 – – from GenBank
67 Gerronema subclavatum Smith-2018 USA MK573888 – – Direct Submission
68 Gerronema subclavatum Mushroom Observer # 243440 USA MK607510 – – Direct Submission
69 Gerronema subclavatum iNaturalist # 8545787 India MN906021 – – from GenBank
70 Gerronema subclavatum S.D. Russell MycoMap # 6854 India MN906138 – – from GenBank
71 Gerronema waikanaense PDD:87667 New Zealand JQ694117 – – from GenBank
72 Gerronema wildpretii BRNM 788347 Madeira LT854045 LT854043 – Antonin et al. (2019)
73 Gerronema xanthophyllum PRM 924657 Czech Republic LT854023 LT854023 – Antonin et al. (2019)
74 Gerronema zhujian FFAAS0364 China OL985973 OL985994 – Na et al. (2022a)
75 Gerronema zhujian FFAAS0370 China OL985974 OL985995 – Na et al. (2022a)
76 Gerronema zhujian FFAAS0376 Holotype China OL985975 OL985996 – Na et al. (2022a)
77 Hydropodia subalpina (≡Hydropus subalpinus) STU:SMNS-STU-F-0900123 Germany MF039248 – – Eberhardt et al. (2018)
78 Hydropodia subalpina (≡Hydropus subalpinus) Montri-291 not indicated MK028414 – – Unpublished
79 Hydropodia subalpina (≡Hydropus subalpinus) Montri-312 not indicated MK028415 – – Unpublished
80 Hydropodia subalpina (≡Hydropus subalpinus) Montri-323 not indicated MK028416 – – Unpublished
81 Hydropodia subalpina (≡Hydropus subalpinus) OKA-TR-K364 Turkey MN701620 MN700170 – Unpublished
82 Hydropodia subalpina (≡Hydropus subalpinus) OKA-TR-K380 Turkey MN701621 MN700171 – Unpublished
83 Hydropodia subalpina (≡Hydropus subalpinus) OKA-TR-B400 Turkey MN701622 MN700172 – Unpublished
84 Hydropus atramentosus 918 Italy JF908050 – – Osmundson et al. (2013)
85 Hydropus fuliginarius S.D. Russell ONT iNaturalist # 130794969 USA OP643427 – – Unpublished
86 Hydropus fuliginarius DAOM196062 USA – AF261368 – Moncalvo et al. (2002)
87 Hydropus marginellus AFTOL-ID 1720 not indicated DQ490627 DQ457674 DQ472722 Matheny et al. (2006)
88 Hydropus marginellus OSC 112834 USA EU669314 EU852808 – Unpublished
89 Hydropus rugosodiscus MGW1257 USA KY777386 – – Unpublished
90 Hydropus rugosodiscus PBM4022 USA KY777390 – – Unpublished
91 Hydropus rugosodiscus Taxon 10 not indicated MW399385 – – Unpublished
92 Leucoinocybe auricoma (≡Mycena auricoma) HKAS126433 China OQ025169 – – Direct Submission
93 Leucoinocybe auricoma (≡Mycena auricoma) AFTOL-ID 1341 (specimen_voucher HKAS41510) China DQ490647 – – Matheny et al. (2006)
94 Leucoinocybe danxiashanensis GDGM79543 China MZ667475 MZ667479 – Unpublished
95 Leucoinocybe danxiashanensis GDGM80113 China MZ667476 MZ667480 – Unpublished
96 Leucoinocybe danxiashanensis GDGM80114 China MZ667477 MZ667481 – Unpublished
97 Leucoinocybe danxiashanensis GDGM80184 China MZ667478 MZ667482 – Unpublished
98 Leucoinocybe flavoaurantia D Italy HM191743 – – Malysheva and Morozova (2011)
99 Leucoinocybe flavoaurantia GDOR Italy HM191744 – – Malysheva and Morozova (2011)
100 Leucoinocybe flavoaurantia LE 262757 Russia HM191745 – – Malysheva and Morozova (2011)
101 Leucoinocybe lenta BOZ (EPITYPE) Italy – LT854032 – Antonín et al. (2019)
102 Leucoinocybe lishuiensis FFAAS 0111 (HOLOTYPE) China MW424488 MW424492 – Na et al. (2021)
103 Leucoinocybe lishuiensis FFAAS 0112 China MW424489 MW424493 – Na et al. (2021)
104 Leucoinocybe lishuiensis FFAAS 0113 China MW424490 MW424494 – Na et al. (2021)
105 Leucoinocybe lishuiensis FFAAS 0115 China MW424491 MW424495 – Na et al. (2021)
106 Leucoinocybe sp. KA12-0435 South Korea KR673482 – – Kim et al. (2015)
107 Leucoinocybe subglobispora FFAAS1034 Holotype China OR238886 OR238898 OR258379 This study
108 Leucoinocybe subglobispora FFAAS1035 China OR238887 OR238899 OR258380 This study
109 Leucoinocybe sulcata CAL 1246 (HOLOTYPE) India KR029720 KR029721 – Latha et al. (2015)
110 Leucoinocybe taniae BCN-SCM B-4064 Italy LT854057 LT854028 – Antonín et al. (2019)
111 Marasmiellomycena albodescendens PDD 96142 New Zealand OL998341 OL998380 – Consiglio et al. (2022)
112 Marasmiellomycena albodescendens PDD 96321 New Zealand OL998343 OL998382 – Consiglio et al. (2022)
113 Marasmiellomycena omphaliiforme (≡Porotheleum omphaliiforme) WU 16775 Italy OM422777 OM423654 – Direct Submission
114 Marasmiellomycena omphaliiforme (≡Porotheleum omphaliiforme) LIP 0401689 France OM422780 OM423655 – Direct Submission
115 Marasmiellomycena omphaliiforme (≡Porotheleum omphaliiforme) AMB 18850 France OM422781 OM423656 – Direct Submission
116 Marasmiellomycena omphaliiforme (≡Porotheleum omphaliiforme) AMB 18845 France OM422782 – – Direct Submission
117 Marasmiellomycena pseudoomphaliiformis BRNM:552721 USA OR913562 OR913566 – Senanayake et al. (2023)
118 Marasmiellomycena pseudoomphaliiformis BRNM:552654 USA OR913560 OR913564 – Senanayake et al. (2023)
119 Marasmiellomycena pseudoomphaliiformis BRNM:552658 USA OR913561 OR913565 – Senanayake et al. (2023)
120 Marasmiellomycena tomentosa FFAAS1036 Holotype China OR238888 OR238900 OR258381 This study
121 Marasmiellomycena tomentosa FFAAS1037 China OR238889 OR238901 OR258382 This study
122 Marasmiellomycena tomentosa FFAAS1038 China OR238890 OR238902 OR258383 This study
123 Megacollybia clitocyboidea TFB11884/TENN60766 USA EU623658 – – Hughes et al. (2007)
124 Megacollybia clitocyboidea TENN62231 USA EU623664 – – Hughes et al. (2007)
125 Megacollybia clitocyboidea TENN62230 clone c4 USA EU623673 – – Hughes et al. (2007)
126 Megacollybia clitocyboidea TENN62230 clone c5 USA EU623674 – – Hughes et al. (2007)
127 Megacollybia fallax MICH 45002 USA EU623714 – – Hughes et al. (2007)
128 Megacollybia fallax TFB11561/TENN59447 USA EU623723 – – Hughes et al. (2007)
129 Megacollybia fallax DAOM208710 USA EU623724 – – Hughes et al. (2007)
130 Megacollybia fallax Mushroom Observer 291302 USA MN176984 – – Direct Submission
131 Megacollybia fallax Mushroom Observer 286893 USA MT437075 – – Direct Submission
132 Megacollybia marginata PRM 860926 Czech Republic LT854022 – – Antonín et al. (2019)
133 Megacollybia marginata PRM 859785 Czech Republic LT854046 LT854042 – Antonín et al. (2019)
134 Megacollybia marginata HR 91642 Czech Republic LT854050 – – Antonín et al. (2019)
135 Megacollybia marginata HR 91607 Czech Republic LT854051 – – Antonín et al. (2019)
136 Megacollybia platyphylla AFTOL-ID 560 USA DQ249275 AY635778 DQ385887 Unpublished
137 Megacollybia platyphylla BRNM 737654 Czech Republic LT854048 LT854036 – Antonín et al. (2019)
138 Megacollybia platyphylla BRNM 766972 Czech Republic LT854049 LT854037 – Antonín et al. (2019)
141 Megacollybia rodmani BHS2009-06 USA GQ397989 – – from GenBank
149 Megacollybia rodmani PUL F27039 USA MW448576 – – from GenBank
150 Megacollybia subfurfuracea TFB11075/TENN59558 clone c3 USA EU623744 – – Hughes et al. (2007)
151 Megacollybia subfurfuracea TFB11075/TENN59558 clone c8 USA EU623745 – – Hughes et al. (2007)
152 Megacollybia texensis DPL7405/TENN62058 clone c1 USA EU623725 – – Hughes et al. (2007)
153 Megacollybia texensis DPL7405/TENN62058 clone c2 USA EU623726 – – Hughes et al. (2007)
154 Megacollybia texensis FLAS-F-61511 USA MH211940 – – from GenBank
155 Mycena purpureofusca HMJAU 43554 China MG654740 MK629356 – Na and Bau (2018)
156 Mycena purpureofusca HMJAU 43624 China MG654741 MK629357 – Na and Bau (2018)
157 Mycena purpureofusca HMJAU 43640 China MG654742 MK629358 – Na and Bau (2018)
158 Porotheleum fimbriatum Dai 12276 China KX081137 KX161656 – from GenBank
159 Porotheleum fimbriatum Dai 12289 China KX081138 KX161654 – from GenBank
160 Porotheleum fimbriatum CLZhao 1120 China MH114870 – – from GenBank
161 Porotheleum fimbriatum CLZhao 2368 China MH114871 – – from GenBank
162 Porotheleum fimbriatum SWFC 006350 China MK894078 – – from GenBank
163 Porotheleum fimbriatum SWFC 006399 China MK894079 – – from GenBank
164 Porotheleum parvulum JBSD131802 Type Dominican Republic NR_182714 OM423657 – Consiglio et al. (2022)
165 Pseudohydropus floccipes AMB 18768 Spain – OM423637 – Consiglio et al. (2022)
166 Pseudohydropus floccipes BRNM 825631 Spain OM422760 OM423636 – Consiglio et al. (2022)
167 Pseudohydropus floccipes BRNM 751633 Spain OM422759 OM423635 – Consiglio et al. (2022)
168 Pseudohydropus globosporus BAP 661 (Holotype, SFSU) USA OM422758 OM423634 – Cooper et al. (2019)
169 Pseudohydropus sp MushroomObserver490861 Jamaica OR879917 – – Direct Submission
170 Pulverulina flavoalba FFAAS1039 Holotype China OR238891 OR238903 OR258384 This study
171 Pulverulina flavoalba FFAAS1040 China OR238892 OR238904 OR258385 This study
172 Pulverulina ulmicola TENN 029208 Holotype USA NR_119887 HQ179668 – Matheny et al. (2020)
173 Pulverulina ulmicola TFB13871 USA MT237476 MT237446 – Matheny et al. (2020)
174 Pulverulina ulmicola KUBOT-KRMK-2020-13 India MW425325 MW425344 – Unpublished
175 Trogia benghalensis CUH AM031 India KU647630 – – Dutta et al. (2017)
176 Trogia benghalensis CUH AM122 India MF967246 – – Dutta et al. (2017)
177 Trogia infundibuliformis KUN_HKAS63661 China JQ031775 JQ031780 – Yang et al. (2012)
178 Trogia infundibuliformis KUN_HKAS56709 China JQ031776 JQ031781 – Yang et al. (2012)
179 Trogia venenata KUN_HKAS54710 China JQ031772 JQ031778 – Yang et al. (2012)
180 Trogia venenata KUN_HKAS56679 China JQ031773 JQ031779 – Yang et al. (2012)
181 Trogia venenata TC2-28 China KT968080 – – Mi et al. (2016)
182 Trogia venenata MHHNU 8750 China KX268227 – – Unpublished
183 Vizzinia domingensis (≡Porotheleum domingense) JBSD131801a Dominican Republic OM422768 OM423646 – Consiglio et al. (2022)
184 Vizzinia nigripes (≡Porotheleum nigripes) JBSD131803 Dominican Republic OM422771 OM423648 – Consiglio et al. (2022)

