683 MycoKeys MycoKeys 114: 239-258 (2025) DOI: 10.3897/mycokeys.114.145299 Research Article Three new species of Gerronema (Agaricales, Basidiomycota) from southern China Wei-Xin Zhang'2, Wang-Qiu Deng2, Chang-Qing Chang’, Ping Zhou’, Min Lin?, Ming Zhang2® 1 Guangdong Provincial Key Laboratory of Microbial Signals and Disease Control, Engineering Research Center of Biological Control, Ministry of Education, State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources, South China Agricultural University, Guangzhou 510642, China 2 State Key Laboratory of Applied Microbiology Southern China, Guangdong Provincial Key Laboratory of Microbial Culture, Collection and Application, Institute of Microbiology, Guangdong Academy of Sciences, Guangzhou 510070, China 3 Guangdong Nanling Forest Ecosystem National Observation and Research Station, Guangzhou, China Corresponding author: Ming Zhang (zhangming@gdim.cn) OPEN Qaceess Academic editor: Maria PR. Martin Received: 25 December 2024 Accepted: 18 January 2025 Published: 4 March 2025 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 Copyright: © Wei-Xin Zhang et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract Three new species of Gerronema are discovered from southern China. Gerronema an- gustum 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. Ger- ronema rhizomorpha is characterized by its yellowish white to brown pileus, well-devel- oped rhizomorphs at the stipe base, absent cheilocystidia and pleurocystidia, and not growing on rotten wood. Molecular phylogenetic analyses of nrlTS + 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, Basidiomyco- ta), 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 pig- mentation, and lignicolous habitat (Singer 1951). In this case, species of Ger- ronema 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 Ger- ronema was regarded as a subgenus of Omphalina. Later, the characteristics of Gerronema were redefined as basidiomata omphalinoid to clitocyboid, pile- us convex to infundibuliform or umbilicate, lamellae decurrent and subdistant, stipe central, basidiospores thin-walled, smooth, inamyloid, tramal tissue sar- 239 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China codimitic, occasionally with cystidia and clamp connections, and lignicolous habitat (Redhead 1986; Norvell et al. 1994). And Chrysomphalina strombodes (Berk. & Mont.) Clémencon and Clitocybe xanthophylla Bres. were transferred into Gerronema (Norvell et al. 1994). Up to now, this definition is widely accept- ed by some scholars (Bafares et al. 2006; Antonin 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 (Clemencon 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 Kihner ex Singer, Megacollybia Kotl. & Pouzar, and Porotheleum Fr. (Norvell et al. 1994; Moncalvo et al. 2002; Antonin et al. 2008, 2019; Yang et al. 2012). Molecular phylogenetic analyses provided new perspectives, and Gerronema was proved to be a poly- phyletic 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 biodi- versity and rich species, but only twelve Gerronema species have been report- ed, 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 de- scribed 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 charac- ters with molecular data in the present study. Materials and methods Sample collection and morphological study Macroscopic morphological characteristics were derived from observation re- cords and color images of fresh specimens collected in the field. Color descrip- tions 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 char- acteristics 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 MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 240 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China red solution, and observe the microstructure and measure it with a light micro- scope (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; L_, W_, and Q_ 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 uL containing 1 pL template DNA, 9.5 uL distilled water, 1 uL of each primer, and 12.5 pL 2 x PCR mix (DreamTaq'™ 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 nrlITS)/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 Lim- ited Company). The newly obtained sequences were submitted to GenBank. Phylogenetic analyses The nrlTS and nrLSU datasets were concatenated, including newly generated sequences, some valuable Gerronema sequences, and related genera (Clito- cybula, Hydropus, Leucoinocybe Singer ex Antonin, Borovicka, Holec & Kolarik, Marasmiellomycena De la Pefia-Lastra, Mateos, Kolarik, Sevéikova & Antonin, 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 out- group 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 Mr- Bayes 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. MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 241 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China Table 1. Information for the sequences used in the phylogenetic analyses. Newly generated sequences are in bold. Clitocybula abundans GC. C Gerronema albidum G. G. G. G. MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 ala) ale|oalala|a|!/@e!