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Research Article
Three new species of Gerronema (Agaricales, Basidiomycota) from southern China
expand article infoWei-Xin Zhang‡§, Wang-Qiu Deng§, Chang-Qing Chang‡, Ping Zhou|, Min Lin§, Ming Zhang§
‡ South China Agricultural University, Guangzhou, China
§ Institute of Microbiology, Guangdong Academy of Sciences, Guangzhou, China
| Guangdong Nanling Forest Ecosystem National Observation and Research Station, Guangzhou, China

Corresponding author: Ming Zhang ( zhangming_1985@163.com )

Academic editor: María P. Martín
© 2025 Wei-Xin Zhang, Wang-Qiu Deng, Chang-Qing Chang, Ping Zhou, Min Lin, Ming Zhang.
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: Zhang W-X, Deng W-Q, Chang C-Q, Zhou P, Lin M, Zhang M (2025) Three new species of Gerronema (Agaricales, Basidiomycota) from southern China. MycoKeys 114: 239-258. https://doi.org/10.3897/mycokeys.114.145299
Open Access

Abstract

Three new species of Gerronema are discovered from southern China. Gerronema angustum is characterized by its small basidiomata, greenish-green pileus, slender stipe, narrow, and close lamellae. Gerronema pubescence is characterized by its pubescent pileus when young, yellowish white to pale yellow lamellae that are lighter towards the margin, narrowly cylindrical to lageniform pleurocystidia, and absent cheilocystidia. Gerronema rhizomorpha is characterized by its yellowish white to brown pileus, well-developed rhizomorphs at the stipe base, absent cheilocystidia and pleurocystidia, and not growing on rotten wood. Molecular phylogenetic analyses of nrITS + nrLSU support the species delimitation. In this study, detailed descriptions, photos of the basidiomata, line drawings, and discussions with related species are comprehensively provided. A key to the known Gerronema species in China is also provided.

Key words:

New taxa, phylogenetic analysis, Porotheleaceae, taxonomy

Introduction

Gerronema Singer (Porotheleaceae, Agaricales, Agaricomycetes, Basidiomycota), typified by G. melanomphax Singer, was first proposed to accommodate the three “rebellious” species from Argentina, which were characterized by their somewhat thick-walled hyphae and consequently more elastic-toughish consistency, strongly irregular hymenophoral trama, strongly intracellular pigmentation, and lignicolous habitat (Singer 1951). In this case, species of Gerronema only differ from species in Omphalina Quél. by the absence of fuscous, intraparietal, or incrusting pigments (Singer 1964, 1986; Bigelow 1970, 1982, 1985). Lange (1981) suggested that the characters defining various segregates of Omphalina are hardly of sufficient significance on the generic level, and Gerronema was regarded as a subgenus of Omphalina. Later, the characteristics of Gerronema were redefined as basidiomata omphalinoid to clitocyboid, pileus convex to infundibuliform or umbilicate, lamellae decurrent and subdistant, stipe central, basidiospores thin-walled, smooth, inamyloid, tramal tissue sarcodimitic, occasionally with cystidia and clamp connections, and lignicolous habitat (Redhead 1986; Norvell et al. 1994). And Chrysomphalina strombodes (Berk. & Mont.) Clémençon and Clitocybe xanthophylla Bres. were transferred into Gerronema (Norvell et al. 1994). Up to now, this definition is widely accepted by some scholars (Bañares et al. 2006; Antonín et al. 2008; Latha et al. 2018; Liu et al. 2019; Jayawardena et al. 2022; Na et al. 2022, 2024).

In previous studies, Gerronema was considered to be heterogenous by some scholars (Clémençon 1982; Moser 1983; Kuyper 1986; Singer 1986; Norvell et al. 1994). However, some researchers regarded Gerronema as a monophyletic group as restricted by Norvell et al. (1994) and included into the hydropoid clade together with Clitocybula (Singer) Singer ex Métrod, Hydropus Kühner ex Singer, Megacollybia Kotl. & Pouzar, and Porotheleum Fr. (Norvell et al. 1994; Moncalvo et al. 2002; Antonín et al. 2008, 2019; Yang et al. 2012). Molecular phylogenetic analyses provided new perspectives, and Gerronema was proved to be a polyphyletic group (Lutzoni 1997; Moncalvo et al. 2002; Redhead et al. 2002; Latha et al. 2018). The genus was resolved into several clades and closely related to the genera Megacollybia and Trogia Fr. in the family Porotheleaceae (Vizzini et al. 2019; Na et al. 2022, 2024). Up to now, the phylogenetic systematic position of Gerronema remains unclear due to the insufficient number of specimens and the limitation of phylogenetic research progress.