Note: Newly-generated sequences are in bold.

Results

Phylogenetic analysis

A data matrix was created for 59 taxa, including 58 taxa of Porotheleaceae and, as an outgroup, one taxon of Mycena. Including gaps, the aligned dataset comprised 2,274 nucleotide sites: 974 for ITS, 610 for nrLSU and 690 for rpb2 exons (all sites without introns). For the ML analysis, the best-fit substitution models selected for ITS, nrLSU and rpb2-exon partitions in the concatenated dataset were TPM2uf+I+G4, GTR +I+G4 and TIM2+I+G4, respectively. For the BI analysis, the best-fit substitution model selected for each of the three DNA regions (ITS, nrLSU and rbp2 exons) was GTR+I+G. Phylogenetic reconstructions, based on BI and ML methods, yielded similar topologies. The BI topology was, therefore, selected as a representative phylogeny (Fig. 1).

Figure 1. 

Phylogenetic consensus tree inferred from the Maximum Likelihood (ML) analysis based on a concatenated ITS, nrLSU and rpb2 dataset (MLB ≥ 75%, BPP ≥ 0.90 are indicated). The tree is rooted with Mycena purpureofusca. The new species and combination are marked by red.

In the tree shown in Fig. 1, 21 major well-supported clades are evident: Chrysomycena, Clitocybula, Delicatula, Hydropodia, Hydropus, Leucoinocybe, Marasmiellomycena, Megacollybia, Pulverulina, Pseudohydropus, Trogia and Vizzinia, all of which form monophyletic groups at the generic level. However, within Porotheleum, two species, totalling three specimens, form two unassigned clades. In addition, Marasmiellomycena, forms a well-supported (MLB = 81%; BPP = 0.90) independent clade comprising four species distinct from Vizzinia and the unassigned Porotheleum. In the phylogenetic tree, Chrysomycena, Gerronema, Hydropus, Leucoinocybe, Megacollybia, Trogia and five taxa of Clitocybula cluster together with high statistical support (MLB = 96%; BPP = 1.00), but one sequence of Delicatula appears outside this large clade in the Maximum Likelihood analysis. The variation in the phylogenetic analysis outcomes for Delicatula specimens can be ascribed to inconsistent sequence coverage. Of the three Delicatula specimens evaluated, two only contained ITS sequences clustered together into a clade (MLB = 99%; BPP = 0.68), suggesting some degree of relatedness. In contrast, the remaining specimen, which only included an LSU sequence, was placed differently across the analyses. Such disparities in sequence coverage are likely to be responsible for the observed discrepancies between different computational algorithms used in the phylogenetic reconstructions. Hydropodia and Pulverulinaform a large, poorly supported clade. Moreover, Hydropus (MLB = 100%; BPP = 1.00), Leucoinocybe (MLB = 99%; BPP = 1.00) and Clitocybula (MLB = 82%; BPP = 1.00) are strongly supported as distinct genera and collectively constitute a distinct clade separate from all other clades. However, Gerronema is polyphyletic (Gerronema 1 to Gerronema 6), with each individual Gerronema clade sister to Megacollybia or Trogia. Finally, Chrysomycena and Hydropodia comprise a single species each.

