683 MycoKeys MycoKeys 120: 317-338 (2025) DOI: 10.3897/mycokeys.120.157997 Research Article Four new species of Marasmius subgenus Globulares (Marasmiaceae, Agaricales) from subtropical regions of China Hong Chen™®, Yu-Qin Xu'™®, Hui Zeng2®, Ya-Ping Hu?®, Sheng-Nan Wang"™, Jun-Qing Yan'® 1 Jiangxi Provincial Key Laboratory of Excavation and Utilization of Agricultural Microorganisms, Jiangxi Agricultural University, Nanchang 330045, China 2 Institute of Edible mushroom, Fujian Academy of Agricultural Sciences, Fuzhou 350011, China 3 Nanjing Institute of Environmental Sciences, Ministry of Ecology and Environment Mountains, Nanjing 210042, China Corresponding author: Jun-Qing Yan (yanjunqing1990@jxau.edu.cn) OPEN Qrceess Academic editor: Maria P. Martin Received: 6 May 2025 Accepted: 25 July 2025 Published: 15 August 2025 Citation: Chen H, Xu Y-Q, Zeng H, Hu Y-P Wang S-N, Yan J-Q (2025) Four new species of Marasmius subgenus Globulares (Marasmiaceae, Agaricales) from subtropical regions of China. MycoKeys 120: 317-338. https://doi.org/10.3897/ mycokeys.120.157997 Copyright: © Hong Chen 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 Marasmius is a large genus in Agaricales, exhibiting rich species diversity and a wide distribution. In this study, four new species of Marasmius belonging to the subgenus Globulares were identified in subtropical regions of China. Morphologically, M. blandus is characterized by a light orange pileus, with a non-striate surface, medium-sized basidiospores, and subfusiform to narrowly utriform pleurocystidia; M. xingshanensis is identified by a brown pileus, variably shaped pleurocystidia, and absence of cheilocystidia; M. vulgaris is identified by large basidiospores, which are up to 16.0 um long, narrowly fusiform to lageniform pleurocystidia, often with capitate, papillate, or constricted moniliform structures at apices; M. subpurpureostriatus is recognized by a grayish-green pileus with deeply sulcate violet striae, lamellae distant, and clavate to fusoid-clavate basidiospores, which are up to 22.0 um long. The distinct taxonomic status of these four new species was confirmed by their positions in the 4-locus (ITS, LSU, tef-1a, rpb2) phylogenetic trees. Detailed descriptions and morphological photographs of four new species are provided in this paper. Key words: Basidiomycetes, new taxa, phylogeny, taxonomy Introduction Marasmius Fr., with M. rotula (Scop.) Fr. designated as the type species, was es- tablished by Fries (1835). According to the Index Fungorum, the genus contains 2,083 records and comprises approximately 700 species (Oliveira et al. 2024). It is characterized by the small to medium-sized basidiomata with convex to campanulate pilei; adnate to adnexed, or free lamellae that may be collariate; an insititious or non-insititious stipe; white spore print; inamyloid, smooth, hy- aline, thin-walled basidiospores; a hymeniform pileipellis. It typically grows on soil, dead branches, or leaf litter in forests and is distributed worldwide, with high diversity in tropical and subtropical areas (Retnowati and Desjardin 2022). Singer classified Marasmius into twelve sections based on cortical struc- ture and spore characteristics (Singer 1958; Singer 1976; Singer 1986). This classification was widely accepted until the end of the last century. 