Gerronema is a small genus; only 75 species names have been recorded in Index Fungorum (http://www.indexfungorum.org, 1 November 2024). Most species of Gerronema are distributed in subtropical to tropical regions (Singer 1970; Norvell et al. 1994). China is one of the countries with the highest biodiversity and rich species, but only twelve Gerronema species have been reported, namely, G. albidum (Fr.) Singer, G. baisanzuens Q. Na, H. Zeng & Y.P. Ge, G. brunneosquamulosum Q. Na & Y.P. Ge, G. chrysocarpum P.G. Liu, G. confusum L. Fan & T.Y. Zhao, G. indigoticum T. Bau & L.N. Liu, G. lapidescens (Horan.) Ming Zhang & W.X. Zhang, G. kuruvense K.P.D. Latha & Manim., G. microcarpum Q. Na, H. Zeng & Y.P. Ge, G. nemorale Har. Takah., G. strombodes (Berk. & Mont.) Singer and G. zhujian Q. Na, H. Zeng & Y.P. Ge, seven of them are originally described from China (Liu 1995; Liu et al. 2019; Dai et al. 2010; Li et al. 2021; Wu et al. 2021; Na et al. 2022, 2024; Zhao and Fan 2022; Zhang et al. 2024). During our field investigation in southern China, three new species of Gerronema were found; they were formally described and introduced in morphological characters with molecular data in the present study.

Materials and methods

Sample collection and morphological study

Macroscopic morphological characteristics were derived from observation records and color images of fresh specimens collected in the field. Color descriptions were obtained according to Kornerup and Wanscher (1978). Samples were dried using an electric dryer at 50 °C and then deposited in the Fungarium of the Institute of Microbiology, Guangdong Academy of Sciences, Guangzhou, China (GDGM). The specific operations of the microscopic morphological characteristics are as follows: Sample each tissue section with tweezers or blades, place it in the slide floating carrier (1 drop of 5% KOH), stain with 1% Congo red solution, and observe the microstructure and measure it with a light microscope (Nikon Ni-U, Nikon Corporation, Japan). Twenty mature spores and 10 basidia were selected for measurement and represented by (a)b–c(d), where a or d indicates the extreme values, and b–c contains 90% of the measurements. L, W, and Q refer to length, width, and L/W ratio, respectively; Lm, Wm, and Qm refer to the mean length, width, and Q value of all basidiospore samples ± the standard deviation, respectively.

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

Genomic DNA samples were extracted from dried specimens using the HiPure Fungal DNA Kit (Magen Biotechnology Co., Ltd., Guangzhou, China) and kept in a -20 °C refrigerator. The internal transcribed spacer (nrITS) and the large subunit nuclear ribosomal DNA gene (nrLSU) were respectively amplified with primer pairs ITS1/ITS4 and LROR/LR7 (White et al. 1990; Hopple and Vilgalys 1999). PCR reactions were 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 2 × PCR mix (DreamTaqtm Green PCR Master Mix, Fermentas) (Zhang et al. 2022). The PCR procedure amplification was as follows: pre-denaturation at 95 °C for 5 min, 35 cycles of denaturation at 95 °C for 30 s, annealing at 56 °C (for nrITS)/50 °C (for nrLSU) for 40 s, extension at 72 °C for 50 s, and final extension at 72 °C for 10 min. After the amplification products were tested by agarose gel electrophoresis, PCR products were sent to Beijing BGI Co., Ltd. (Guangzhou, China) for sequencing. Then the sequences of the bidirectional sequencing were checked and assembled by the Geneious Pro trial 4.8.4 (Biomatters Limited Company). The newly obtained sequences were submitted to GenBank.

Phylogenetic analyses

The nrITS and nrLSU datasets were concatenated, including newly generated sequences, some valuable Gerronema sequences, and related genera (Clitocybula, Hydropus, Leucoinocybe Singer ex Antonín, Borovička, Holec & Kolařík, Marasmiellomycena De la Peña-Lastra, Mateos, Kolařík, Ševčíková & Antonín, Megacollybia, Porotheleum, Pseudohydropus Vizzini & Consiglio, Pulverulina Matheny & K.W. Hughes, and Trogia) sequences in GenBank were selected for phylogenetic analyses based on previous studies (Na et al. 2022, 2024) and listed in Table 1. Mycena purpureofusca (Peck) Sacc. was selected as the outgroup according to a previous study (Na et al. 2022). Using the auto strategy with MAFFT v7.505 (Katoh and Standley 2013) and then manually edited in MEGA v11.0.10 (Koichiro et al. 2021). The best model of nucleotide evolution for the dataset (nrITS + nrLSU) was identified using PartitionFinder 2 (Lanfear et al. 2016). Bayesian Inference (BI) and Maximum Likelihood (ML) bootstrap analyses were performed using the best-fit substitution models identified in PhyloSuite v1.2.3 (Zhang et al. 2020). The BI analysis was carried out in MrBayes 3.2.6 (Ronquist et al. 2012) under the best-fit substitution model, in which the initial 25% of sampled data were discarded as burn-in. Maximum likelihood phylogenies were inferred using IQ-TREE (Nguyen et al. 2015) under the edge-linked partition model for 5000 ultrafast (Minh et al. 2013) bootstraps. Phylogenetic trees were visualized using FigTree v.1.4.4. The maximum likelihood bootstrap over 50% (MLB ≥ 50%) and the Bayesian posterior probability over 0.90 (BPP ≥ 0.90) were shown.