In the phylogenetic tree, samples of the new species and new combination are placed in Marasmiellomycena, where they constitute monophyletic lineages, each with high statistical support (M. albodescendens: MLB = 93%, BPP = 1.00; M. tomentosa: MLB = 100%, BPP = 1.00). The four other new species are strongly supported as members of Gerronema 3, Pulverulina, Leucoinocybe and Clitocybula clades (C. fuscostriata: MLB = 92%, BPP = 1.00; G. brunneosquamulosum: MLB = 100%, BPP = 1.00; L. subglobispora: MLB = 100%, BPP = 1.00; and Pulverulina flavoalba: MLB = 100%, BPP = 1.00). Marasmiellomycena tomentosa is closely related to a clade containing two species and a new combination, M. albodescendens, M. omphaliiforme and M. pseudoomphaliiformis. Pulverulina flavoalba sp. nov. is grouped with high statistical support (MLB = 100%; BPP = 1.00) with three sequences of Pulverulina ulmicola (H.E. Bigelow) Matheny & K.W. Hughes from India and the USA (including holotype voucher no. TENN 029208). Within the Leucoinocybe clade, L. subglobispora constitutes a monophyletic lineage that is most closely related to Leucoinocybe lishuiensis, a new species recently described from China (Na et al. 2021). Clitocybula fuscostriata is placed along with C. lacerata (Scop.) Métrod in an unresolved lineage that is treated as C. lacerata agg. by Antonín et al. (2019) and in our studies.

Clitocybula albida A.K. Dutta, K. Acharya & Antonín, reported from India as a new species, was transferred to Porotheleum [as Porotheleum albidum (A.K. Dutta, K. Acharya & Antonín) Vizzini & Consiglio] and Porotheleum parvulum Angelini, Vizzini, Consiglio & M. Marchetti as a new species from the Dominican Republic (Dutta et al. 2018; Consiglio et al. 2022). The phylogenetic status of Clitocybula albida is currently unclear and treated as unassigned clades in the study of Senanayake et al. (2023). On the other hand, Porotheleum parvulum is known to cluster with Marasmiellomycena and Vizzinia, forming a clade. Within this clade, Porotheleum parvulum is specifically determined to be a sister group to Marasmiellomycena. In the research conducted by Senanayake et al. (2023), Pseudohydropus and Pulverulina were identified as forming a monophyletic group. Contrastingly, in our phylogenetic tree, Pseudohydropus emerges as an independent lineage, receiving robust support (MLB =100%; BPP = 1.00) and not aligning as a sister group with any other genera. The observed differences might stem from variances in sequence coverage and the evolutionary rates of the genes. While Senanayake et al. (2023) utilised ITS and LSU sequences for their phylogenetic construction, our study encompassed ITS, LSU and RPB2 in the combined phylogenetic analysis. (Fig. 1).

Taxonomy

Clitocybula fuscostriata Q.Na & Y.P.Ge, sp. nov.

MycoBank No: 849407
Figs 2, 3, 4

Diagnosis

Pileus with dark-brown striae. Differs from C. striata in having broader basidiospores and lacking hymenial cystidia.

Figure 2. 

Basidiomata of Clitocybula fuscostriata A–D collection FFAAS1029 E–F collection FFAAS1030, holotype G–H collection FFAAS1031. Scale bars: 10 mm (A–H).

Holotype

China. Zhejiang Province: Baiyun National Forest Park, Liandu District, Lishui City, 2 Aug 2021, Qin Na, Yupeng Ge, Zewei Liu, Yaping Hu, Changjing Liu and Hui Ding, FFAAS1030 (collection number MY0460).

Figure 3. 

Morphological features of Clitocybula fuscostriata (FFAAS1030, holotype) A basidiomata B basidiospores C basidia D caulocystidia E pileipellis and pileocystidia. Scale bars: 10 mm (A); 5 μm (B); 10 μm (C–E).

Etymology

Name refers to the pileus with radially fuscous striae.

Figure 4. 

Microscopic features of Clitocybula fuscostriata (FFAAS1030, holotype) A–E basidiospores F basidia G margin of lamellae H lamellar trama I pileipellis and pileocystidia J caulocystidia. Scale bars: 5 μm (A–E); 10 μm (F–J). Structures were stained with 1% Congo Red aqueous solution before photographing.

Description

Pileus 3.0–28.5 mm in diameter, hemispherical at first, then convex with depressed centre, expanded with age, infundibuliform with deeply umbilicate at the centre when old, thin-fleshed, dry, surface innately radially Fuscous (XLVI13′′′′k) to Fuscous-Black (XLVI13′′′′m) striate, surface somewhat fibrillose, becoming glabrous, radially cracked at margin when old, Benzo Brown (XLVI13′′′′i), Hair Brown (XLVI17′′′′i), Fuscous (XLVI13′′′′k) to Fuscous-Black (XLVI13′′′′m) at the centre, Pale Smoke Grey (XLVI21′′′′f) in the margin when young, Pale Smoke Grey (XLVI21′′′′f) to Smoke Grey (XLVI21′′′′d) with Bone Brown (XL13′′′m) at the centre when old. Context thin, white, fragile. Lamellae subdecurrent, white, with 1–3 tiers of lamellulae, irregularly intervenose, edges concolorous with the face. Stipe 17.0–52.0 × 1.0–2.5 mm, hollow, cylindrical, strongly and coarsely grooved, slightly bulbous at the base, fragile, finely whitish fibrillose, white in the upper part, Citrine Drab (XL21′′′i) in the base, base covered with a few white fibrils. Odour and taste inconspicuous.

Basidiospores (80/4/3) (5.2) 5.4–5.8–6.2 (6.5) × (4.2) 4.3–4.7–5.0 (5.1) μm [Q = 1.13–1.34, Q = 1.25 ± 0.050] [holotype (40/2/1) (5.3) 5.5–5.8–6.2 (6.5) × (4.2) 4.4–4.6–5.0 (5.1) μm, Q = 1.17–1.32, Q = 1.26 ± 0.040], broadly ellipsoid, hyaline in 5% KOH, smooth, thin-walled, guttulate, amyloid. Basidia 22–32 × 5–9 μm, 2- or 4-spored, clavate, sterigmata 2.5–4.7 × 0.6–1.6 μm. Hymenial cystidia absent. Lamellae edge cells scattered, cylindrical, narrowly clavate, thin-walled. Lamellar trama subregular; hyphae 3–7 μm wide, thin-walled, hyaline, non-dextrinoid. Pileipellis hyphae 4–9 μm wide, smooth; pileocystidia 70–162 × 7–19 μm, cylindrical or narrowly clavate, apically obtuse, thin-walled, hyaline, smooth. Stipitipellis a cutis made up of 3–8 μm wide hyphae, smooth, thin-walled; caulocystidia 27–63 × 5–8 μm, cylindrical, clavate, fusoid, apically obtuse, thin-walled base, smooth, transparent. Clamps present in all tissues.

Habit and habitat

Scattered on rotten branches or twigs in Acer, Armeniaca, Cercidiphyllum, Emmenopterys and Picea mixed forests.

Known distribution

Zhejiang Province, China.

Additional material examined

China. Zhejiang Province: Baiyun National Forest Park, Liandu District, Lishui City, 2 Aug 2021, Qin Na, Yupeng Ge, Hui Zeng and Yulan Sun, FFAAS1029 (collection number MY0459); Zhejiang Province: Baiyun National Forest Park, Liandu District, Lishui City, 2 Aug 2021, Qin Na, Yupeng Ge, Zewei Liu, Yaping Hu, Changjing Liu and Hui Ding, FFAAS1031 (collection number MY0466).