317 Hong Chen et al.: Four new species of Marasmius subgenus G/obulares from China Recent molecular phylogenetic studies have shown that Marasmius is poly- phyletic (Wilson and Desjardin 2005). By analyzing nrlTS and nLSU rDNA se- quences, Wilson and Desjardin (2005), Wannathes et al. (2009a), Tan et al. (2009), and Jenkinson et al. (2014) restricted the genus to a monophyletic lineage containing only the five sections defined by Singer, including the sec- tions Marasmius, Globulares, Sicci, Leveilleani and Neosessiles. Other sections recognized by Singer are currently excluded from Marasmius. Subsequently, Oliveira et al. (2024) proposed two subgenera — Marasmius and Globulares, based on morphological and phylogenetic analysis of multilocus (SSU, LSU, ITS, rpb2 and tef-1a). The subgenus Marasmius is characterized by thin basid- iomata with an insititious stipe, inamyloid or dextrinoid trama, and includes sect. Crinis-eques, sect. Marasmius, sect. Sanguirotales, sect. Variabilispori, and sect. Sicciformes. The subgenus Globulares is characterized by a great variety of basidiomata, stipe with a basal mycelium (non-insititious), exclu- sively dextrinoid trama, and includes sect. Globulares and sect. Sicci. Macrofungal species diversity in China is remarkably rich, with the sub- tropical regions emerging as hotspots for biodiversity research due to their complex topography and climatic conditions. In recent years, at least thir- teen new species within Marasmius have been discovered in China (Deng and Li 2011; Deng et al. 2011; Deng et al. 2012; Yang et al. 2013; Deng et al. 2015; Wang et al. 2015; Deng et al. 2017; Liang et al. 2017; Li et al. 2023; Zhang et al. 2023). During biodiversity surveys in China’s subtropical regions, we discovered four new species within the subgenus G/obulares. This study provides a detailed elucidation of the newly discovered species through morphological comparisons and phylogenetic analyses. Materials and methods Morphological studies Specimens were collected from Hubei, Jiangxi, and Zhejiang provinces of China between 2019 and 2024 and were deposited in the Herbarium of Fungi, Jiangxi Agricultural University (HFJAU). Macroscopic characteristics were recorded from fresh specimens. Color codes are from Kornerup and Wanscher (1978). Microscopic features were described from dried material mounted in H,O, 5% aqueous KOH, Melzer’s reagent and Congo Red using an Olympus BX-53 microscope (Olympus Corporation, Tokyo, Japan) (Horak 2005). For each collection, at least 40 basidiospores, basidia, and cystidia were measured. The range of spore size is expressed in the form (a) b-c (d), in which “a” and “d” represent the minimum and maximum values; 90% of the spores fall within the range “b-c”. The meanings of the other spore characteristics are as follows: “Q” stands for the ratio of length to width (Chen et al. 2024; Yan et al. 2024). DNA extraction, PCR amplification, and sequencing Genomic DNA was extracted from dried specimens using the NuClean Plant Genomic DNA kit (CWBIO, China) (Hopple and Vilgalys 1999; Wang et al. 2022). The ITS, LSU, tef-7a, and rpb2 regions were amplified using the respective MycoKkeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 318 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China primer pairs of ITS1F/ITS4, LROR/LR7, EF983F/EF1567R, and rpb2-6F/rpb2- 7.1R (Hopple and Vilgalys 1999; Sanchez-Ramirez et al. 2014). PCR amplification was conducted in a 25 uL reaction system as follows: 1 uL DNA, 2 uL primers, 9.5 pL ddH,0, and 12.5 pL 2x Taq Master Mix (Dye Plus). For ITS, PCR was carried out using a touchdown amplification procedure: initial 95 °C for 5 min, and then 14 cycles of denaturing at 95 °C for 30 s, annealing at 65 °C for 45s (-1 °C per cycle), extension at 72 °C for 1 min, and then 30 