Table 1.
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Information for the sequences used in the phylogenetic analyses. Newly generated sequences are in bold.

Taxon Voucher Locality GenBank accession No. Reference
nrITS nrLSU
Clitocybula abundans STU:SMNS-B-FU-2017/00898 Germany MF627833 – Direct Submission
C. familia PRM 921866 Czech Republic JF730327 JF730320 Antonín et al. (2011)
C. fuscostriata FFAAS1030 China OR238882 OR238894 Na et al. (2024)
Gerronema albidum H:6050710 USA – MF318923 Direct Submission
G. albidum H:6059277 USA – MF318924 Direct Submission
G. angustum GDGM 88662 China PQ452698 PQ350413 This study
G. angustum GDGM 88663 China PQ452699 – This study
G. atrovirens BKF10264 Thailand MZ452088 MZ452671 Jayawardena et al. (2022)
G. atrovirens BKF10265 Thailand MZ452668 MZ452672 Jayawardena et al. (2022)
G. baishanzuense FFAAS0359 China OL985962 OL985984 Na et al. (2022)
G. baishanzuense FFAAS0360 China OL985963 – Na et al. (2022)
G. baishanzuense FFAAS0361 China OL985964 – Na et al. (2022)
G. baishanzuense FFAAS0362 China OL985965 OL985986 Na et al. (2022)
G. baishanzuense FFAAS0363 China OL985966 OL985987 Na et al. (2022)
G. baishanzuense FFAAS0366 China OL985967 OL985988 Na et al. (2022)
G. brunneosquamulosum FFAAS1032 China OR238884 OR238896 Na et al. (2024)
G. brunneosquamulosum FFAAS1033 China OR238885 OR238897 Na et al. (2024)
G. citrinum G7458 French MN994795 – Jaouen et al. (2019)
G. citrinum G7785 French MN994822 – Jaouen et al. (2019)
G. citrinum PC0713130 French MN994747 – Jaouen et al. (2019)
G. citrinum PC0714037 French MN994655 – Jaouen et al. (2019)
G. confusum BJTC FM1592 China OK161262 – Zhao and Fan (2022)
G. confusum BJTC FM1624 China OK161271 – Zhao and Fan (2022)
G. indigoticum HMJAU47636 China MK693727 MK693732 Liu et al. (2019)
G. indigoticum HMJAU47942 China MK693728 MK693733 Liu et al. (2019)
G. indigoticum HMJAU47943 China MK693729 MK693734 Liu et al. (2019)
G. keralense BKF10263 Thailand MZ452107 MZ452144 Direct Submission
G. keralense CAL 1666 India MH156555 MH153979 Latha et al. (2018)
G. kuruvense BKF10266 Thailand MZ452090 MZ452669 Direct Submission
G. kuruvense CAL 1665 India MH156554 MH153978 Latha et al. (2018)
G. kuruvense DCY3362(HGASMF01-15010) China MZ951144 – Direct Submission
G. kuruvense FCATAS9085 China PP622159 – Direct Submission
G. kuruvense KUC20220701_03 Korea OR600252 – Cho et al. (2024)
G. lapidescens GDGM 85271-1 China OR736197 – Zhang et al. (2024)
G. lapidescens GDGM 85271-2 China OR736198 – Zhang et al. (2024)
G. lapidescens GDGM 86705 China OR736202 – Zhang et al. (2024)
G. microcarpum FFAAS0371 China OL985968 OL985990 Na et al. (2022)
G. microcarpum FFAAS0372 China OL985969 OL985991 Na et al. (2022)
G. microcarpum FFAAS0373 China OL985970 OL985992 Na et al. (2022)
G. microcarpum FFAAS0374 China OL985971 – Na et al. (2022)
G. microcarpum FFAAS0375 China OL985972 OL985993 Na et al. (2022)
G. nemorale HMJAU59063 China OK560883 – Direct Submission
G. nemorale HMJAU59064 China OK560871 – Direct Submission
G. nemorale FA236 Pakistan MN744687 – Aqdus and Khalid (2021)
G. nemorale FA239 Pakistan MN744688 – Aqdus and Khalid (2021)
G. nemorale FA249 Pakistan MN744686 – Aqdus and Khalid (2021)
G. nemorale FFAAS0389 China OL985981 OL986002 Na et al.(2022)
G. nemorale FFAAS0392 China OL985982 OL986003 Na et al.(2022)
G. nemorale FFAAS0410 China OL985983 OL986004 Na et al.(2022)