/@|]ala!l@e|ealalalaloal!]alala|@|/ea!lala|]e/.a@|a/a!a]e Taxon familia fuscostriata albidum angustum angustum atrovirens atrovirens baishanzuense baishanzuense baishanzuense baishanzuense baishanzuense baishanzuense brunneosquamulosum brunneosquamulosum citrinum citrinum citrinum citrinum confusum confusum . indigoticum . indigoticum . indigoticum . keralense . keralense . kuruvense . kuruvense . kuruvense . kuruvense . kuruvense . lapidescens . lapidescens . lapidescens . Microcarpum . Microcarpum . Microcarpum . Microcarpum Voucher STU:SMNS-B-FU-2017/00898 PRM 921866 FFAAS1030 H:6050710 H:6059277 GDGM 88662 GDGM 88663 BKF 10264 BKF10265 FFAASO0359 FFAASO360 FFAASO361 FFAAS0362 FFAAS0363 FFAASO366 FFAAS1032 FFAAS1033 G7458 G7785 PC0713130 PC0714037 BJTC FM1592 BJTC FM1624 HMJAU47636 HMJAU47942 HMJAU47943 BKF10263 CAL 1666 BKF10266 CAL 1665 DCY3362(HGASMF01-15010) FCATAS9085 KUC20220701_03 GDGM 85271-1 GDGM 85271-2 GDGM 86705 FFAASO371 FFAASO372 FFAASO0373 FFAAS0374 Locality Germany Czech Republic China USA USA China China Thailand Thailand China China China China China China China China French French French French China China China China China Thailand India Thailand India China China Korea China China China China China China China GenBank accession No. nrITS MF627833 JF730327 OR238882 PQ452698 PQ452699 MZ452088 MZ452668 OL985962 OL985963 OL985964 OL985965 OL985966 OL985967 OR238884 OR238885 MN994795 MN994822 MN994747 MN994655 OK161262 OK161271 MK693727 MK693728 MK693729 MZ452107 MH156555 MZ452090 MH156554 MZ951144 PP622159 OR600252 OR7/36197 OR736198 OR7/36202 OL985968 OL985969 OL985970 OL985971 nrLSU JF730320 OR238894 MF318923 MF318924 PQ350413 MZ452671 MZ452672 OL985984 OL985986 OL985987 OL985988 OR238896 OR238897 MK693732 MK693733 MK693734 MZ452144 MH153979 MZ452669 MH153978 OL985990 OL985991 OL985992 Reference Direct Submission Antonin et al. (2011) Na et al. (2024) Direct Submission Direct Submission This study This study Jayawardena et al. (2022) Jayawardena et al. (2022) Na et al. (2022) Na et al. (2022) Na et al. (2022) Na et al. (2022) Na et al. (2022) Na et al. (2022) Na et al. (2024) Na et al. (2024) Jaouen et al. (2019) Jaouen et al. (2019) Jaouen et al. (2019) Jaouen et al. (2019) Zhao and Fan (2022) Zhao and Fan (2022) Liu et al. (2019) Liu et al. (2019) Liu et al. (2019) Direct Submission Latha et al. (2018) Direct Submission Latha et al. (2018) Direct Submission Direct Submission Cho et al. (2024) Zhang et al. (2024) Zhang et al. (2024) Zhang et al. (2024) Na et al. (2022) Na et al. (2022) Na et al. (2022) Na et al. (2022) 242 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China Taxon G. microcarpum G. nemorale G. nemorale G. nemorale G. nemorale G. nemorale G. nemorale G. nemorale G. nemorale G. pubescence G. pubescence G. rhizomorpha G. rhizomorpha Gerronema sp. strombodes strombodes strombodes strombodes strombodes strombodes subclavatum subclavatum G. subclavatum G. subclavatum G. subclavatum G. subclavatum G. subclavatum G. viridilucens G. waikanaense G. xanthophyllum G. xanthophyllum G. zhujian G. zhujian Hydropus fuliginarius H. marginellus H. rugosodiscus Leucoinocybe danxiashanensis L. lishuiensis L. subglobispora MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 Voucher FFAASO0375 HMJAU59063 HMJAU59064 FA236 FA239 FA249 FFAAS0389 FFAAS0392 FFAAS0410 GDGM 93936 GDGM 94001 GDGM 87835 GDGM 92067 HMJAU59018 FLAS-F-60957 FLAS-F-71339 TENN:F-60009 TFB12519/TENN60718 TFB12783/TENN61350 DJLO5NC72 FLAS-F-60986 FLAS-F-61518 FLAS-F-71359 iNaturalist # 8545787 Mushroom Observer # 243440 S.D. Russell MycoMap # 6854 Smith-2018 iNaturalist # 17333993 DED 7822 PDD:87667 PRM 924657 SYKOf3970 FFAASO0370 FFAAS0376 S.D. Russell ONT iNaturalist # 130794969 OSC 112834 MGW1257 GDGM 80184 FFAASO115 FFAAS1034 Locality China China China Pakistan Pakistan Pakistan China China China China China China China China USA USA USA USA USA USA USA USA USA India USA India USA Brazil New Zealand Czech Republic Russia China China USA USA USA China China China GenBank accession No. nrITS OL985972 OK560883 OK560871 MN744687 MN744688 MN744686 OL985981 OL985982 OL985983 PQ452700 PQ452701 PQ452702 PQ452703 0K491123 MH016911 OR438652 KY271083 EU623640 EU623641 EU623639 MH016932 MH211945 OR242635 MN906021 MK607510 MN906138 MK573888 JQ694117 LT854023 OR915457 OL985974 OL985975 OP643427 EU669314 KY777386 MZ667478 MW424491 OR238886 nrLSU 0L985993 OL986002 OL986003 OL986004 PQ350414 PQ350415 PQ350416 OR449361 LT854023 OL985995 OL985996 EU852808 MZ667482 MW424495 OR238898 Reference Na et al. (2022) Direct Submission Direct Submission Aqdus and Khalid (2021) Aqdus and Khalid (2021) Aqdus and Khalid (2021) Na et al.(2022) Na et al.(2022) Na et al.