cycles of denaturing at 95 °C for 30 s, annealing at 52 °C for 30 s, and extension at 72 °C for 1 min, with the final extension at 72 °C for 10 min (Yan et al. 2022). For the other regions, the procedure was initial 98 °C for 5 min, and then 8 cycles of denaturing at 98 °C for 5 s, annealing at 61 °C for 40 s (-1 °C per cycle), exten- sion at 72 °C for 2 min, and then 35 cycles of denaturing at 98 °C for 5s, anneal- ing at 54 °C for 1.5 min, extension at 72 °C for 2 min, with the final extension at 72 °C for 10 min (Sanchez-Ramirez et al. 2014; Chen et al. 2024). The PCR prod- ucts were sequenced by Qing Ke Biotechnology Co. Ltd. (Wuhan City, China). Alignment and phylogenetic analyses Sequence reads were assembled and edited using Sequencher v.5.4 and were deposited in GenBank (GB) database. Based on the research by Oliveira et al. (2024), and the similarity of these new species to the most closely related se- quences identified in the BLAST results of ITS, a total of 187 sequences, in- cluding 78 ITS, 55 LSU, 27 tef-1a and 27 rpb2 regions, were used in subsequent analyses. Four species of Marasmius subg. Marasmius were designated as out- groups. Details are presented in Table 1. Sequence datasets, containing intron regions, were separately aligned using MAFFT v.7 (Katoh and Standley 2013). BI and ML phylogenetic analyses of the processed sequences were run using Mrbayes v.3.2.7a and IQ-TREE v.2.1.2, respectively (Ronquist and Huelsenbeck 2003; Nguyen et al. 2014). The best-fit models for BI were determined by PartitionFinder, complying with Corrected Akaike information criterion (AlCc) (Lanfear et al. 2017). For the ML analysis, 10000 replicates are performed based on the ultrafast bootstrap option of ML that allowed partitions from different seeds. For the BI analysis, the Markov chains were run for 1 million generations. The first 25% of trees were discarded as burn-in. Branches with Bayesian posterior probability (BI-PP) = 0.95 and ML bootstrap support (ML-BP) = 75% are shown in the tree (Fig. 1). The designation of the clade names is based on the study by Oliveira et al. (2024). Results Phylogenetic analysis A total of 4086 characters from 78 taxa were used in phylogenetic analyses (ITS 931 bp; LSU 1328 bp; tef-7a 1081 bp; rpb2 746 bp), of which 2673 were constant, 1135 were parsimony-informative, and 278 were singleton. The best- fit models of ML: HKY + F + G4 for ITS, TIM3e +1 + G4 for LSU, TIM2e + 1 + G4 for tef-1a and rpb2. The best-fit models of BI: HKY + F + G4 for ITS, SYM +1 + G4 for LSU, SYM +1 + G4 for tef-7a and rpb2. For Bayes analysis, the average standard deviation of split frequencies less than 0.01 after 610000 generations. Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 319 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China Table 1. Details of sequences used in the phylogenetic analyses. spews720r(rloype) | KPIZ7E7A KPIDTETE Species Marasmius albopurpureus M. albopurpureus M. auranticapitatus MC4554(holotype) ON502670 | ONSO27345 | M. avellaneus FK1616(holotype) M. cohaerens var. lachnophyllus -—-DED4071——s«|s«OROBE637 | ORESENEZ M. cohaerens var. mandshuricus —-LE295991 |: KF774167 | KF896247,— | M. cohaerens var. mandshuricus -—-LE295986(holotype) | KF774171 | KF896245 M. confertus var. tenuicystidiatus ~- BRNM718808 =| HQ607374. HQ607375, | M. confertus var. tenuicystidiatus —-HMAS 293268 =| OR236975 M. fusicystidiosus BRNM 714567(holotype) Fugi7624 | Fu936144 | | M. galbinus GDGM27251 (holotype) | HQ709445, M. gracilis JO90(holotype) M. grandiviridis NW152(holotype) | OR636648 | OR6S6977, M. indopurpureostriatu KD 14-001(holotype) 7777 ee M. nivicola KPM-NC 0006038 FJ904973 | FJ904955 M. odoratus CAL:1264(holotype) |ktigoz32{ | | M. oreades __NNos5694 | JN943604 | JNGATI44 | | 0031099 | M. oreades Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 Reference Wang et al. (2015) Wang et al. (2015) Oliveira et al. (2022) Oliveira et al. (2022) Oliveira et al. (2020a) Shay et al. (2017) this study this study this study this study Wannathes et al. (2009a) Wannathes et al. (2009a) Antonin et al. (2010a) Liang et al. (2017) Antonin (2003) Oliveira et al. (2022) Antonin et al. (2010b) Antonin et al. (2010b) Oliveira et al. (2024) Kiyashko et al. (2014) Kiyashko et al. (2014) Antonin et al. (2011) Antonin et al. (2011) Oliveira et al. (2024) Oliveira et al. (2024) Oliveira et al. (2024) Oliveira et al. (2020b) Oliveira et al. (2020b) Oliveira et al. (2024) Antonin et al. (2010a) Deng and Li (2011) Oliveira et al. (2020a) Wannathes et al. (2009a) Wannathes et al. (2009a) Oliveira et al. (2022) Dutta et al. (2015) Kiyashko et al. (2014) Kiyashko et al. (2014) Magnago et al. (2016) Magnago et al. (2016) Antonin et al. (2010b) Antonin et al. (2010b) Wannathes et al. (2009b) Kiyashko et al. (2014) Oliveira et al. (2024) Antonin et al. (2010b) Antonin et al. (2010b) Farook and Manimohan (2015) Kotowski et al. (2019) unpublished 320 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China Species M. pinicola M. pinicola M. pseudoconfertus M. pseudopurpureostriatus M. puerariae M. purpureostriatus M. rhabarbarinus M. rhabarbarinus M. rhabarbarinoides M. roseus M. rotula M. rubicundus M. silvicola M. silvicola M. silvicola M. spegazzinii M. subpurpureostriatus M. subvigintifolius M. vulgaris M. vulgaris M. vulgaris M. vulgaris M. vulgaris M. vulgaris M. wynneae M. xingshanensis M. xingshanensis M. xingshanensis Voucher Li20220706-13 Li20220706-15(holotype) HKAS 49088(holotype) NW286(holotype) R Kirschner & C-J Chen 2139 (type) NW158 J0494 J0457 J066 J0352(holotype) AW274 J0464(holotype) J0357 J0362 J0366 J0467 HFJAU4740(holotype) J0242(holotype) HFJAU0904 HFJAU1901 HFJAU1976 HFJAU2719 HFJAU2748 HFJAU2875(holotype) HCCN-G86 HFJAU5S344 HFJAU5540 HFJAU5544(holotype) ITS 0Q941785 0Q941786 HQ832733 OR636672 JX470333 EU935539 KP635192 KP635191 KP635190 ON502678 MN714042 ON502658 KP635194 KP635195 KP635196 KP635197 PV363556 MN714035 PV363550 PV363551 PV363552 PV363553 PV363554 PV363555 FJ904979 PV363547 PV363548 PV363549 LSU OR657016 JX470332 KP635147 KP635146 KP635145 ONS02745 OR657020 ON5S02728 KP635149 KP635150 KP635151 PV363575 OR657031 PV363571 PV363572 PV363574 FJ904961 PV363577 PV363578 PV363579 tef-1a PP026070 PP026069 PP026068 ON553954 PP026113 ON553948 PP026060 PP026061 PP026075 PV390073 PP026132 PV390070 PV390071 PV390074 PV390075 PV390076 rpb2 OR896451 OR896450 ON553944 OR896415 ON553937 OR896485 OR896486 OR896488 PV394695 OR896411 PV394688 PV394689 PV394690 PV394692 PV394693 PV394697 Reference Li et al. (2023) Li et al. (2023) Deng et al. (2011) Wannathes et al. (2009a) Kirschner et al. (2013) Wannathes et al. (2009a) Oliveira et al. (2020b) Oliveira et al. (2020b) Oliveira et al. (2020b) Oliveira et al. (2022) Oliveira et al. (2020a) Oliveira et al. (2022) Oliveira et al. (2024) Oliveira et al. (2020b) Oliveira et al. (2020b) Wannathes et al. (2009a) this study Oliveira et al. (2020a) this study this study this study this study this study this study Antonin et al. (2010b) this study this study this study The results of the phylogenetic analysis are shown in Fig. 1. Four new spe- cies respectively