G. pubescence GDGM 93936 China PQ452700 PQ350414 This study
G. pubescence GDGM 94001 China PQ452701 PQ350415 This study
G. rhizomorpha GDGM 87835 China PQ452702 – This study
G. rhizomorpha GDGM 92067 China PQ452703 PQ350416 This study
Gerronema sp. HMJAU59018 China OK491123 – Direct Submission
G. strombodes FLAS-F-60957 USA MH016911 – Direct Submission
G. strombodes FLAS-F-71339 USA OR438652 – Direct Submission
G. strombodes TENN:F-60009 USA KY271083 – Direct Submission
G. strombodes TFB12519/TENN60718 USA EU623640 – Hughes et al. (2007)
G. strombodes TFB12783/TENN61350 USA EU623641 – Hughes et al. (2007)
G. strombodes DJL05NC72 USA EU623639 – Hughes et al. (2007)
G. subclavatum FLAS-F-60986 USA MH016932 – Direct Submission
G. subclavatum FLAS-F-61518 USA MH211945 – Direct Submission
G. subclavatum FLAS-F-71359 USA OR242635 – Direct Submission
G. subclavatum iNaturalist # 8545787 India MN906021 – Direct Submission
G. subclavatum Mushroom Observer # 243440 USA MK607510 – Direct Submission
G. subclavatum S.D. Russell MycoMap # 6854 India MN906138 – Direct Submission
G. subclavatum Smith-2018 iNaturalist # 17333993 USA MK573888 – Direct Submission
G. viridilucens DED 7822 Brazil – OR449361 Direct Submission
G. waikanaense PDD:87667 New Zealand JQ694117 – Direct Submission
G. xanthophyllum PRM 924657 Czech Republic LT854023 LT854023 Antonín et al. (2019)
G. xanthophyllum SYKOf3970 Russia OR915457 – Direct Submission
G. zhujian FFAAS0370 China OL985974 OL985995 Na et al. (2022)
G. zhujian FFAAS0376 China OL985975 OL985996 Na et al. (2022)
Hydropus fuliginarius S.D. Russell ONT iNaturalist # 130794969 USA OP643427 – Direct Submission
H. marginellus OSC 112834 USA EU669314 EU852808 Direct Submission
H. rugosodiscus MGW1257 USA KY777386 – Direct Submission
Leucoinocybe danxiashanensis GDGM 80184 China MZ667478 MZ667482 Direct Submission
L. lishuiensis FFAAS0115 China MW424491 MW424495 Na et al. (2021)
L. subglobispora FFAAS1034 China OR238886 OR238898 Na et al. (2024)
Oudemansiella aff. platyphylla 360-630 Japan AB509870 – Direct Submission
Marasmiellomycena pseudoomphaliiformis BRNM:552721 USA OR913562 OR913566 Senanayake et al. (2023)
M. tomentosa FFAAS1036 China OR238888 OR238900 Na et al. (2024)
Megacollybia clitocyboidea TENN62231 USA EU623664 – Hughes et al. (2007)
M. marginata HR 91607 Czech Republic LT854051 – Antonín et al. (2019)
M. platyphylla BRNM 737654 Czech Republic LT854048 LT854036 Antonín et al. (2019)
Mycena purpureofusca HMJAU43554 China MG654740 MK629356 Na and Bau (2018)
M. purpureofusca HMJAU43624 China MG654741 MK629357 Na and Bau (2018)
Porotheleum fimbriatum CLZhao 1120 China MH114870 – Direct Submission
P. fimbriatum Dai 12276 China KX081137 KX161656 Direct Submission
Pseudohydropus floccipes BRNM 816173 Czech Republic OM422758 OM423634 Direct Submission
P. floccipes BRNM 825631 Spain OM422760 OM423636 Consiglio et al. (2022)
P. globosporus BAP 661 USA MH414566 MH385340 Cooper et al. (2019)
Pulverulina flavoalba FFAAS1039 China OR238891 OR238903 Na et al. (2024)
P. flavoalba FFAAS1040 China OR238892 OR238904 Na et al. (2024)
P. ulmicola TFB13871 USA MT237476 MT237446 Matheny et al. (2020)
Trogia benghalensis CUH AM031 India KU647630 – Dutta et al. (2017)
T. infundibuliformis KUN_HKAS56709 China JQ031776 JQ031781 Yang et al. (2012)
T. venenata KUN_HKAS54710 China JQ031772 JQ031778 Yang et al. (2012)