Notes

Clitocybula fuscostriata is considered to be a distinct species in the genus on account of its pileus with dark-brown striae, broadly ellipsoid basidiospores, absence of cheilocystidia and pleurocystidia and thin-walled pileipellis and stipitipellis hyphae. Five recorded species morphologically resemble this new species: C. familia (Peck) Singer, C. lacerata (Scop.) Métrod, C. oculata (Murrill) H.E. Bigelow, C. striata Dähncke, Contu & Vizzini and C. tilieti (Singer) Singer (Singer 1943; Romagnesi 1968; Bigelow 1973; Lennox 1979; Ludwig 2000, 2001; Dähncke et al. 2010; Antonín et al. 2011). Clitocybula striata, a new taxon reported from Spain, has certain morphological similarities to C. fuscostriata, namely, a grey-brown to brown pileus with dark-brown striae, but differs from C. striata in having ellipsoid basidiospores (5–7 × 3.5–4.8 μm; Q = 1.5) and presence of utriform or lageniform cheilocystidia (Dähncke et al. 2010). In contrast to C. fuscostriata, C. tilieti can be easily mistaken for C. striata, but the pileus of C. tilieti is distinctly viscid and its stipitipellis and caulocystidia are thick-walled (Singer 1943; Antonín et al. 2011). Clitocybula lacerata (Scop.) Métrod, the type species of Clitocybula, is characterised by its caespitose stipes, beige-grey to pale-grey brown pileus, presence of clavate cheilocystidia and a pileipellis with pale encrusting pigmentation, differentiating it from C. fuscostriata (Peck 1878; Breitenbach and Kranzlin 1991; Ludwig 2000, 2001; Antonín et al. 2019). Clitocybula oculata (Murrill) H.E. Bigelow and C. familia resemble C. fuscostriata in colour and size of the pileus and stipe, but can be distinguished from the new species by the size and shape of the basidiospores [C. oculata basidiospores (8.5–)10–12(–13) × 6–9 μm, broadly ellipsoid or ovate; C. familia basidiospores 3.5–5.3(–5.5) × 3.5–5.0 μm, globose, subglobose to broadly ellipsoid] (Romagnesi 1968; Bigelow 1973; Lennox 1979; Ludwig 2000, 2001; Antonín et al. 2011).

Gerronema brunneosquamulosum Q.Na & Y.P.Ge, sp. nov.

MycoBank No: 849408
Figs 5, 6, 7

Diagnosis

Differs from G. zhujian in having a fuscous stipe densely covered with deep-brown pubescence or scales and by the presence of large basidiospores.

Figure 5. 

Basidiomata of Gerronema brunneosquamulosum A, B FFAAS1032, holotype C, D collection FFAAS1033 E, F pileus with granules, fur or scales G, H stipe covered with dark brown scales. Scale bars: 10 mm (A–E); 5 mm (F–H).

Holotype

China. Zhejiang Province: Baiyun National Forest Park, Liandu District, Lishui City, 2 Aug 2021, Qin Na, Yupeng Ge, and Hui Zeng, FFAAS1032 (collection number MY0481).

Figure 6. 

Morphological features of Gerronema brunneosquamulosum (FFAAS1032, holotype) A basidiomata B basidia C basidiospores D cheilocystidia E caulocystidia F pileipellis. Scale bars: 10 mm (A); 10 μm (B–F).

Etymology

Name refers to the pileus and stipe covered with dark-brown scales.

Figure 7. 

Microscopic features of Gerronema brunneosquamulosum (FFAAS1032, holotype) A–E basidiospores F basidia G–J cheilocystidia K lamellar trama L pileipellis and pileocystidia M caulocystidia. Scale bars: 5 μm (A–E); 10 μm (structures A–K, M were stained with 1% Congo Red aqueous solution and L in 5% KOH aqueous solution before photographing).

Description

Pileus 4.5–42.0 mm in diam., applanate and centrally depressed, subumbilicate to umbilicate when young, concave to deeply infundibulate with age, pellucid-striate or sulcate, always ± distinctly radially striped with darkened lines, Buffy Brown (XL17′′′k) at the centre, Olive Buff (XL21′′′d) in margin when young, Olive Brown (XL17′′′k), Clove Brown (XL17′′′m), Light Greyish-Olive (XLVI21′′′′b) in margin with age, densely covered with tiny, Warm Blackish-Brown (XXXIX1′′′m) granules, pubescence or scales, slightly sparse with age, dry, lustreless, with a slightly involuted margin. Context white, thin, tough. Lamellae narrowly adnexed to subdecurrent, moderately broad, pure white, edges concolorous with the sides. Stipe 6.0–32.0 × 1.5–2.0 mm, central, cylindrical, almost equal above, white, densely covered with Warm Blackish-Brown (XXXIX1′′′m) scales, hollow, base Light Seal Brown (XXXIX9′′′m), slightly swollen with tiny, inconspicuous fine white hairs. Odourless. Taste mild.

Basidiospores [60/3/2] (9.0) 9.2–10.0–11.2 (12.9) × (4.9) 5.2–5.8–6.6 (7.2) μm [Q = 1.54–1.91, Q = 1.73 ± 0.097] [holotype [40/2/1] (9.0) 9.2–10.2–11.2 (12.9) × (5.3) 5.5–5.9–6.5 (7.2) μm, Q = 1.54–1.90, Q = 1.71 ± 0.086], ellipsoid to narrowly ellipsoid, hyaline, guttulate, thin-walled, inamyloid. Basidia 22–39 × 7–9 μm, hyaline, clavate, 2- or 4-spored, sterigmata 2.3–6.0 × 0.8–2.2 μm. Cheilocystidia 23–59 × 6–9 μm, subfusiform, clavate, apex usually swollen, hyaline. Pleurocystidia absent. Lamellar trama subregular; hyphae 2–7 μm wide, thin-walled, hyaline, inamyloid. Pileus trama subregular, sarcodimitic. Pileipellis hyphae 3–7 μm wide, a cutis, light yellow (2B2); terminal elements clavate or utriform with rounded apex, 53–95 × 7–16 μm, Dark Citrine (IV21m), Olive Brown (XL17′′′k) to Clove Brown (XL17′′′k) pigmented; true pileocystidia absent. Hyphae of the stipitipellis 5–11 μm wide, hyaline, smooth; caulocystidia long cylindrical, sometimes with rounded apex, 40–76 × 5–12 μm, hyaline, thin-walled. All tissues non-reactive in iodine. Clamps present in all tissues.

Habit and habitat

Solitary to scattered on rotten wood, branches and twigs in Acer, Ginkgo, Liriodendron, Picea and Tsuga.

Known distribution

Fujian Province, Zhejiang Province, China.

Additional material examined

China. Fujian Province: Wuyi Mountain, Nanping City, 13 Aug 2021, Qin Na, Yupeng Ge, Junqing Yan, Hui Zeng, and Zewei Liu, FFAAS1033 (collection number MY0571).

Notes

Gerronema brunneosquamulosum is unique amongst members of Gerronema on account of its fuscous pileus and stipe with dark-brown to blackish-brown pubescence or scales, larger basidiospores and a dark-pigmented pileipellis. Gerronema zhujian, reported from Anhui and Fujian Provinces in China, is the most closely allied congener of G. zhujian on the basis of the brown colouration of the umbilicus of its pileus, its whitish stipe and similarly-shaped cheilocystidia and terminal elements of the pileipellis (Na et al. 2022a). This taxon differs from G. brunneosquamulosum in having a pruinose white stipe, subdecurrent to decurrent lamellae and possessing smaller basidiospores (Na et al. 2022a). Two species of Omphalina Quél., characterised by dark pigments in the pileus, have been described from Argentina–Omphalina depauperata (Singer) Raithelh. and O. subpallida (Singer) Raithelh., formerly named Gerronema subpallidum Singer and G. depauperatum Singer, respectively. These two species most closely resemble G. brunneosquamulosum, but differ in having an unornamented stipe, ellipsoid basidiospores and no cheilocystidia (Singer 1970). Other species of Gerronema, such as G. nemorale and G. strombodes, are well characterised with a distinctly yellow, yellowish-orange, olive-yellow to yellowish-brown pileus and their micromorphological features are also different (Singer 1970; Antonín et al. 2008; Latha et al. 2018). Species of Trogia, especially Trogia fulvochracea Corner (p.31) and Trogia mycenoides (p.53) Corner, share some similarities with the new taxon (Corner 1991). Trogia fulvochracea, however, has a fulvous or cinnamon-ochraceous pileus, a smooth white stipe and smaller basidiospores (7–9.5 × 4.5–6.0 μm). Trogia mycenoides differs in having a smaller pileus (5–30 mm in diam.), ellipsoid basidiospores and clavate to subglobose cheilocystidia; in addition, true pileocystidia are present, but are soon evanescent (Corner 1991).

Leucoinocybe subglobispora Q.Na & Y.P.Ge, sp. nov.

MycoBank No: 849409
Figs 8, 9, 10

Diagnosis

Pileus dark brown. Basidiospores subglobose to broadly ellipsoid. Pileocystidia and caulocystidia thick-walled. Differs from L. lishuiensis in having broader basidiospores.