formed distinct and stable branches. Marasmius blandus belongs to /brunneospermus clade and groups together with M. brunneosper- mus Har. Takah. and M. fusicystidiosus Antonin, Ryoo & H.D. Shin (BI-PP = 1, ML-BP = 100%). Marasmius xingshanensis belongs to /brunneospermus clade and groups together with M. macrocystidiosus Kiyashko & E.F. Malysheva. and M. magnus A.C. Magnago & J.S. Oliveira (BI-PP = 1, ML-BP = 98%). Marasmius vulgaris belongs to /confertus clade and is closely related to M. graminipes Wannathes, Desjardin & Lumyong, but this grouping has unstable support in Bayesian analysis. Marasmius subpurpureostriatus belongs to /purpureostria- tus clade and is closely related to M. purpureostriatus Hongo. Taxonomy Marasmius blandus J.Q. Yan & H. Chen, sp. nov. MycoBank No: 858718 Fig. 2 Etymology. “blandus” refers to its smooth and non-striate pileus. Holotype. CHINA * JiangXi Province, Jiangxi Agricultural University, 6 October 2020, collected by Jia-Yue Sun, HFJAU2362. Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 321 Hong Chen et al.: Four new species of Marasmius subgenus G/obulares from China -/92, Marasmius xingshanensis China HFJAU5344 1/99 Marasmius xingshanensis China HFJAU5544(holotype) Marasmius xingshanensis China HFJAU5540 00; Marasmius macrocystidiosus Pakistan KP-13 Marasmius macrocystidiosus Russian LE 295996(holotype) 1/100; Marasmius magnus Magnago AC 1001 (holotype) Marasmius magnus Canada NP 514 Marasmius blandus China HEJAU&046 arasmius blandus China Marasmius blandus China HFJAU2362(holotype) /b runneospermus Marasmius blandus China HFJAUS635 Ser. Brunneospermi Marasmius brunneospermus Japan KPM-NC 0005011 (holotype) Marasmius fusicystidiosus Korea BRNM 714567(holotype) -/98 Marasmius silvicola Brazil JO362 Marasmius silvicola Brazil JO366 Marasmius silvicola Brazil JO357 /silvicol 1/100 Marasmius nigrodiscus USA TENN 93556 S SH eee a arasmius nigrodiscus er. Silvicolae 1/100; Marasmius maximus Korea BRNM 714571 -/1001' Marasmius maximus Korea BRNM 714671 Marasmius wynneae Korea HCCN-G86 00; Marasmius nivicola Korea BRNM 714573 Marasmius nivicola Japan KPM-NC 0006038 (holotype) 0.95/97, -/9] 1/99 1/100 P-P8/? Marasmius oreades Denmark NN055694 Marasmius oreades China ZRL2015086 / wynneae Marasmius decipiens USA DED4146 Ser. Wynnearum 1/100; Marasmius cohaerens var. cohaerens Korea BRNM652833 1/10 Marasmius cohaerens var. cohaerens Korea BRNM695761 1/10 Marasmius cohaerens var. mandshuricus Russia LE 295991 1/100— Marasmius cohaerens var. mandshuricus Russia LE BSR NPE) /cohaerens Marasmius cohaerens var. lachnophyllus USA DED4071 r. Cohaerentes 1/100; Marasmius albopurpureus China GDGM57089 | ' Marasmius albopurpureus China ee ey ie Marasmius pseudopurpureostriatus Thailand NW286(holotype) Marasmius purpureostriatus Thailand NW158 Marasmius SUDB LIRA TE OS ria as China HFJAU4740(holotype) Marasmius grandiviridis Thailand NW152(holotype) Marasmius odoratus India CAL: 12oaGiolotype) Marasmius puerariae TaiWan R Kirschner & C-J Chen 2139 (TNM)(holotype) Marasmius indopurpureostriatu India KD 14-001(holotype Marasmius campestris Hainan, China HKAS 80857(holotype) 1/100' Marasmius campestris Hainan, China HKAS 80858 Marasmius galbinus China GDGM27251 (holotype) Marasmius bekolacongoli USA Lockwood2131638 0.97/80 /purpureostriatus Marasmius mokfaensis Thailand DED7726(holotype) mas Marasmius decipiens USA DED3612 Ser. Purpureostriati 1/98 Marasmius vulgaris China HFJAU1976 Marasmius vulgaris China HFJAU2875(holotype) Marasmius vulgaris China HFJAU1901 Marasmius vulgaris China HFJAU0904 Marasmius