(2022) This study This study This study This study Direct Submission Direct Submission Direct Submission Direct Submission Hughes et al. (2007) Hughes et al. (2007) Hughes et al. (2007) Direct Submission Direct Submission Direct Submission Direct Submission Direct Submission Direct Submission Direct Submission Direct Submission Direct Submission Antonin et al. (2019) Direct Submission Na et al. (2022) Na et al. (2022) Direct Submission Direct Submission Direct Submission Direct Submission Na et al. (2021) Na et al. (2024) Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China Taxon Oudemansiella aff. Marasmiellomycena pseudoomphaliiformis M. tomentosa Megacollybia clitocyboidea M. marginata M. platyphylla Mycena purpureofusca M. purpureofusca Porotheleum fimbriatum P. fimbriatum Pseudohydropus floccipes P floccipes P. globosporus Pulverulina flavoalba P flavoalba P ulmicola Trogia benghalensis T. infundibuliformis T. venenata GenBank accession No. Voucher Locality ire nae) Reference platyphylla 360-630 Japan AB509870 = Direct Submission BRNM:552721 USA OR913562 OR913566 Senanayake et al. (2023) FFAAS1036 China OR238888 OR238900 Na et al. (2024) TENN62231 USA EU623664 - Hughes et al. (2007) HR 91607 Czech LT854051 = Antonin et al. (2019) Republic BRNM 737654 Czech LT854048 LT854036 Antonin et al. (2019) Republic HMJAU43554 China MG654740 MK629356 Na and Bau (2018) HMJAU43624 China MG654741 MK629357 Na and Bau (2018) CLZhao 1120 China MH114870 7 Direct Submission Dai 12276 China KX081137 KX161656 Direct Submission BRNM 816173 Czech OM422758 0M423634 Direct Submission Republic BRNM 825631 Spain OM422760 OM423636 Consiglio et al. (2022) BAP 661 USA MH414566 MH385340 Cooper et al. (2019) FFAAS1039 China OR238891 OR238903 Na et al. (2024) FFAAS1040 China OR238892 OR238904 Na et al. (2024) TFB13871 USA M1T237476 MT237446 Matheny et al. (2020) CUH AM031 India KU647630 i Dutta et al. (2017) KUN_HKAS56709 China JQ031776 JQ031781 Yang et al. (2012) KUN_HKAS54710 China JQ031772 JQ031778 Yang et al. (2012) Phylogenetic trees were visualized using FigTree v.1.4.4. The maximum likeli- hood bootstrap over 50% (MLB = 50%) and the Bayesian posterior probability over 0.90 (BPP = 0.90) were shown. Results Molecular phylogenetic results The final concatenated dataset consisted of 95 nrlTS and 52 nrLSU sequences from 46 taxa of 11 genera of Physalacriaceae, Porotheleaceae, and Mycen- aceae, which comprised 1927 nucleotide sites (942 for nrlTS, 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 se- lected for nrlTS 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). In the phylogenetic tree (Fig. 1), Clitocybula, Gerronema, Hydropus, Leucoinocy- be, Marasmiellomycena, Megacollybia, Porotheleum, Pseudohydropus, Pulverulina, MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 244 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China 0.08 100/1 Gerronema alps da FFAAS0371 oI Gerronema keralense BKF 10263 9710.98 Gerronema keralense CAL 1666 1o~1- ~Megacollybia marginata HR_91607 62/-2 LOO/ Megacollybia play hylla BRNM 737654 Megacollybia clitocyboidea TENN62231 ipo) Gerronema confusum BJTC FM1592 Gerronema confusum BJTC FM1624 50/- Gerronema atrovirens BKF 10264 Gerronema atrovirens BKF10265 oot Gerronema indigoticum HMJAU47636 Gerronema indigoticum HMJAU47942 83/0.09 Gerronema indigoticum HMJAU47943 100/0.9 Trogia mafandtoult ormis KUN HKAS56709 LOO Trogia venenata KU KASS547T0 rogia benghalensis CUH AMO31 63/-- Gerronema nemorale FA2?36 Gerronema nemorale HMJAU59063 Gerronema nemorale FA249 Gerronema nemorale HMJAU59064 Gerronema nemorale FA2 Gerronema nemorale FFAAS0392 94/0.91N"- Gerronema nemorale FFAAS0410 Gerronema nemorale FFAAS0389 Gerronema subclavatum FLAS-F-71359 71/--| L004) Gerronema subclavatum Mushroom Observer.# 243440 Gerronema subclavatum Smith-2018 iNaturalist # 17333993 Gerronema subclavatum iNaturalist # 8545787 Gerronema subclavatum $.D, Russell EON # 6854 Gerronema subclavatum FLAS-F-6098 Gerronema subclavatum FLAS-F-61518 65/-= Gerronema brunneosquamulosum FFAAS1032 Gerronema brunneosquamulosum FFAAS1033 ogi Gerronema lapidescens GDGM 85271-] —' Gerronema lapidescens GDGM 85271-2 Gerronema lapidescens GDGM 86705 100/0.99| | 100) Gerronema angustum GDGM 88662 loo! ' Gerronema angustum GDGM 88663 lowi- Gerronema sp. HMJAU59018 Gerronema waikanaense PDD:87667 63/-- Gerronema xanthophyllum PRM 924657 PSN | Gerronema xanthophyllum SY KOf3970 78/0 erronema citrinum PCQ713 Gerronema citrinum G7785 63/-{L00y Gerronema citrinum G7458 Gerronema citrinum PC0714037 Gerronema kuruvense BKF 10266 : Gerronema kuruvense CAL 1665 ce cutRL: Gerronema kuruvense FCATAS9085 thon? Gerronema kuruvense KUC20220701_ 03 Gerronema kuruvense DCY3362(HGASMF01-15010) loo, Gerronema zhujian FFAAS037 | Gerronema zhujian FFAAS0376 ae Gerronema baishanzuense FFAAS0359 Gerronema baishanzuense FFAAS0361 of Gerronema baishanzuense FFAAS0362 —+ Gerronema baishanzuense FFAAS0363 Gerronema baishanzuense FFAAS0360 Gerronema baishanzuense FFAAS0366 100/) Gerronema pubescence GDGM 93936 Gerronema pubescence GDGM 94001 Gerronema strombodes TFB12519/TENN60718 Gerronema strombodes TFB12783/TENN61350 98/1 Gerronema strombodes FLAS-F-713 ggiit Gerronema strombodes DJILOSNC72 Gerronema strombodes TENN:F-60009 Gerronema strombodes FLAS-F-60957 DGM 87835 DGM 92067 2 93/0.95 Oudemansiella aff. platyphylla 360-630 100/1 qavarcps marginellus OSC 112834 ropus rugosodiscus 0/1 vdrop d MGW1257 } : 'ydropus puleniartes S.D. Russell ONT iNaturalist # 130794969 eikoy akan lishuiensis FFAASO115 Leucoinocybe SENSE OF FFAAS 1034 Leucoinocybe danxiashanensis GDGM80184 - Clitocybula abundans STU:SMNS-B-FU-2017/00898 Clitocybula fuscostriata FFAAS1030 Clitocybula familia PRM 921866 1o0/) Pseudohydropus floccipes BRNM 81 Pseudohydropus floccipes BRNM 82 