Figure 8. 

Basidiomata of Leucoinocybe subglobispora A–E collection FFAAS1034, holotype F–G collection FFAAS1035. Scale bars: 10 mm (A–G).

Holotype

China. Zhejiang Province: Tianmu Mountain, Hangzhou City, 1 Aug 2021, Qin Na, Yupeng Ge, Zewei Liu and Yulan Sun, FFAAS1034 (collection number MY0444).

Figure 9. 

Morphological features of Leucoinocybe subglobispora (FFAAS1034, holotype) A basidiomata B basidiospores C cheilocystidia D basidia E caulocystidia F pileocystidia. Scale bars: 5 mm (A); 10 μm (B–F).

Etymology

Name refers to the subglobose to broadly ellipsoid basidiospores.

Figure 10. 

Microscopic features of Leucoinocybe subglobispora (FFAAS1034, holotype) A–E basidiospores F basidia G–J cheilocystidia K lamellar trama L pileipellis and pileocystidia M caulocystidia. Scale bars: 5 μm (A–E); 10 μm (F–M). Structures were stained with 1% Congo Red aqueous solution before photographing.

Description

Pileus 2.5–8.0 mm in diameter, hemispherical or campanulate when young, becoming campanulate with age, umbilicate at the centre, sulcate, finely granulose all over, Dark Livid Brown (XXXIX1′′′k), Benzo Brown (XLVI13′′′′i) to Fuscous (XLVI13′′′′k) at the centre, Pale Smoke Grey (XLVI21′′′′f) in the margin, uplifted or recurved at the margin and sometimes rimose in age, dry. Context white, thin, fragile. Lamellae adnexed to slightly subdecurrent, white, with 1–2 tiers of lamellulae, edges concolorous with the face. Stipe 9.5–14.0 × 1.0–1.5 mm, equal or slightly broadened at the base, hollow, fragile, white, sometimes inconspicuous Pale Olive-Buff (XL21′′′d) at the base, densely pruinose, but sparsely with age, base covered with small white fibrils. Odour and taste indistinctive.

Basidiospores (60/3/2) (5.6) 5.8–6.4–7.1 (7.5) × (4.8) 5.0–5.6–6.5 (6.8) μm [Q = 1.06–1.27, Q = 1.16 ± 0.054] [holotype (40/2/1) (5.7) 5.9–6.5–7.2 (7.5) × (4.9) 5.0–5.5–6.5 (6.8) μm, Q = 1.07–1.27, Q = 1.18 ± 0.052], subglobose to broadly ellipsoid, hyaline in 5% KOH, smooth, thin-walled, guttulate, amyloid. Basidia 28–37 × 7–9 μm, 4-spored, clavate, sterigmata 1.4–2.7 × 0.8–1.7 μm. Cheilocystidia 28–62 × 9–15 μm, distinct, flexuose, narrowly utriform, fusoid or lageniform, subcapitate, thin-walled, hyaline. Pleurocystidia absent. Lamellae trama subregular; hyphae 2–6 μm wide, thin-walled, hyaline, amyloid. Pileipellis hyphae 2–8 μm wide, smooth; pileocystidia 62–116 × 10–19 μm, lageniform, subulate, apically obtuse, distinctly 0.8–1.8 μm thick-walled, with a thin-walled base, hyaline, smooth. Stipitipellis a cutis made up of 3–9 μm wide hyphae, smooth, thin-walled; caulocystidia 34–62 × 5–10 μm, subulate, fusoid, lageniform, sometimes clavate, always thick-walled in the middle part and with a thin-walled base, smooth, transparent. Clamps present in all tissues.

Habit and habitat

Solitary or scattered on rotten wood or branches in Acer, Armeniaca, Cercidiphyllum, Emmenopterys and Picea mixed forests.

Known distribution

Zhejiang Province, China.

Additional material examined

China. Zhejiang Province: Baiyun National Forest Park, Liandu District, Lishui City, 2 Aug 2021, Qin Na, Yupeng Ge, Zewei Liu, Yaping Hu and Hui Ding, FFAAS1035 (collection number MY0475).

Notes

Leucoinocybe subglobispora is considered to be a distinct species of Leucoinocybe on account of its subdecurrent lamellae, subglobose to broadly ellipsoid basidiospores, thick-walled pileocystidia and caulocystidia and saprophytic habitat. Leucoinocybe lenta, the type species of Leucoinocybe, also has a white stipe and lamellae, similarly-shaped cheilocystidia and thick-walled pileocystidia, but differs from the new species by the presence of a reddish-brown pileus with pinkish shades or pale pinkish-beige at the centre that fades to white towards the margin, larger basidiomata and ellipsoid basidiospores [(5.3)6.0–7.3(7.9) × (3.8)4.0–4.5(5.1) μm] (Gröger 2006; Eyssartier and Roux 2011; Antonín et al. 2019; Kaygusuz et al. 2020). Leucoinocybe taniae (= Clitocybula flavoaurantia) resembles L. subglobispora in having a brown pileus, white and decurrent lamellae and a white stipe with a brownish base, but differs in possessing the following features: a reddish-yellow pileus when old, larger and broadly amygdaliform spores (6.2–7.8 × 4.8–7.0 μm) and thin-walled pileocystidia and caulocystidia (Vila 2002; Contu 2003; Malysheva and Morozova 2011; Antonín et al. 2019). Leucoinocybe sulcata, recently described as a new taxon from India, is easily distinguished from the new species by the presence of greyish-orange to brown basidiomata, a larger pileus (13–52 mm in diam.), broadly ellipsoid to subamygdaliform basidiospores (5.0–6.5 × 4.0–5.5 μm; Q = 1.1–1.5) and thin-walled caulocystidia and the absence of pileocystidia (Latha et al. 2015). Leucoinocybe lishuiensis, reported as a new species from south-eastern China in our previous study, can be easily mistaken for L. subglobispora on account of having an identical habit and habitat, a small, pure-brown pileus, slightly decurrent lamellae, similarly-shaped cheilocystidia and thick-walled pileocystidia and caulocystidia; however, the narrowly ellipsoid basidiospores and smaller pileocystidia of L. lishuiensis can be used to distinguish this species from L. subglobispora (Na et al. 2021). Another new combination of Leucoinocybe, L. auricoma (Har. Takah.) Matheny, originally named Mycena auricoma Har. Takah., is also comparable to the present species in having thick-walled pileocystidia and caulocystidia; however, L. auricoma has a yellowish-orange flocculent pileus and stipe, ovoid-ellipsoid to ellipsoid basidiospores (5–7 × 3–4 μm) and pileocystidia and caulocystidia with yellow contents (Takahashi 1999; Matheny et al. 2020).

Marasmiellomycena albodescendens (J.A. Cooper) Q.Na & Y.P.Ge, comb. nov.

MycoBank No: 851718

Basionym

Porotheleum albodescendens J.A. Cooper, in Consiglio, Vizzini, Cooper, Marchetti, Angelini, Brugaletta & Setti, Riv. Micol. 64(2): 117, 2022.

Type specimen

Holotype : New Zealand: North Island, Taupo, Tauhara Centre, 15 May 2011, PDD 96321.

Selected description

Consiglio et al. (2022).

Distribution

New Zealand.

Notes

Marasmiellomycena albodescendens has marasmielloid basidiomes, a pure-white pileus, relatively large spores, no hymenial cystidia and abundant, thick-walled pileocystidia and caulocystidia with yellowish contents. Unlike other species of Marasmiellomycena possessing a yellow, reddish-brown or yellowish-brown pileus, M. albodescendens can be easily recognised by its white pileus. The pileus of Marasmiellomycena albodescendens is macromorphologically more similar to some species of Marasmiellus Murrill (Stevenson 1964); however, its micromorphological characteristics place this species in Marasmiellomycena, consistent with the results of our phylogenetic analysis (Fig. 1). Marasmiellomycena albodescendens has been infrequently collected in New Zealand, but is probably common and widespread and grows on small, dead, fallen branches and twigs in indigenous scrub and broad-leaf forests in late summer and autumn (Consiglio et al. 2022).

Marasmiellomycena tomentosa Q.Na & Y.P.Ge, sp. nov.