vulgaris China HEFJAU2748 Marasmius vulgaris China HFJAU2719 Marasmius graminipes Thailand A eat Marasmius confertus var. tenuicystidiatus Korea M 718808 Marasmius confertus var. tenuicystidiatus China HMAS 293268 -/95; Marasmius bondoi Thailand NW390 Marasmius bondoi Thailand NW3 eal a Marasmius pseudoconfertus China H 49088(holotype) Marasmius cystidiatus Canada DED4594 Marasmius floridanus Canada DEDS164 Marasmius spegazzinii Brazil JO467 7100 Marasmius pinicola China Li20220706-13(holotype) Oe LY 4/97) 1/100" Marasmius pinicola China Li - olotype : Meee altoribeirensis Brazil Tos3? i Ser. Conferti 1/100; Marasmius rhabarbarinus Brazil JO494 yi aey Pia IaR Saas, Brae J WoLe farasmius rhabarbarinoides Brazi * /100; seule dimorphus Brazil JO298 /rhabarbarinus arasmius dimorphus Brazil JO334 Ser. Rhabarbarini 1/97, Marasmius EET ae TEI DAITS Brazil JO523 (holotype) -/85| — Marasmius roseus Brazil Meee ) 1/100|' Marasmius haematocephalus Brazil JO5 Sect. Globulares 1/99 Fert Marasmius auranticapitatus Brazil MC4554(holotype Sect. Sicci NEEL Weenie inns cielo ote) EET Ta6a holoty pep ype) -/82 Marasmius gracilis Brazil JO90(holotype) Marasmius rotula Canada AW274 ; 1/100 Marasmius subvigintifolius Brazil JO242(holotype) | Outgroup Subg. Marasmius Marasmius avellaneus Brazil FK1616(holotype) Figure 1. Phylogram of Marasmius generated by Bayesian inference (BI) analysis based on sequences of ITS + LSU + tef-1a + rpb2. It was rooted with M. avellaneus, M. gracilis, M. rotula and M. subvigintifolius. Bayesian inference (BI-PP) = 0.95 and ML bootstrap proportions (ML-BP) = 75 are indicated as PP/BP. The new taxa are marked in bold. Diagnosis. Marasmius blandus is mainly characterized by the rather small basidiomata, light orange pileus, with a non-striate surface; basidiospores mainly shorter than 7.5 um; variably shaped pleurocystidia, subfusiform, nar- rowly utriform, with a short or long papilla at the apex and rarely with nodulose on the surface; cheilocystidia clavate, subfusiform, apex obtuse, rarely with short papilla or branched. It differs from M. brunneospermus by having smaller cheilocystidia, which are mainly shorter than 30 um in length. MycoKkeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 322 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China Figure 2. Morphological structures of M. blandus. A-C. Basidiomata; D. Basidiospores; E. Pileipellis; F-J. Pleurocystidia; K-M. Cheilocystidia. All microscopic structures were observed in 5% KOH, and used 1% Congo red. Scale bars: 20 mm (A-C); 10 um (D); 30 um (E-J); 20 um (K-M). Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 323 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China Description. Pileus 25-50 mm, plano-convex to plane, with or without a slight obtuse umbo at the center, smooth, non-striate, hygrophanous, light orange (6A5-6) at center, slightly paler to white towards margin, drying out to white, the center and the margin with pale brown. Context thin, white. Lamel- lae 3.0-5.0 mm broad, adnexed, ventricose, subdistant, white with slightly brown, with 2-3 tiers of lamellules, edges even, concolorous. Stipe 20-50 mm long, 2.0-3.0 mm thick, central, cylindric, equal, fibrous, hollow, light yellow (443-4), becoming reddish brown (8D5-6) as stipe dries, smooth, apex velu- tinous, and the base covered with white mycelium. Basidiospores (5.0)5.5-7.5(8.0) x 2.4-3.5 um (av = 6.5 x 3.0 um), Q = (1.6)1.8-2.6(3.0), elongated-ellipsoid to cylindrical, slightly flattened on one side in profile, 2.4-4.0 um broad, elongated-ellipsoid to