95/0,95 : Pseudoh Grape grobosporis AP 661 tr Pulverulina flavoalba FFAAS103 Pulverulina flavoalba FFAAS1040 Pulverulina ulmicola TFB13871 Marasmiellomycena pseudoomphaliiformis BRNM:552721 Marasmiellomycena tomentosa FFAAS1036 Gerronema viridilucens DED 7822 100/1) Porotheleum fimbriatum CLZhao 1120 Porotheleum fimbriatum Dai 12276 10/1) Gerronema albidum H:6050710 Gerronema albidum H:6059277 6173 5631 cena purpureofusca HMJAU43554 cena purpureofusca HMJAU43624 SS Figure 1. Phylogenetic consensus tree of Gerronema species inferred from the maximum likelihood and Bayesian in- ference 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. MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 245 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China 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. angus- tum, 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. bais- hanzuense with a well-supported (MLB = 94%, BPP = 0.95). Gerronema rhizomor- pha 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 x 2.5-3.5 um, main- ly clavate to narrowly utriform cheilocystidia. 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). Etymology. angustum (Latin), referring to the narrow lamellae of this species. Description. Basidiomata small-sized. Pileus 10-18 mm broad, infundibuli- form, 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 x 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 x 2.5-3.5 um, L_ = 4.88 + 0.51 um, W_ = 2.96 + 0.32 um, Q = (1.33)1.43-1.83, Q_ = 1.66 + 0.18, ellipsoid to oblong, smooth, thin-walled, hyaline, guttulate, inamyloid. Basidia 18-26 x 5.5—7 um, clavate, thin-walled, hyaline, 4-spored, with sterigmata 2.3—4.4 um long. Cheilocystidia 26-45 x 6-9.5 um, clavate, fusiform to narrowly utriform, thin-walled, hyaline. Pleurocystidia not seen. Lamellar trama regular to subregular, hyphae 3-22 um wide, cylindrical, thin-walled, hyaline. Pileipellis a cutis, hyphae 1.5-24.5 um wide, smooth, hyaline; pileocystidia 22.5-65 x 8-15.5 um, oblong to utriform, thin-walled, greyish brown to light brown pigmented in KOH. Pileus trama sub- regular, sarcodimitic. Stipitipellis a cutis, hyphae 3.5-25 um wide, sometimes upturned hyphae, smooth, thin-walled, hyaline; caulocystidia 56-72 x 10.5- 20.5 um, 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 for- ests. 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). MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 246 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China Figure 2. Basidiomata of Gerronema angustum A collection GDGM 88662, holotype B collection GDGM 88663. Photo- graphed by Bin Song. Scale bars: 10 mm. Figure 3. Microscopic features of Gerronema angustum (GDGM 88662, holotype) A Basidiospores B Basidia C Cheilocys- tidia D Pileipellis E Stipitipellis. Scale bars: 10 um (A-C); 50 um (D-E). MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 247 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China 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, yel- lowish white to pale yellow lamellae are paler towards the margin, the absence of cheilocystidia, and the narrow cylindrical to utriform pleurocystidia. 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). Etymology. pubescence (Latin), referring to the species pileus usually cov- ered with pubescence when young. Description. Pileus 12-70 mm broad, hemispherical to plano-convex, de- pressed 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 i i 4 a we vl 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). MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 248 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China 15-40 x 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 x (3.5)4—4.5 pm, L_ = 7.13 + 0.57 um, W_ = 4.08 + 0.29 um, Q = (1.5)1.56-2, Qm = 1.75 + 0.18, ellipsoid to oblong, smooth, thin- walled, hyaline, guttulate, inamyloid. Basidia 24.5-39 x 5-7.5 um, clavate, thin- walled, hyaline, 2- or 4-spored, with sterigmata 2.5—5 um long. Cheilocystidia absent. Pleurocystidia 40-104 x 8.5-12.5 um, narrowly cylindrical to lageni- form, thin-walled, hyaline. Lamellar trama regular to subregular, hyphae 3.5- 26.5 um wide, thin-walled, hyaline. Pileipellis a cutis, hyphae 3-25 um wide, smooth, hyaline; pileocystidia 39-100 x 10-21 um, oblong to narrowly clav- ate, apex sometimes rostrate, thin-walled, greyish brown pigmented in KOH. Pileus trama regular to subregular, sarcodimitic. Stipitipellis a cutis, hyphae 2.5-38.5 um wide, smooth, thin-walled, hyaline; caulocystidia 38.5-84.5 x 8.5- 19 um, 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: Zhaoging 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). D OULIP EES Figure 5. Microscopic features of Gerronema pubescence (GDGM 94001, holotype) A Basidiospores B Basidia C Pleuro- cystidia D Pileipellis E Stipitipellis. Scale bars: 10 mm (A, B); 50 mm (C-E). MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 249 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China 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. 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). 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, hemi- spherical 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, subdis- tant, ventricose, even, white to yellowish white (1A1-—2), with 1-3 lamellulae. 