MycoBank No: 851717
Figs 11, 12, 13, 14

Diagnosis

Pileus and stipe distinctly tomentose. Pileus dark brown, subsquamulose. Basidiospores narrowly ellipsoid, slightly amyloid. Hymenial cystidia absent. Pileipellis and stipitipellis sarcodimitic, hyphae thick-walled with yellowish-brown pigments. Pileocystidia and caulocystidia thick-walled with yellow contents. Differs from M. pseudoomphaliiformis by possessing a distinctly tomentose, dark-brown subsquamulose pileus, narrowly ellipsoid basidiospores and absence of cheilocystidia.

Figure 11. 

Basidiomata of Marasmiellomycena tomentosa A–D collection FFAAS1036, holotype E, F collection FFAAS1037 G, H collection FFAAS1038. Scale bars: 10 mm (A–H).

Holotype

China. Zhejiang Province: Tianmu Mountain, Hangzhou City, 30 Jul 2021, Qin Na, Zewei Liu, Yulan Sun and Yupeng Ge, FFAAS1036 (collection number MY0421).

Figure 12. 

Morphological features of Marasmiellomycena tomentosa (FFAAS1036, holotype) A basidiomata B basidiospores C basidia D pileipellis and pileocystidia. Scale bars: 10 mm (A); 10 μm (B–D).

Etymology

Name refers to the tomentose to subsquamulose pileus.

Figure 13. 

Morphological features of Marasmiellomycena tomentosa (FFAAS1036, holotype) A stipitipellis and caulocystidia. Scale bars: 10 μm (A).

Description

Pileus 0.5–18.5 mm in diameter, at first convex or campanulate, soon expanding to plano-convex, always depressed to umbilicate at the centre, surface dry, densely covered with minute white (LIII) pubescence, tomentose all over, subsquamulose, ground colour Verona Brown (XXIX13′′k) to Warm Sepia (XXIX13′′m), Mustard Yellow (XVI19′b), Old Gold (XVI19′i) to Buffy Citrine (XVI19′k), Saccardo’s Olive (XVI19′m) at the centre, fading to Wax Yellow (XLVI21′′′′f) when old, margin slightly sulcate, uplifted or recurved in age. Context thin, Primrose Yellow (XXX23′′d). Lamellae decurrent to subdecurrent, Wax Yellow (XLVI21′′′′f), Mustard Yellow (XVI19′b), with 1–2 tiers of lamellulae, edges concolorous with the face, slightly fimbriate edge. Stipe 7.5–21.0 × 1.0–1.6 mm, central, terete, curved, equal or slightly broadened at the base, hollow or stuffed, dry, Mustard Yellow (XVI19′b) in the upper part, Saccardo’s Olive (XVI19′m), Benzo Brown (XLVI13′′′′i), Fuscous (XLVI13′′′′m), Deep Greyish-Olive (XLVI21′′′′b) towards the base, densely and minutely silky-fibrillose and white (LIII) pruinose-floccose to tomentose throughout, base covered with white mycelium. Odour indistinct to fungoid, taste mild.

Figure 14. 

Microscopic features of Marasmiellomycena tomentosa (FFAAS1036, holotype) A–E basidiospores F basidia G–J lamellae margin. Scale bars: 5 μm (A–E); 10 μm (F–J). Structures were stained in 5% KOH aqueous solution before photographing.

Basidiospores (80/4/3) (6.8) 7.2–7.6–8.2 (8.4) × (3.7) 3.9–4.1–4.5 (4.6) μm [Q = 1.75–1.98, Q = 1.83 ± 0.052] [holotype (40/2/1) (6.8) 7.2–7.7–8.4 × 3.9–4.2–4.6 μm, Q = 1.75–1.98, Q = 1.82 ± 0.050], narrowly ellipsoid, hyaline in 5% KOH, smooth, thin-walled, multiguttulate, slightly amyloid. Basidia 20–35 × 5–8 μm, 2- or 4-spored, clavate, sterigmata 2.2–4.8 × 0.6–1.6 μm. Hymenial cystidia absent. Lamellar trama subregular; hyphae 3–10 μm wide, with 0.5–1.0 µm thick-walled, light yellow, dextrinoid. Pileipellis hyphae 3–8 μm wide, sarcodimitic, cutis, smooth, 0.4–1.0 μm thick-walled, with intracellular yellowish-brown pigment; pileocystidia 38–223 × 5–12 μm, in clusters, narrowly subulate or narrowly lageniform to fusiform with very long and tapering neck, distinctly 0.6–1.5 μm thick-walled, yellow, smooth. Stipitipellis made up of cylindrical, 4–9 µm wide hyphae, sarcodimitic, smooth, 0.5–1.0 μm thick-walled, with intracellular brownish-orange pigment; caulocystidia 45–327 × 5–9 μm, similar to the pileocystidia, but usually longer, 0.5–1.3 μm thick-walled, smooth, with intracellular yellowish pigment. Clamps present in all tissues.

Habit and habitat

Solitary or scattered on rotten branches, twigs and wood debris in Acer, Armeniaca, Cercidiphyllum, Emmenopterys and Picea mixed forests.

Known distribution

Zhejiang Province, China.

Additional material examined

China. Zhejiang Province: Tianmu Mountain, Hangzhou City, 30 Jul 2021, Qin Na, Zewei Liu, Yulan Sun and Yupeng Ge, FFAAS1037 (collection number MY0422); Zhejiang Province: Tianmu Mountain, Hangzhou City, 1 Aug 2021, Qin Na, Zewei Liu, Yulan Sun and Yupeng Ge, FFAAS1038 (collection number MY0443).

Notes

Marasmiellomycena tomentosa is a rare thermophilous species reported from south-eastern areas of China from July to August on rotten branches, twigs and woody debris of deciduous and coniferous trees (Acer, Armeniaca, Cercidiphyllum, Emmenopterys and Picea). The most distinctive characteristics of this species are a tomentose, brown subsquamulose pileus, a tomentose stipe, narrowly ellipsoid and slightly amyloid basidiospores, the absence of hymenial cystidia and thick-walled pileipellis, stipitipellis, pileocystidia and caulocystidia with yellow or brownish-orange contents. Species morphologically most closely allied to Marasmiellomycena tomentosa include M. omphaliiforme, M. pseudoomphaliiformis and M. albodescendens. Marasmiellomycena pseudoomphaliiformis resembles M. tomentosa by the presence of a pale beige to brown pileus with finely tomentose to pubescent pileus, but differs in having white to cream-white or beige lamellae rather than yellow, ellipsoid to ellipsoid-fusiform basidiospores [(6.5–)7.0–9.0(–9.5) × 4.0–5.5 µm] and clavate, fusiform to lageniform cheilocystidia (Senanayake et al. 2023). Marasmiellomycena omphaliiforme is considered to be a closely-related taxon with evident affinities to M. tomentosa–not only regarding its phylogenetic placement, but also in terms of morphological features (Kühner and Romagnesi 1954; Antonín and Noordeloos 1993, 1997; Consiglio et al. 2022; Senanayake et al. 2023). The two species resemble one another in having a similarly-coloured pileus and stipe, similarly-shaped basidiospores, pileocystidia and caulocystidia and a yellowish-pigmented pileipellis and stipitipellis; however, the minutely pubescent, granulose to subsquamulose pileus, as well as the relative abundance of cheilocystidia, appear to be variable characters in M. omphaliiforme in contrast to the new species (Kühner and Romagnesi 1954; Antonín and Noordeloos 1993, 1997; Consiglio et al. 2022). According to the description of Consiglio et al. (2022), Marasmiellomycena albodescendens from New Zealand has a pure-white pileus, a thin-walled pileipellis and larger basidiospores (9.6 ± 0.7 µm × 5.2 ± 0.4 µm).

Pulverulina flavoalba Q.Na & Y.P.Ge, sp. nov.

MycoBank No: 849410
Figs 15, 16, 17

Diagnosis

Pileus white to light orange yellow. Basidiospores cylindrical. Hymenial cystidia absent. Lamellar trama, pileipellis and stipitipellis hyphae thin-walled. Differs from Pu. ulmicola in having larger and longer basidiospores and possessing thin-walled lamellar trama, pileipellis and stipitipellis hyphae.