cylindrical in face view, smooth, colorless, hyaline, inamyloid, thin-walled. Basidia 24.0-30.5 x 4.0-6.5 um, clavate, 4-spored. Pleurocystidia 37.0-80.0 x 7.5-18.0 um, variable, subfusiform, narrowly utriform, apex with short or long papilla, sur- face rarely with nodulose, smooth, slightly thick-walled, yellowish in 5% KOH. Cheilocystidia 18.5-31.0(38.0) x 5.0-15.5 um, variable, clavate, subfusiform, apex obtuse, rarely with short papilla or branched, smooth, thin-walled. Pile- ipellis a hymeniderm composed of cells 16.5-31.5 x 7.0-16.0 um, pyriform or broadly clavate, smooth, hyaline, thin-walled. Lamellae trama interwoven, with hyphae 4.0-6.5 um in diam, hyaline, dextrinoid, thin-walled. Stipitipellis a cutis composed of cylindrical hyphae, 4.0-9.5 um wide, parallel, smooth. Caulocystidia absent. Clamp connections present. Habitat. Scattered on soil in broad-leaved forest or mixed coniferous and broad-leaved forests. Additional specimens examined. CHINA + JiangXi Province, Jiangxi Agri- cultural University, 1 May 2019, collected by Jia-Yue Sun, HFJAU3367; 23 May 2023, collected by Lin-Gen Chen, Cheng-Feng Nie HFJAU4946; Hubei Province, Xingshan County, Yichang City, 23 July 2024, collected by Jun-Qing Yan, Lin- Gen Chen, Hong Chen, Ling Ding, HFJAU5635. Note. Based on molecular systematics and morphological analysis, M. blan- dus belongs to subg. Globulares ser. Brunneospermi (Oliveira et al. 2020b; Ol- iveira et al. 2024), within this series, M. blandus is morphologically similar to M. brunneospermus, M. fusicystidiosus, and M. macrocystidiosus. However, M. brunneospermus has an irregularly wrinkled to rugulose reticulate pileus and larger cheilocystidia (30-57 x 4-13 um) (Takahashi 1999); M. fusicystidiosus differs in the reddish ochre-brown pileus at the center, slightly paler towards margin and larger basidiospores (8.5—10 x 3.5—4.0 um) (Antonin et al. 2010a); M. macrocystidiosus is distinguished from M. blandus by a light brown or gray- ish brown pileus, larger basidiospores (6.9-10.5 x 3.3—4.4 um) and larger pleu- rocystidia (78.3-123.0 x 12.5-13.8 tm) (Kiyashko et al. 2014). In addition, morphologically, among the known species of sect. Globulares, only M. desjardinii K. Das, Antonin & D. Chakr., M. muramwyanensis Antonin, M. goossensiae Beeli, and M. phlebodiscus Desjardin & E. Horak. share simi- lar morphological characteristics with the new species M. blandus, including a hymenidermal pileipellis composed of Globulares-type cells, the basidiospores range in size from 5.0-8.0 x 2.0-4.0 um, and have the well-developed pleu- rocystidia. However, M. desjardinii differs from M. blandus by a longer stipe (70-180 x 4-10 mm), grayish orange to apricot pileus when dry, pleurocystidia Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 394 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China sometimes lageniform and possesses pileocystidia (Das et al. 2019); M. mu- ramwyanensis differs by having a stipe white at apex, ochraceous to brown- ish-orange at base, pleurocystidia cylindrical, clavate, fusoid (Antonin 2003); M. goossensiae differs from M. blandus in forming a cream colored pileus with fuligineous or ochraceous brown centre, clavate, and rostrate pleurocys- tidia (Antonin 2007); M. phlebodiscus has a pale beige to tan colored pileus with reticulate wrinkles at center and only fusoid pleurocystidia, fusoid and mucronate cheilocystidia (Petrini et al. 1997). Marasmius xingshanensis J.Q. Yan & H. Chen, sp. nov. MycoBank No: 858719 Fig. 3 Etymology. “xingshanensis” refers to its type specimen originating from