Results

Molecular phylogenetic results

The final concatenated dataset consisted of 95 nrITS and 52 nrLSU sequences from 46 taxa of 11 genera of Physalacriaceae, Porotheleaceae, and Mycenaceae, which comprised 1927 nucleotide sites (942 for nrITS, 985 for nrLSU), of which 752 were parsimony-informative, 277 were singleton sites, and 898 were constant sites. For the ML analyses, the best-fit substitution models selected for nrITS and nrLSU region partitions in the concatenated dataset were HKY+I+G and GTR+I+G, respectively. For the BI analysis, the best-fit substitution model selected for each of the two DNA regions was GTR+I+G (2 parallel runs, 2000000 generations), and the average standard deviation of split frequencies was stably dropped under 0.01. The phylogenetic trees generated from BI and ML analyses show almost similar topologies and few variations in statistical support, so only the ML tree is displayed (Fig. 1).

Figure 1. 

Phylogenetic consensus tree of Gerronema species inferred from the maximum likelihood and Bayesian inference based on a concatenated nrITS and nrLSU (MLB ≥ 50%, BPP ≥ 0.90 are shown). The tree is rooted with Mycena purpureofusca. New taxa are shown in bold.

In the phylogenetic tree (Fig. 1), Clitocybula, Gerronema, Hydropus, Leucoinocybe, Marasmiellomycena, Megacollybia, Porotheleum, Pseudohydropus, Pulverulina, and Trogia were nested in the core clade of Porotheleace with significant support (MLB = 100%, BPP = 1.00). The proposed new species formed three independent lineages within the genus Gerronema (MLB = 98%, BPP = 1.00), named G. angustum, G. pubescence, and G. rhizomorpha. In addition, G. angustum is sister to G. waikanaense (G. Stev.) J.A. Cooper and an unnamed specimen (HMJAU59018) (MLB = 100%, BPP = 1.00). Gerronema pubescence closely related to G. baishanzuense with a well-supported (MLB = 94%, BPP = 0.95). Gerronema rhizomorpha is placed at the base of the Gerronoma clade.

Taxonomy

Gerronema angustum Ming Zhang & W.X. Zhang, sp. nov.

Fungal Names: FN 572081
Figs 2, 3

Diagnosis.

Distinguished from other Gerronema species by the combination characters, including caespitose habit, greenish green pileus, narrow and close lamellae, slender stipe, baidiospores measuring (4)4.5–5.5 × 2.5–3.5 μm, mainly clavate to narrowly utriform cheilocystidia.

Figure 2. 

Basidiomata of Gerronema angustum A collection GDGM 88662, holotype B collection GDGM 88663. Photographed by Bin Song. Scale bars: 10 mm.

Holotype.

China • Guangdong Province: Shaoguan City, Nanling National Forest Park; 24°53'54"N, 113°2'24"E; 210 m asl.; 7 July 2022; Bin Song, Guo-Rui Zhong, and De-Chun Xie (GDGM 88662).

Figure 3. 

Microscopic features of Gerronema angustum (GDGM 88662, holotype) A Basidiospores B Basidia C Cheilocystidia D Pileipellis E Stipitipellis. Scale bars: 10 μm (A–C); 50 μm (D–E).

Etymology.

angustum (Latin), referring to the narrow lamellae of this species.

Description.

Basidiomata small-sized. Pileus 10–18 mm broad, infundibuliform, umbilicate to deeply umbilicate at center, greyish brown to brown (6E3–4) when young, greyish green (30B3–4, 30C2–3) when old, greyish green (30E5–6) at center, surface moist, glabrous, margin inflexed, radially striped with greenish grey to dull green (30B4–5, 30D4–5) lines. Lamellae decurrent, close, narrow, arcuate, even, white (30A1) to greenish grey (30B2), with 1–3 lamellulae. Stipe 45–60 × 2–5 mm, slender, centric, cylindrical, hollow, fragile, grey to greyish green (30C1–3), covered with white (30A1) fibrils. Odor and taste not recorded.

Basidiospores (4)4.5–5.5 × 2.5–3.5 μm, Lm = 4.88 ± 0.51 µm, Wm = 2.96 ± 0.32 µm, Q = (1.33)1.43–1.83, Qm = 1.66 ± 0.18, ellipsoid to oblong, smooth, thin-walled, hyaline, guttulate, inamyloid. Basidia 18–26 × 5.5–7 μm, clavate, thin-walled, hyaline, 4-spored, with sterigmata 2.3–4.4 µm long. Cheilocystidia 26–45 × 6–9.5 μm, clavate, fusiform to narrowly utriform, thin-walled, hyaline. Pleurocystidia not seen. Lamellar trama regular to subregular, hyphae 3–22 μm wide, cylindrical, thin-walled, hyaline. Pileipellis a cutis, hyphae 1.5–24.5 μm wide, smooth, hyaline; pileocystidia 22.5–65 × 8–15.5 µm, oblong to utriform, thin-walled, greyish brown to light brown pigmented in KOH. Pileus trama subregular, sarcodimitic. Stipitipellis a cutis, hyphae 3.5–25 μm wide, sometimes upturned hyphae, smooth, thin-walled, hyaline; caulocystidia 56–72 × 10.5–20.5 µm, narrowly cylindrical to oblong, thin-walled, hyaline. Stipe trama regular, sarcodimitic. Clamp connections present in all tissues.

Habit and distribution.

Caespitose on the rotten wood in broad-leaved forests. Currently only known from the type locality in China.

Additional specimen examined.

China • Guangdong Province: Shaoguan City, Nanling National Forest Park; 24°55'39"N, 113°3'20"E; 225 m asl.; 7 July 2022; Bin Song, Guo-Rui Zhong, and De-Chun Xie (GDGM 88663).

Gerronema pubescence Ming Zhang & W.X. Zhang, sp. nov.

Fungal Names: FN 572082
Figs 4, 5

Diagnosis.