Figure 15. 

Basidiomata of Pulverulina flavoalba A–D collection FFAAS1039, holotype E–H collection FFAAS1040. Scale bars: 5 mm (A–H).

Holotype

China. Guangxi Zhuang Autonomous Region: Liangfengjiang National Forest Park, Nanning City, 13 Jul 2022, Yupeng Ge and Renxiu Wei, FFAAS1039 (collection number MY0863).

Figure 16. 

Morphological features of Pulverulina flavoalba (FFAAS1039, holotype) A basidiomata B basidia C basidiospores D caulocystidia E pileipellis. Scale bars: 2 mm (A); 10 μm (B, D, E); 5 μm (C).

Etymology

Name refers to the white to light-yellow pileus and stipe.

Figure 17. 

Microscopic features of Pulverulina flavoalba (FFAAS1039, holotype) A–E basidiospores F basidia G lamellae margin H lamellar trama I pileipellis J caulocystidia. Scale bars: 5 μm (A–E); 10 μm (F–J). Structures A–F were stained in 5% KOH aqueous solution and G–J with 1% Congo Red aqueous solution before photographing.

Description

Pileus 1.2–5.8 mm in diameter, arched or plano-convex with a slight depression at the centre when young, becoming more depressed with age; translucent striate, floccose or granulose, glabrescent when old, surface dull, dry; white (LIII) when young, aniline yellow (IV19i) or light orange-yellow (III17d) at the centre and in the margin with age, margin decurved. Context white, thin, not fragile. Lamellae decurrent, white, orange citrine (IV19k) tinged when old, with 1–2 tiers of lamellulae, edges even, medium-broad. Stipe 1.6–14.4 × 0.5–1.0 mm, terete or slightly broadened at the base, curved, dry, white, with a pruinose, pubescent or fibrillose surface, sparser with age, hollow, not fragile, white, sometimes aniline yellow (IV19i), light orange-yellow (III17d) in the middle and at the base; base covered with white mycelium. Odour absent, taste mild.

Basidiospores (60/3/2) (6.8) 7.0–7.9–8.8 (9.1) × (3.3) 3.7–4.1–4.4 (4.7) μm [Q = 1.81–2.19, Q = 1.93 ± 0.099] [holotype (40/2/1) (6.8) 7.0–7.8–8.9 (9.1) × (3.3) 3.7–4.1–4.4 (4.7) μm, Q = 1.77–2.19, Q = 1.92 ± 0.084], cylindrical, hyaline in 5% KOH, smooth, thin-walled, guttulate, inamyloid, with a small, but discernible apiculus. Basidia 21–30 × 4–6 μm, 2- or 4-spored, clavate, sterigmata 1.9–5.6 × 0.6–1.6 μm. Hymenial cystidia absent. Lamellar trama subregular to interwoven; hyphae 5–15 µm wide, hyaline, thin-walled. Pileipellis a cutis of cylindrical hyphae 3–7 µm wide, smooth; end cells often protruding, 35–105 × 3–12 μm, cylindrical, subfusiform, apically obtuse, thin-walled, hyaline, smooth. Stipitipellis hyphae 3–8 μm wide, smooth, thin-walled; caulocystidia 19–50 × 4–9 μm, clavate, subfusiform, thin-walled, smooth, transparent. Clamps present in all tissues.

Habit and habitat

Scattered to gregarious on rotten wood, branches or fruits in mixed forests of Acacia, Ficus, Ilex, Parashorea, Picea and Trachycarpus etc.

Known distribution

Guangxi Zhuang Autonomous Region, China.

Additional material examined

China. Guangxi Zhuang Autonomous Region: Liangfengjiang National Forest Park, Nanning City, 13 Jul 2022, Yupeng Ge and Renxiu Wei, FFAAS1040 (collection number MY0865).

Notes

Clitocybe ulmicola H.E. Bigelow was established by Bigelow in 1982 and published as a new combination, Pulverulina ulmicola (H.E. Bigelow) Matheny & K.W. Hughes (Matheny et al. 2020). The description of Pulverulina ulmicola modified from Bigelow (1982) includes observations based on recent American material (Matheny et al. 2020). As far as we know, only Pulverulina ulmicola has previously been included in the genus and has had morphological features described in detail (Bigelow 1982; Matheny et al. 2020). In appearance, Pulverulina ulmicola is a small, whitish, marasmioid fungus, with small basidiomata, distant decurrent lamellae, a tough texture, interwoven gill trama, long cylindrical caulocystidia and short, ellipsoid, smooth basidiospores and occurs on the bark of living Ulmus and Quercus trees. Our collections of Pulverulina flavoalba from the Guangxi Zhuang Autonomous Region represent a taxon that is distinct from Pulverulina ulmicola, as compared to the macroscopic and microscopic characters described by Matheny et al. (2020). Pulverulina ulmicola differs from P. flavoalba in having a white or whitish to very pale brown or faintly greyish pileus, broadly ellipsoid to ovoid basidiospores and lamellar trama, pileipellis and stipitipellis hyphae with thickened walls (Matheny et al. 2020). The Pulverulina genus comprises two additional species besides Pulverulina ulmicola, namely Pulverulina cyathella (J. Favre & Schweers ex Kuyper) Chalange & P.-A. Moreau and Pulverulina praticola (Kuyper, Arnolds & P.-J. Keizer) Chalange & P.-A. Morea. These two species were transferred to Pulverulina by Chalange and Moreau (2023) from their previous classification under Omphalina. Both species can be readily distinguished from Pulverulina flavoalba based on their spore size and morphology. Specifically, the spores of Pulverulina praticola [(6.0-)6.5-8.0(-8.5) × (5.0-)5.5-6.5(-7.0) μm] are noticeably wider than those of Pulverulina flavoalba, resulting in a significantly lower Q value (Q = 1.1-1.3, Qmean = 1.2) compared to Pulverulina flavoalba (Kuyper et al. 1997). Similarly, Pulverulina cyathella also exhibits wider spores [(5.5-)6.5-7.0 × (5.0-)6.0-6.5 μm] and are (sub)globose in shape, distinguishing them from the cylindrical spores of Pulverulina flavoalba (Kuyper 1996).

Key to 22 species belonging to nine genera of Porotheleaceae in China

1 Lamellae not well developed Delicatula integrella
– Lamellae well developed 2
2 Pileocystidia present 3
– Pileocystidia absent 9
3 Cheilocystidia not seen Marasmiellomycena tomentosa
– Cheilocystidia abundant 4
4 Basidiospores inamyloid Megacollybia 5
– Basidiospores amyloid 6
5 Cheilocystidia digitate, narrowly or broadly clavate or sphaeropedunculate, rarely with short apical outgrowths Me. clitocyboidea
– Cheilocystidia clavate, without outgrowths Me. platyphylla
6 Cheilocystidia distinctly thick-walled overall Leucoinocybe 7
– Cheilocystidia thin-walled or slightly thick-walled in the base Clitocybula 8
7 Basidiospores narrowly ellipsoid L. lishuiensis
– Basidiospores subglobose to broadly ellipsoid L. subglobispora
8 Basidiospores (5.2) 5.4–5.8–6.2 (6.5) × (4.2) 4.3–4.7–5.0 (5.1) μm, broadly ellipsoid C. fuscostriata
– Basidiospores 3.5–5.3(–5.5) × 3.5–5.0 μm, globose, subglobose to broadly elliptic C. familia
9 Pileus trama sarcodimitic 10
– Pileus trama not sarcodimitic 18
10 Basidiospores inamyloid Trogia venenata
– Basidiospores amyloid Gerronema 11
11 Basidiomata distinctly small (Pileus < 9 mm in diam.) G. microcarpum
– Basidiomata moderately small (Pileus > 9 mm in diam.) 12
12 Pleurocystidia present G. chrysocarpum
– Pleurocystidia absent 13
13 Pileus blue G. indigoticum
– Pileus not blue 14
14 Pileus and stipe pure white G. albidum
– Pileus yellow to brown, stipe white to yellowish-brown 15
15 Pileus without pubescence or scales 16
– Pileus densely covered with deep brown pubescence or scales 17
16 Cheilocystidia up to 48 μm long G. baishanzuense
– Cheilocystidia less than 35 μm long G. nemorale
17 Stipe without fuscous pubescence or scales, basidiospores (6.3) 6.7–7.4–8.0 (8.5) × (3.2) 3.7–4.1–4.6 (4.8) μm G. zhujian
– Stipe with deep brown fuscous pubescence or scales, basidiospores (9.0) 9.2–10.0–11.2 (12.9) × (4.9) 5.2–5.8–6.6 (7.2) μm G. brunneosquamulosum
18 Cheilocystidia absent Pulverulina flavoalba
– Cheilocystidia present 19
19 Dermatocystidia inconspicuous and rare Pseudohydropus floccipes
– Dermatocystidia abundant Hydropus 20
20 Carpophore blackening when touched or bruised H. nigrita
– Carpophore not blackening in any part when touched or bruised 21
21 Basidiospores ellipsoid H. marginellus
– Basidiospores broadly ellipsoid H. atriceps