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). MycoKeys 114; 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 250 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China Stipe 40-110 x 4-10 mm, central to eccentric, cylindrical, hollow, white to yel- lowish 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) x 5-6(6.5) um, L_ = 7.80 + 0.71 um, W_ = 5.53 + 0.42 um, Q = (1.23)1.27-1.58(1.6), Q_ = 1.41 + 0.13, broadly ellipsoid to ellip- soid, smooth, thin-walled, hyaline, guttulate, inamyloid. Basidia 25.5-32 x 7.5- 11.5 um, clavate, thin-walled, hyaline, 4-spored, with sterigmata 2.8-6 um long. Hymenial cystidia absent. Lamellar trama regular to subregular, hyphae 3-22 um wide, thin-walled, hyaline. Pileipellis a cutis, hyphae 3-25 um wide, thin- walled, light yellow in KOH; pileocystidia 25.5-63.5 x 6-14 um, narrowly utri- form to cylindrical, thin-walled, greyish brown to light brown pigmented in KOH. Pileus trama subregular, sarcodimitic. Stipitipellis a cutis, hyphae 1.5-11 um wide, smooth, thin-walled, hyaline; caulocy stidia 12.5-33 x 3.5-8 um, narrow- ly 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). ——— — IETS |15 Figure 7. Microscopic features of Gerronema rhizomorpha (GDGM 92067, holotype) A Basidiospores B Basidia C Pileipel- lis D Stipitipellis. Scale bars: 10 mm (A, B); 50 mm (C, D). MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 251 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China Discussion Morphologically, Gerronema angustum can be easily distinguished from oth- er species in Gerronema by its caespitose habit, slender basidiomata, green- ish-green pileus, narrow and close lamellae, and ellipsoid to oblong basidio- spores measuring (4)4.5-5.5 x 2.5-3.5 um. 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-con- vex 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. ker- alense 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 x 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 cheilocys- tidia 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 rhizo- morphs, 1—2-spored basidia, and abundant subcylindrical to cylindrical cheilo- cystidia (Zhao and Fan 2022). Gerronema rhizomorpha is also similar to G. atri- album (Murrill) Borovicka & Kolarik, 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 basid- ia (36-50 x 6.5-9.5 um), which mainly has 2-spored. In addition, G. atrialbum mainly grows on humus or rotting hardwood (Murrill 1913; Antonin et al. 2019). Phylogenetically (Fig. 1), our new species formed three distinct lineages ac- cording 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). How- ever, 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 cheilocys- tidia 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 MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 252 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China (MLB = 100%, BPP = 1.00) and shows that they represent the same phyloge- netic species. It is noteworthy that G. rhizomorpha with well-developed rhizo- morphs 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 misiden- tified, only ten of the 20 species reported in Asia are from China. Recent investi- gations 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 phy- logenetic relationships of Gerronema still remain highly controversial. Upon de- fining 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 rhi- ZORMO TINS ere ss Fal eect Petre cal Bic tian Metta dal tees SN een vsdede caterer vise ee ate sei 3 2 Sclerotia can be found at the base..............cccecceeeseeeseeeees G. lapidescens - Sclerotia not be found or recorded.............cccccesecesseeeeteeeees G. rhizomorpha ro mel oi: c=), 01001; Ae 1274 5) |0| = pene Pee Dee de aCe Syl ox On DMR ae fe ae ee G. indigoticum =) "BaSidiomatanwathOut IWC: TING Cis scix cee - roe hecrene hnaa Sth micdencensmenetteadterstbanteeen 4 4 Basidiomata greyish Qreen...............cccssssccsssscecssscecsssseeesesseeeees G. angustum - Basidiomata without greyish green tinge .............cccceesceeseteeeeeeeeesseeeeeeees 5 Si UBasiconiaraawiite. Saha cpnct eel bharieende sip hnwtantetuelt oi ahatnsanteee: G. albidum = riBasidiomata without White tine. cc jeccccosnccveaces can se ctenacusnanespenebebenaydenuncctosth 6 6" SPICEFOCYSTGIa PESOMiAe acho 5. acgtanseccaccteuedeneaa tt deeacial taaeaadeecene fas datien Mteaceacbe 7 Fa "SPICUPOCY SHIA ADSCMN 1 atch, oS i bear ie | Oe secede be 8 PA” MENCUS WISCIC te c- ccacte consde acai eat ensen te aene aes tee Satoneee eae eee G. chrysocarpum - Pileus dry, usually covered with pubescence when young. . G. pubescence 8 Pileus densely covered with deep brown fur or SCal@S..............:ccccceceseeees 9 = “APHBUSAWITNOUETUIOT SCALES ye sitet oon caelint pvedosvesazaecaese dered eeccnne bevcta Faversveeaebels 10 9 Stipe fuscous, basidiospores 9.0-12.9 X 4.9-7.2 UM... cccccccceeceeeteeeeeees etreaehd/SsVouh cht hereevd Sites cles ttscee Www veut piettrteed set ute ne ateres G. brunneosquamulosum - Stipe white, basidiospores 6.3-8.5 x 3.2-4.8 UM .........c eee G. zhujian 10 Basidiomata small-sized, Pileus usually < 11 mm in diam...................0 11 - Basidiomata small to medium-sized, Pileus = 11 mm in diam................. 12 11. Basidiospores 6.1-7.5 x 3.5-4.3 um, cheilocystidia present.................00 sd feet Beh Se ASR CAPR LCE Seca Pret co ANUS SiS SORA MARNE UT CM, SCb St atti G. microcarpum - Basidiospores 8-11 x 4-6 um, cheilocystidia absent............ G. kuruvense 1A SA BASHA al = 2S PON OC ves vcs adaeascevs unseeesetuneetuawenceads neat nensete G. confusum p | BasiGlia A-SOOne,, Mihi. Secs Oil unc beued Map svesatis Aes Mia ancora bee R ta maaat abel 13 MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 253 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China 13 Pileus up to 25 mm in diam, stipe 4.5—26.0 x 0.5-2.0 MM............cceeeeeees Bi, iptarte OL arated] gM een mea er or ke fe Oe te th, AOE, |W el Rits G. baishanzuense - Pileus less than 20 mm in diam, stipe 19.0-36.0 x 1.0-2.5 mm................. Mee Mick ied eNotes Hot wets at Cems wascade ents tes yecaedae Oe ues etese ee ciel 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 Acad- emy 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 Investiga- tion Program (grant no. 2022FY100500), the National Natural Science Foundation of China (grant no. 32360008), the Science and Technology Project of Forestry Admin- istration of Guangdong Province (grant no. LC-2021124), the Open Fund of Guang- dong 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. References Antonin V, Ryoo R, Shin HD (2008) Gerronema nemorale (Basidiomycota, Agaricomy- cetes): Anatomic-morphological, cultivational, enzymatic and molecular characteris- tics and its first records in the Republic of Korea. Czech Mycology 60(2): 197-212. https://doi.org/10.33585/cmy.60204 Antonin V, Borovicka J, Holec J, Beran M, Dvorak D (2011) Clitocybula familia (Fungi, Agaricales) — taxonomy, distribution, ecology and first records in the Czech Republic and Slovakia. Czech Mycology 63(1): 1-11. https://doi.org/10.33585/cmy.63101 MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 254 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China Antonin V, Borovicka J, Holec J, Piltaver A, Kolarik M (2019) Taxonomic update of Clito- cybula sensu lato with a new generic classification. Fungal Biology 123(6): 431-447. https://doi.org/10.1016/j.funbio.2019.03.004 Aqdus F, Khalid AN (2021) Gerronema nemorale: First report of the genus and species from Pakistan. Mycotaxon 136(1): 249-259. https://doi.org/10.5248/136.249 Banares A, Beltran E, Bon M (2006) Gerronema wildpretii sp. nov. (Agaricales, Basidio- mycetes) a new species from the Canary Islands. Mycologia 98(3): 455-459. https:// doi.org/10.1080/15572536.2006.11832680 Bigelow HE (1970) Omphalina in North America. Mycologia 62(1): 1-32. https://doi.org /10.1080/00275514.1970.12018941 Bigelow HE (1982) North America species of Clitocybe Part |. J. Cramer, Vaduz, 280 pp. Bigelow HE (1985) North America species of Clitocybe Part II. J. Cramer, Berlin Stutt- gart, 471 pp. Cho M, Kwon SL, Jang §S, Yoo Y, Lee SH, Kwon DY, Kim C, Lim YW, Kim JJ (2024) Notes of 15 unrecorded macrofungi in Korea. Journal of Species Research 13(1): 67-88. https://doi.org/10.12651/JSR.2024.13.1.067 Clémencon H (1982) Kompendium der Blatterpilze. Europaische omphalinoide Tricholo- mataceae. Zeitschrift fur Mykologie 48: 195-237. Consiglio G, Vizzini A, Cooper J, Marchetti M, Angelini C, Brugaletta E, Setti L (2022) The agaricoid members of the genus Porotheleum (Porotheleaceae, Agaricales), Po- rotheleum emend., Porotheleaceae s. stricto, and new genera for Agaricus floccipes and Mycena subalpina. Rivista di Micologia 64(2): 99-190. Cooper JA (2014) New species and combinations of some New Zealand agarics belong- ing to Clitopilus, Lyophyllum, Gerhardtia, Clitocybe, Hydnangium, Mycena, Rhodocolly- bia and Gerronema. Mycosphere: Journal of Fungal Biology 5(2): 263-288. https:// doi.org/10.5943/mycosphere/5/2/2 Cooper A, Desjardin DE, Perry BA (2019) The genus Mycena (Basidiomycota, Agaricales, Mycenaceae) and allied genera from Republic of SAdo Tomé and Principe, West Africa. Phytotaxa 383(1): 1-47. https://doi.org/10.11646/phytotaxa.383.1.1 Dai YC, Zhou LW, Yang ZL, Wen HA, BauT, Li TH (2010) A revised checklist of edible fungi in China. Junwu Xuebao 29(1): 1-21. Dutta AK, Nandi S, Tarafder E, Sikder R, Roy A, Acharya K (2017) Trogia benghalensis (Marasmiaceae, Basidiomycota), a new species from India. Phytotaxa 331(2): 273- 280. https://doi.org/10.11646/phytotaxa.331.2.11 Hopple Jr JS, Vilgalys R (1999) Phylogenetic relationships in the mushroom genus Co- prinus and dark-spored allies based on sequence data from the nuclear gene cod- ing for the large ribosomal subunit RNA: Divergent domains, outgroups, and mono- phyly. Molecular Phylogenetics and Evolution 13(1): 1-19. https://doi.org/10.1006/ mpev.1999.0634 Hughes KW, Petersen RH, Mata JL, Psurtseva NV, Kovalenko AE, Morozova OV, Lickey EB, Blanco JC, Lewis DP, Nagasawa E, Halling RE, Takehashi S, Aime MC, Bau T, Henkel T (2007) Megacollybia (Agaricales). Reports of the Tottori Mycological Institute 45: 1-57. Jaouen G, Sagne A, Buyck B, Decock C, Louisanna E, Manzi S, Baraloto C, Roy M, Schi- mann H (2019) Fungi of French Guiana gathered in a