Distinguished from other Gerronema species by the combination characters of the pastel grey pileus covered with pubescence when young, yellowish white to pale yellow lamellae are paler towards the margin, the absence of cheilocystidia, and the narrow cylindrical to utriform pleurocystidia.

Figure 4. 

Basidiomata of Gerronema pubescence A, B collection GDGM 94001, holotype C, D collection GDGM 93936. A, B Photographed by Hao Huang C, D photographed by Ming Zhang. Scale bars: 30 mm (A, B); 10 mm (C, D).

Holotype.

China • Guangdong Province: Huizhou City, Xiangtou Mountain Nature Reserve; 23°26'N, 114°37'E; 335 m asl.; 19 September 2023; Hao Huang and Wei-Xin Zhang (GDGM 94001).

Figure 5. 

Microscopic features of Gerronema pubescence (GDGM 94001, holotype) A Basidiospores B Basidia C Pleurocystidia D Pileipellis E Stipitipellis. Scale bars: 10 mm (A, B); 50 mm (C–E).

Etymology.

pubescence (Latin), referring to the species pileus usually covered with pubescence when young.

Description.

Pileus 12–70 mm broad, hemispherical to plano-convex, depressed at center, grey (1E1), covered with pubescence when young, white to yellowish white (1A1–2) with age, grey to greyish brown (5E2–3, 6C1–2) at center, shallowly sulcate, surface dry, glabrous or pubescent, distinctly radially striped with grey to brownish orange (4C1–2, 6C2–3) lines, margin inflexed to reflexed. Lamellae subdecurrent, subdistant, ventricose, even, yellowish white to pale yellow (4A2–3), white (4C1) towards margin, with 1–5 lamellulae. Stipe 15–40 × 2–7 mm, central, cylindrical, hollow, white (1A1) to grey (1C1), covered with white granulose or fibrils. Odor and taste not recorded.

Basidiospores (6)6.5–8 × (3.5)4–4.5 μm, Lm = 7.13 ± 0.57 µm, Wm = 4.08 ± 0.29 µm, Q = (1.5)1.56–2, Qm = 1.75 ± 0.18, ellipsoid to oblong, smooth, thin-walled, hyaline, guttulate, inamyloid. Basidia 24.5–39 × 5–7.5 μm, clavate, thin-walled, hyaline, 2- or 4-spored, with sterigmata 2.5–5 µm long. Cheilocystidia absent. Pleurocystidia 40–104 × 8.5–12.5 µm, narrowly cylindrical to lageniform, thin-walled, hyaline. Lamellar trama regular to subregular, hyphae 3.5–26.5 μm wide, thin-walled, hyaline. Pileipellis a cutis, hyphae 3–25 μm wide, smooth, hyaline; pileocystidia 39–100 × 10–21 μm, oblong to narrowly clavate, apex sometimes rostrate, thin-walled, greyish brown pigmented in KOH. Pileus trama regular to subregular, sarcodimitic. Stipitipellis a cutis, hyphae 2.5–38.5 μm wide, smooth, thin-walled, hyaline; caulocystidia 38.5–84.5 × 8.5–19 μm, narrowly cylindrical to clavate, thin-walled, hyaline. Stipe trama regular, sarcodimitic. Clamp connections present in all tissues.

Habit and distribution.

Scattered or caespitose on the rotten wood in broad-leaved forests. Currently only known from the type locality in China.

Additional specimen examined.

China • Guangdong Province: Zhaoqing City, Dinghu Mountain Nature Reserve; 23°10'43"N, 112°33'10"E; 150 m asl.; 9 April 2024; Ming Zhang, Guo-Rui Zhong, and Wen-Xiao Xia (GDGM 93936).

Gerronema rhizomorpha Ming Zhang & W.X. Zhang, sp. nov.

Fungal Names: FN 572083
Figs 6, 7

Diagnosis.

Distinguished from other Gerronema species by the combination characters of medium-sized basidiomata, yellowish white to brown pileus, well-developed rhizomorphs at stipe base, the absence of cheilocystidia, and pleurocystidia.

Figure 6. 

Basidiomata of Gerronema rhizomorpha A, B collection GDGM 92067, holotype C, D collection GDGM 87835. Photographed by Ming Zhang. Scale bars: 30 mm (A, B); 50 mm (C); 30 mm (D).

Holotype.

China • Guangdong Province: Fengkai County, Zhaoqing City, Heishiding provincial natural reserve; 23°26'30"N, 111°53'28"E; 340 m asl.; 25 May 2023; Ming Zhang and Guo-Rui Zhong (GDGM 92067).

Figure 7. 

Microscopic features of Gerronema rhizomorpha (GDGM 92067, holotype) A Basidiospores B Basidia C Pileipellis D Stipitipellis. Scale bars: 10 mm (A, B); 50 mm (C, D).

Etymology.

rhizomorpha (Latin), referring to the species, usually has long and well-developed rhizomorphs at stipe base.

Description.