Discussion

Previous molecular phylogenetic analyses of the so-called hydropoid clade and the Porotheleaceae have been conducted, based on various combinations of ITS, 28S, 18S, 5.8S, 25S, rpb1 and rpb2 loci (Moncalvo et al. 2002; Matheny et al. 2006, 2020; Antonín et al. 2019; Vizzini et al. 2019, 2022; Consiglio et al. 2022; Senanayake et al. 2023). In the present study, we chose three regions, namely, ITS, nrLSU and rpb2, to analyse phylogenetic relationships in Porotheleaceae. Phylogenetic analyses, based on a combined dataset of these three loci, indicated that Marasmiellomycena comprising four species and Pulverulina, comprising two species, constitute monophyletic clades within Porotheleaceae. We thus report new records in China for two genera, Marasmiellomycena and Pulverulina, which cover two new species and a new combination. Marasmiellomycena now includes two new species, namely M. tomentosa and M. albodescendens. Additionally, the species previously identified as Porotheleum albodescendens has been combined as Marasmiellomycena albodescendens, representing a new combination within the Marasmiellomycena, all well characterised by having agaricoid basidiomata. On the basis of macromorphology and phylogenetic affinities, we have only retained one species in Porotheleum–the type species, Porotheleum fimbriatum (Pers.) Fr., which is distinguished by its fruiting clusters of small cup-shaped to tubular cream cyphelloid basidiomes that are densely crowded on a common membranous, resupinate subiculum/stroma with a broad rhizomorphic margin (Cooke 1989). Our results also agreed with Senanayake et al. (2023) that the genus Vizzinia contains two species V. dominingense and V. nigripes, which forms a well-supported lineage and the phylogenetic positions of Porotheleum albidum and Porotheleum parvulum are unclear.

Morphologically, Marasmiellomycena is easily recognisable as an omphalinoid mushroom in the field owing to its pileus that is depressed to umbilicate at the centre, decurrent to subdecurrent lamellae, dark-coloured stipe, sarcodimitic structure and thick-walled caulocystidia with contents. Marasmiellomycena is most similar to Vizzinia, but Vizzinia differs in basidiomata turning brownish on handling, distinctly squamulose pileus, weakly amyloid spores and absence of cheilocystidia. Pulverulina resembles Clitocybula in being an omphalinoid basidiocarps with decurrent lamellae, but can be distinguished by pruinose stipes, inamyloid basidiospores and absence of hymenial cystidia. Gerronema, Megacollybia and Trogia are more similar to Marasmiellomycena on the basis of their sarcodimitic structure. Marasmiellomycena can be readily discriminated in possessing dark-coloured stipe, inamyloid basidiospores and thick-walled caulocystidia with yellow to yellowish-brown pigments. Pulverulina species are characterised by their inamyloid basidiospores, non-sarcodimitic structure, thin-walled caulocystidia and non-pigmented pileocystidia and caulocystidia.

Our multi-gene phylogenetic analysis divided Gerronema into several highly-supported clades. This finding is consistent with the analyses of Antonín et al. (2019), Vizzini et al. (2019, 2022), Matheny et al. (2020) and Na et al. (2022a), who have reported that Gerronema is a non-monophyletic genus comprising several unrelated clades. The type of Gerronema has not been sequenced so it is unclear which belongs to Gerronema sensu stricto. Other genera in Porotheleaceae, namely, Chrysomycena, Clitocybula, Delicatula, Hydropodia, Hydropus, Leucoinocybe, Marasmiellomycena, Megacollybia, Pulverulina, Trogia and Vizzinia are monophyletic in previous phylogenetic studies as well as the present one (Matheny et al. 2020; Consiglio et al. 2022; Vizzini et al. 2022; Senanayake et al. 2023). Hydropodia subalpina (Höhn.) Vizzini, Consiglio & M. Marchetti, a new combination from Hydropus, is not related to Hydropus s. s.–which corresponds to the clade including the type species Hydropus fuliginarius (Batsch) Singer in the phylogenetic classification of Consiglio et al. (2022). In addition, Consiglio et al. (2022) consider Hydropodia to be sister to the Porotheleum clade; in our studies, however, Hydropodia is closer to Pseudohydropus and forms a sister clade.

Several species of Porotheleaceae have been reported to be edible or have toxic or ecological effects. Megacollybia platyphylla (Pers.) Kotl. & Pouzar (Dai et al. 2010), are known to be edible, whereas Trogia venenata Zhu L. Yang, Yan C. Li & L.P. Tang has caused hundreds of deaths in south-western China (Yang et al. 2012). Current evidence regarding the edibility and ecological functions of other Porotheleaceae species is insufficient. Specifically, whether they engage in symbiotic or saprophytic relationships with plants, as well as their roles within ecosystems, remains unclear. Although it is uncertain if these species exhibit symbiosis (and likely absent), future studies may uncover their capabilities to promote seed germination, similar to some Mycena species or possessing characteristics like bioluminescence. Further research is needed to investigate the edibility and ecological role of Porotheleaceae.

Acknowledgements

We thank Dr Junqing Yan (Jiangxi Agriculture University), Ms. Zewei Liu (Ludong University), Ms. Yulan Sun (Ludong University) and Ms. Renxiu Wei (Ludong University) for their kind help during fieldwork. We sincerely thank the reviewers for their corrections and suggestions to improve our work.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study was supported by the National Natural Science Foundation of China (grant no. 32200008), the Natural Science Foundation of Shandong Province (grant no. ZR2020QC001), the 5511 Collaborative innovation project of Fujian Province (grant no. XTCXGC2021007), the Central Public-Interest Scientific Institution Basal Research Fund (grant no. GYZX200203), the East-west cooperation project, FAAS (grant no. DKBF-2022-12), the Natural Science Foundation of Fujian Province (grant no. 2023J01379), the Project of Biological Resources Survey in Wuyishan National Park (grant no. HXQT2020120701), the Project of Biodiversity Conservation in Lishui, Zhejiang Province (grant no. HXYJCP2021110648), the Biodiversity investigation, observation and assessment program of Ministry of Ecology and Environment of China (grant no. 2110404 and 2019-2023), the Shandong Agricultural Industry Technology System (2021 grant no. 26, SDAIT-07-03) and the Soft Science project of Ministry of Public Security (grant no. 2022LL75).

Author contributions

Qin Na, Xianhao Cheng, and Yupeng Ge were responsible for designing the research and contributed significantly to data analysis and interpretation. Hui Zeng, Yaping Hu, Zhiheng Zeng, Bingrong Ke, Changjing Liu, and Hui Ding actively participated in the field investigation. All authors have contributed to the manuscript and approved the version submitted for publication.

Author ORCIDs

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

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

Yaping Hu https://orcid.org/0000-0003-1242-1139

Hui Ding https://orcid.org/0000-0003-4490-2105

Binrong Ke https://orcid.org/0009-0008-7209-7362

Zhiheng Zeng https://orcid.org/0009-0008-4208-2629

Xianhao Cheng https://orcid.org/0000-0002-5922-9913

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

Data availability

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

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