taxonomic, environmental and molecular dataset. Scientific Data 6(1): 206. https://doi.org/10.1038/s41597-019- 0218-z Jayawardena RS, Hyde KD, Wang S, Sun YR, Suwannarach N, Sysouphanthong P, Ab- del-Wahab MA, Abdel-Aziz FA, Abeywickrama PD, Abreu VP Armand A, Aptroot A, Bao DF, Begerow D, Bellanger JM, Bezerra JDP, Bundhun D, Calabon MS, Cao T, Cantillo T, MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 255 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China Carvalho JLVR, Chaiwan N, Chen CC, Courtecuisse R, Cui BK, Damm U, Denchev CM, Denchev TT, Deng CY, Devadatha B, de Silva NI, Dos Santos LA, Dubey NK, Dumez S, Ferdinandez HS, Firmino AL, Gafforov Y, Gajanayake AJ, Gomdola D, Gunaseelan S, He S, Htet ZH, Kaliyaperumal M, Kemler M, Kezo K, Kularathnage ND, Leonardi M, Li JP, Liao C, Liu S, Loizides M, Luangharn T, Ma J, Madrid H, Mahadevakumar S, Ma- harachchikumbura SSN, Manamgoda DS, Martin MP, Mekala N, Moreau PA, Mu YH, Pahoua P, Pem D, Pereira OL, Phonrob W, Phukhamsakda C, Raza M, Ren GC, Rinaldi AC, Rossi W, Samarakoon BC, Samarakoon MC, Sarma VV, Senanayake IC, Singh A, Souza MF, Souza-Motta CM, Spielmann AA, Su W, Tang X, Tian X, Thambugala KM, Thongklang N, Tennakoon DS, Wannathes N, Wei D, Welti S, Wijesinghe SN, Yang H, Yang Y, Yuan HS, Zhang H, Zhang J, Balasuriya A, Bhunjun CS, Bulgakov TS, Cai L, Camporesi E, Chomnunti P, Deepika YS, Doilom M, Duan WJ, Han SL, Huanraluek N, Jones EBG, Lakshmidevi N, Li Y, Lumyong S, Luo ZL, Khuna S, Kumla J, Manawasing- he IS, Mapook A, Punyaboon W, Tibpromma §S, Lu YZ, Yan J, Wang Y (2022) Fungal di- versity notes 1512-1610: Taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 117(1): 206-212. https://doi.org/10.1007/ $13225-022-00513-0 Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30(4): 772-780. https://doi.org/10.1093/molbev/mst010 Koichiro T, Glen S, Sudhir K (2021) MEGA11: Molecular Evolutionary Genetics Analysis Version 11. Molecular Biology and Evolution 7(7): 7. https://doi.org/10.1093/molbev/ msab120 Kornerup A, Wanscher JH (1978) Methuen Handbook of Colour. Eyre Methuen, London, 196-265. Kuyper TW (1986) Generic delimitation in Euripean omphalinoid Tricholomataceae. In La Famiglia delle Tricholomataceae, Italy, 83-104. https://doi.org/10.1038/nmeth.4285 Lanfear R, Frandsen PB, Wright AM, Senfeld T, Calcott B (2016) PartitionFinder 2: New methods for selecting partitioned models of evolution formolecular and morpholog- ical phylogenetic analyses. Molecular Biology and Evolution 34(3): 772-773. https:// doi.org/10.1093/molbev/msw260 Lange M (1981) Typification and delimitation of Omphalina Quél. Nordic Journal ocf. Nordic Journal of Botany 1(5): 691-696. https://doi.org/10.1111/j.1756-1051.1981. tb01424.x Latha KPD, Nanu S, Sharafudheen SA, Manimohan P (2018) Two new species of Ger- ronema (Agaricales, Basidiomycota) from Kerala State, India. Phytotaxa 364(1): 081- 091. https://doi.org/10.11646/phytotaxa.364.1.5 Li TH, Jiang QC, Xing JH, Deng WQ, Zhang M (2021) Atlas of Macrofungi in Zhongshan. Guangdong Sceience & Technolocy Press, Guangzhou, 121 pp. Liu PG (1995) Five new species of Agaricales from Southern and Southeastern Yunnan, China. Mycotaxon 6: 89-105. Liu LN, Mou GF, Bau T (2019) A new Gerronema species with striking colours from China. Phytotaxa 405(2): 074-082. https://doi.org/10.11646/phytotaxa.405.2.2 Lutzoni F (1997) Phylogeny of lichen- and non-lichen-forming omphalinoid mushrooms and the utility of testing for combinability among multiple data sets. Systematic Biol- ogy 46(3): 373-406. https://doi.org/10.1093/sysbio/46.3.373 Matheny PB, Hughes KW, Kalichman J, Lebeuf R (2020) Pulverulina, a new genus of Agaricales for Clitocybe ulmicola. Southeastern Naturalist (Steuben, ME) 19(3): 447- 459. https://doi.org/10.1656/058.019.0301 MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 256 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China Minh BQ, Nguyen MA, von Haeseler A (2013) Ultrafast approximation for phyloge- netic bootstrap. Molecular Biology and Evolution 30(5): 1188-1195. https://doi. org/10.1093/molbev/mst024 Moncalvo JM, Vilgalys R, Redhead RA, Johnson JE, James TY, Catherine Aime M, Hof- stetter V, Verduin SJUW, Larsson E, Baroni TJ, Greg Thorn R, Jacobsson S, Clemencon H, Miller Jr OK (2002) One hundred and seventeen clades of euagarics. Molecu- lar Phylogenetics and Evolution 23(3): 357-400. https://doi.org/10.1016/S1055- 7903(02)00027-1 Moser M (1983) Keys to Agarics and Boleti. The White Friars Press, Tonbridge, 535 pp. Murrill WA (1913) The Agaricaceae of the Pacific Coast-IV. New Species of Clitocybe and Melanoleuca. Mycologia 5(4): 206-223. https://doi.org/10.1080/00275514.191 3.12018520 Na Q, Bau T (2018) New species of Mycena (Mycenaceae, Agaricales) with colored lamellae and three new species records from China. Phytotaxa 361(3): 266-278. https://doi.org/10.11646/phytotaxa.361.3.2 Na Q, Hu YP, Zeng H, Ding H, Cheng XH, Ge YP, Liu ZW, Qi LL (2021) The first reported oc- currence of Leucoinocybe (Porotheleaceae, Agaricales) in China: Leucoinocybe lishui- ensis sp. nov. from Zhejiang Province. Nova Hedwigia 113(3-4): 453-469. https:// doi.org/10.1127/nova_hedwigia/2021/0661 Na Q, Hu YP, Zeng H, Ding H, Cheng XH, Ge YP, Song ZZ (2022) Updated taxonomy on Gerronema (Porotheleaceae, Agaricales) with three new taxa and one new record from China. MycoKeys 89: 87-120. https://doi.org/10.3897/mycokeys.89.79864 Na Q, Hu YP, Zeng H, Ding H, Cheng XH, Ge YP Ke BR, Zeng ZH, Liu CJ (2024) Morpho- logical and phylogenetic analyses reveal five new species of Porotheleaceae (Agar- icales, Basidiomycota) from China. MycoKeys 105: 49-95. https://doi.org/10.3897/ mycokeys.105.118826 Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Bi- ology and Evolution 32(1): 268-274. https://doi.org/10.1093/molbev/msu300 Norvell L, Redhead S, Ammirati J (1994) Omphalina sensu lato in North America 1-2. 