Basidiomata medium-sized. Pileus 35–95 mm broad, hemispherical to plano-convex, depressed at center, brown (5E5–6) when young, yellowish white (2A2–3) to brown (5E4–5) at maturity, usually brown (5E4–5) to dark brown (6F4–5) at center, surface dry, distinctly radially striped with brown (5E6–7, 6E4–5) lines, margin inflexed. Lamellae subdecurrent, subdistant, ventricose, even, white to yellowish white (1A1–2), with 1–3 lamellulae. Stipe 40–110 × 4–10 mm, central to eccentric, cylindrical, hollow, white to yellowish white (1A1–2), sometimes yellowish grey to greyish yellow (2C2–3) at the base, covered with yellowish grey to greyish yellow (2C2–4) fibrils, base with developed and white (1A1) rhizomorphs. Odor and taste not recorded.

Basidiospores 7–8.5(9.5) × 5–6(6.5) μm, Lm = 7.80 ± 0.71 µm, Wm = 5.53 ± 0.42 µm, Q = (1.23)1.27–1.58(1.6), Qm = 1.41 ± 0.13, broadly ellipsoid to ellipsoid, smooth, thin-walled, hyaline, guttulate, inamyloid. Basidia 25.5–32 × 7.5–11.5 μm, clavate, thin-walled, hyaline, 4-spored, with sterigmata 2.8–6 µm long. Hymenial cystidia absent. Lamellar trama regular to subregular, hyphae 3–22 μm wide, thin-walled, hyaline. Pileipellis a cutis, hyphae 3–25 μm wide, thin-walled, light yellow in KOH; pileocystidia 25.5–63.5 × 6–14 μm, narrowly utriform to cylindrical, thin-walled, greyish brown to light brown pigmented in KOH. Pileus trama subregular, sarcodimitic. Stipitipellis a cutis, hyphae 1.5–11 μm wide, smooth, thin-walled, hyaline; caulocy stidia 12.5–33 × 3.5–8 μm, narrowly cylindrical to oblong, thin-walled, hyaline. Stipe trama regular, sarcodimitic. Clamp connections present in all tissues.

Habit and distribution.

Solitary or scattered, growing on the damp soil under broad-leaved forests. Currently known from China and Japan.

Additional specimen examined.

China • Guangdong Province: Shaoguan City, Nanling National Forest Park; 24°56'48"N, 113°3'19"E; 490 m asl.; 9 June 2022; Ming Zhang, Guo-Rui Zhong, and Shi-Zheng Wang (GDGM 87835).

Discussion

Morphologically, Gerronema angustum can be easily distinguished from other species in Gerronema by its caespitose habit, slender basidiomata, greenish-green pileus, narrow and close lamellae, and ellipsoid to oblong basidiospores measuring (4)4.5–5.5 × 2.5–3.5 μm. Gerronema angustum is similar to G. albidum, which has been recorded in China (Singer 1962). But the distinctly white basidiomata and the absence of cheilocystidia can be used to distinguish G. albidum from G. angustum.

Gerronema pubescence is characterized by its hemispherical to plano-convex pileus covered with pubescence when young, yellowish white to pale yellow lamellae paler towards the pileus margin, absent cheilocystidia, and narrowly cylindrical to utriform pleurocystidia. Gerronema pubescence is similar to G. keralense K.P.D. Latha & Manim. and G. zhujian. However, G. keralense, originally described from India, can be distinguished by its small yellowish-brown pileus (4–17 mm broad), greyish-yellow stipe gradually greyish-brown towards the base, flexuose or irregular cheilocystidia, and absent pleurocystidia (Latha et al. 2018). Gerronema zhujian can be distinguished by the greyish-white pileus with a brown tinge at the center, slightly brown and narrow stipe (19–25 × 1.0–1.5 mm), subfusiform cheilocystidia, and the absence of pleurocystidia (Na et al. 2022).

Gerronema rhizomorpha is mainly characterized by its yellowish white to brown pileus, well-developed rhizomorphs at stipe base, and absent cheilocystidia and pleurocystidia. Gerronema rhizomorpha is similar to G. confusum in some extent as sharing relatively large basidiomata and brown pileus. But G. confusum from the north of China can be distinguished by its lignicolous habitat, greyish-brown stipe surface covered with dark brown granulose, absent rhizomorphs, 1–2-spored basidia, and abundant subcylindrical to cylindrical cheilocystidia (Zhao and Fan 2022). Gerronema rhizomorpha is also similar to G. atrialbum (Murrill) Borovička & Kolařík, with the stipe base often with rhizomorphs, white lamellae, and the absence of cheilocystidia and pleurocystidia. However, G. atrialbum, which is originally described from the USA, can be distinguished by its greyish brown to grey pileus and brown to pale stipe, relatively longer basidia (36–50 × 6.5–9.5 μm), which mainly has 2-spored. In addition, G. atrialbum mainly grows on humus or rotting hardwood (Murrill 1913; Antonín et al. 2019).

Phylogenetically (Fig. 1), our new species formed three distinct lineages according to the ML and BI phylogenetic analyses of the concatenated dataset and can be easily distinguished from other species with known sequences. Gerronema angustum formed a distinct lineage in Gerronema and is sister to G. waikanaense with high statistical support (MLB = 100%, BPP = 1.00). However, G. waikanaense, reported from New Zealand, differs by its dark leaden grey basidiomata, minutely fibrillose stipe surface, distant lamellae, and absent cheilocystidia (Cooper 2014).

Gerronema pubescence is phylogenetically related to G. baishanzuense (MLB = 94%, BPP = 0.95). But G. baishanzuense can be distinguished by its relatively small pileus (3–25.5 mm broad), relatively short stipe (4.5–26 mm) densely covered with pruinose when young, clavate or subfusiform cheilocystidia usually swollen at apex, and absent pleurocystidia (Na et al. 2022).

Gerronema rhizomorpha was gathered together with an ITS sequence (AB509870) named Oudemansiella sp. (platyphylla 360–630) from Japan (MLB = 100%, BPP = 1.00) and shows that they represent the same phylogenetic species. It is noteworthy that G. rhizomorpha with well-developed rhizomorphs at stipe base and not growing on rotten wood is rare in Gerronema, which is morphologically more similar to species of the genus Megacollybia, but it does not belong to Megacollybia in phylogenetic analyses. Additionally, G. rhizomorpha has significant variability in the ITS1 region, has a low similarity rate with Gerronema, and is placed at the base of the Gerronema clade, possibly representing a separate evolutionary lineage. As the Gerronema genus is not a monophyletic group, G. rhizomorpha was temporarily classified as a member of Gerronema in the present study.

Due to Gerronema being widely distributed and many species being misidentified, only ten of the 20 species reported in Asia are from China. Recent investigations have found a high species diversity of Gerronema in southern China, and there are still many other species waiting to be reported, which can enrich the species diversity of the genus. In addition, the intraspecific and intergeneric phylogenetic relationships of Gerronema still remain highly controversial. Upon defining the diversity of species explicitly, the systematic phylogenetic framework of the genus Gerronema needs to be further refined based on more samples and sequence fragments to solve the classification problem of the Gerronema.

Key to species of Gerronema in China

1 Baisidiomata grow on soil, stipe base with well-developed rhizomorphs 2
– Baisidiomata grow on decaying woods, stipe base without developed rhizomorphs 3
2 Sclerotia can be found at the base G. lapidescens
– Sclerotia not be found or recorded G. rhizomorpha
3 Basidiomata blue G. indigoticum
– Basidiomata without blue tinge 4
4 Basidiomata greyish green G. angustum
– Basidiomata without greyish green tinge 5
5 Basidiomata white G. albidum
– Basidiomata without white tinge 6
6 Pleurocystidia present 7
– Pleurocystidia absent 8
7 Pileus viscid G. chrysocarpum
– Pileus dry, usually covered with pubescence when young. G. pubescence
8 Pileus densely covered with deep brown fur or scales 9
– Pileus without fur or scales 10
9 Stipe fuscous, basidiospores 9.0–12.9 × 4.9–7.2 μm G. brunneosquamulosum
– Stipe white, basidiospores 6.3–8.5 × 3.2–4.8 μm G. zhujian
10 Basidiomata small-sized, Pileus usually ≤ 11 mm in diam. 11
– Basidiomata small to medium-sized, Pileus ≥ 11 mm in diam 12
11 Basidiospores 6.1–7.5 × 3.5–4.3 μm, cheilocystidia present G. microcarpum
– Basidiospores 8–11 × 4–6 μm, cheilocystidia absent G. kuruvense
12 Basidia 1–2-spored G. confusum
– Basidia 4-spored 13
13 Pileus up to 25 mm in diam, stipe 4.5–26.0 × 0.5–2.0 mm G. baishanzuense
– Pileus less than 20 mm in diam, stipe 19.0–36.0 × 1.0–2.5 mm G. nemorale

Acknowledgments

We would like to thank Bin Song, De-Chun Xie, Guo-Rui Zhong, Hao Huang, Shi-Zheng Wang, and Wen-Xiao Xia (Institute of Microbiology, Guangdong Academy of Sciences, Guangzhou, China) for their help during the field trips. We would also like to sincerely thank the editors and anonymous reviewers for their valuable suggestions and comments.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work is financed by the Science & Technology Fundamental Resources Investigation Program (grant no. 2022FY100500), the National Natural Science Foundation of China (grant no. 32360008), the Science and Technology Project of Forestry Administration of Guangdong Province (grant no. LC-2021124), the Open Fund of Guangdong Nanling Forest Ecosystem National Observation and Research Station (grant no. NLFP202403), and the GDAS’ Project of Science and Technology Development (Grant no. 2022GDASZH-2022010201-01).

Author contributions

All authors have contributed to the manuscript and approved the version submitted for publication.

Author ORCIDs

Ming Zhang https://orcid.org/0000-0001-6420-2531

Data availability

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

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