1: Omphalina wynniae and the genus Chrysomphalina. 2: Omphalina sensu Bigelow. Mycotaxon 50: 379-407. Redhead SA (1986) Mycological observations: 17-20, nomenclatural notes on some omphaloid genera in Canada: Chrysomphalina, Rickenella, Gerronema, Omphalina. Acta Mycologica Sinica (Suppl): 297-304. Redhead SA, Moncalvo JM, Vilgalys R, Lutzoni F (2002) Phylogeny of agarics: Partial systematics solutions for bryophilous omphalinoid agarics outside of the agaricales (euagarics). Mycotaxon 82: 151-168. https://doi.org/10.1007/s12225-009-9102-x Ronquist F, Teslenko M, van der Mark P Ayres DL, Darling A, Hohna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539-542. https://doi.org/10.1093/sysbio/sys029 Senanayake C, Rossi W, Leonardi M, Weir A, McHugh MC, Rajeshkumar K, Verma RK, Karunarathna SC, Tibpromma S, Ashtekar N, Ashtamoorthy SK, Raveendran S, Kour G, Singh A, De la Pefia-Lastra S, Mateos A, Kolafik M, Antonin V, Sevéikova H, Es- teve-Raventos F, Larsson E, Pancorbo F, Moreno G, Altes A, Turégano Y, Du T-Y, Lu L, Li Q-R, Kang J-C, Gunaseelan S, Kezo K, Kaliyaperumal M, Fu J, Samarakoon MC, Gafforov Y, Teshaboeva S, Kunjan PC, Chamaparambath A, Flakus A, Etayo J, Rodri- guez-Flakus P Zhurbenko MP. de Silva NI, Tennakoon DS, Latha KPD, Manimohan MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 257 Wei-Xin Zhang et al.: Three new species of Gerronema (Agaricales, Basidiomycota) from southern China P Raj KNA, Calabon MS, Ahmadpour A, Heidarian Z, Alavi Z, Alavi F, Ghosta Y, Azizi R, Luo M, Zhao M-P, Kularathnage ND, Hua L, Yang Y-H, Liao C-F, Zhao H-J, Lestari AS, Jayasiri SC, Yu F-M, Lei L, Liu J-W, Karimi O, Tang S-M, Sun Y-R, Wang Y, Zeng M, Htet ZH, Linaldeddu BT, Alves A, Phillips AJL, Bregant C, Montecchio L, De Kesel A, Hustad VP, Miller AN, Fedosova AG, Kucera V, Raza M, Hussain M, Chen Y-P, Thi- yagaraja V, Gomdola D, Rathnayaka AR, Dissanayake AJ, Suwannarach N, Hongsanan S, Maharachchikumbura SSN, Dissanayake LS, Wijayawardene NN, Phookamsak R, Lumyong S, Jones EBG, Yapa N, Wanasinghe DN, Xie N, Doilom M, Manawasinghe IS, Liu J-K, Zhao Q, Xu B, Hyde KD, Song J (2023) Fungal diversity notes 1611-1716: Taxonomic and phylogenetic contributions on fungal genera and species emphasis in south China. Fungal Diversity 122(1): 161-403. https://doi.org/10.1007/s13225- 023-00523-6 Singer R (1951) New genera of fungi. V. Mycologia 43(5): 598-604. https://doi.org/10.1 080/00275514.1951.12024157 Singer R (1962) Diagnoses fungorum novorum Agaricalium II. Sydowia 15: 45-83. Singer R (1964) Die Gattung Gerronema. Nova Hedwigia 7: 53-92. Singer R (1970) Omphalinae (Clitocybeae-Tricholomataceae, Basidiomycetes). Organi- zation for Flora Neotropica, University of California, 131 pp. Singer R (1986) The Agaricales in modern taxonomy, 4" edn. Koeltz Scientific Books, KOnigstein, 981 pp. Vizzini A, Picillo B, Luigi P Dovana F (2019) Chrysomtcena perplexa gen. et sp. nov. (Agaricales, Porotheleaceae), a new entity from the Lazio region. Rivista Micologica Romana 107: 96-107. White TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. PCR protocols, a guide to methods and ap- plications. Academic Press, 315-322. https://doi.org/10.1016/B978-0-12-3721 80- 8.50042-1 Wu LQ, Xu JY, Zhang M, Chen ZX, Liu ZX, Li TH (2021) Macrofungal Diversity in Danx- iashan National Nature Reserve and Four Rew Records in China. Acta Edulis Fungi 37(3): 135-146. https://doi.org/10.16488/j.cnki.1005-9873 Yang ZL, Li YC, Tang LP Shi GQ, Zeng G (2012) Trogia venenata (Agaricales), a novel poisonous species which has caused hundreds of deaths in southwestern China. Mycological Progress 11(4): 937-945. https://doi.org/10.1007/s11557-012-0809-y Zhang D, Gao F, Jakovlié |, Zou H, Zhang J, Li WX, Wang GT (2020) PhyloSuite: An in- tegrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Molecular Ecology Resources 20(1): 348-355. https://doi.org/10.1111/1755-0998.13096 Zhang M, Xie DC, Wang CQ, Deng WQ, Li TH (2022) New insights into the genus Gyropo- rus (Gyroporaceae, Boletales), with establishment of four new sections and descrip- tion of five new species from China. Mycology 13(3): 223-242. https://doi.org/10.10 80/21501203.2022.2094012 Zhang WX, Wang CQ, Deng WQ, Chang CQ, Zhang M (2024) A taxonomic revision of rare medicinal fungus Leiwan. Acta Edulis Fungi 31(3): 90-99. Zhao TY, Fan L (2022) Gerronema confusum (Porotheleaceae), a new species from North China. Phytotaxa 550(2): 186-194. https://doi.org/10.11646/phytotaxa.550.2.7 MycoKeys 114: 239-258 (2025), DOI: 10.3897/mycokeys.114.145299 258