the Xingshan County of China. Holotype. CHINA * Hubei Province, Xingshan County, Yichang City, 29 June 2024, collected by Jun-Qing Yan, Lin-Gen Chen, Hong Chen, Ling Ding, HFJAU5544. Diagnosis. Marasmius xingshanensis is mainly characterized by the rather small basidiomata, pileus dark brown, reddish brown at center, slightly paler to white towards margin; lamellae rarely forked; basidiospores mainly shorter than 7.0 um; well-developed pleurocystidia, up to 100 um long, subcylindrical, ventricose, narrowly utriform, apex with obtuse or short papilla, the base is con- stricted into a curved long or short stipe; lamellae edge is composed of a large number of basidioles and rarely basidia; cheilocystidia absent. It differs from M. riparius Singer by having smaller spores which are shorter than 7 um. Description. Pileus 15-30 mm, plano-convex to plane, center with slightly ob- tuse umbo, smooth, hygrophanous, dark brown (7F7), reddish brown (8D5-6) at center, slightly paler to white towards margin, striate up to 1/3 from the margin, drying out to white, with grayish orange at center. Context thin, white. Lamellae 2.0-3.0 mm broad, adnexed, ventricose, moderately close, grayish red (7B3), light brown (7D5), with 2-3 tiers of lamellules, rarely forked, edges even, white. Stipe 20-41 mm long, 1.5-3.0 mm thick, central, cylindric, equal, fibrous, hollow, gray- ish red (7B3), light brown (7D5), smooth, the base covered with white mycelium. Basidiospores (5.0)5.5—7.0(7.5) x 2.2-3.5 um (av = 6.3 x 2.9 um), Q =(1.6)1.8- 2.7(3.0), elongated-ellipsoid to cylindrical, slightly flattened on one side in pro- file, 2.3-3.5 um broad, elongated-ellipsoid to cylindrical in face view, smooth, colorless, hyaline, inamyloid, thin-walled. Basidia 24.5-33.5 x 4.0-6.5 um, clav- ate, 4-spored. Pleurocystidia (43.0)53.0-90.0(100.0) x (5.7)7.0-16.0 um, vari- ously, subcylindrical, ventricose, narrowly utriform, rarely branched, apex obtuse or with short papilla, the base is constricted into a curved long or short stipe, surface rarely with nodulose, smooth, slightly thick-walled, yellowish in 5% KOH. Lamellae edge is composed of a large number of basidioles and rarely basidia, cheilocystidia absent. Pileipellis a hymeniderm composed of cells 19.0-38.0 x 7.5-18.0 um, pyriform or broadly clavate, smooth, hyaline, thin-walled. Lamel- lae trama interwoven, with hyphae 5.0-8.8 um in diam, hyaline, dextrinoid, thin- walled. Stipitipellis a cutis composed of cylindrical hyphae 4.0-8.5 um wide, parallel, smooth. Caulocystidia absent. Clamp connections present. Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 325 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China } , : LE oo ee BY : a F ‘ es “ et ee a ee ee Figure 3. Morphological structures of M. xingshanensis. A, B. Basidiomata; C. Basidiospores; D. Lamellar edge; E-N. Pleurocystidia; O. Pileipellis. All microscopic structures were observed in 5% KOH, structures of C and E-O were stained by 1% Congo red. Scale bars: 20 mm (A, B); 10 um (C); 30 um (D-N); 20 um (0). Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 326 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China Habitat. Scattered on soil in broad-leaved forest or mixed coniferous and broad-leaved forests. Additional specimens examined. CHINA * Hubei Province, Xingshan County, Yichang City, 29 June 2024, collected by Jun-Qing Yan, Lin-Gen Chen, Hong Chen, Ling Ding, HFJAU5344; 4 July 2024 collected by Jun-Qing Yan, Lin-Gen Chen, Hong Chen, Ling Ding, HFJAU5540. Note. Based on molecular systematics and morphological analysis, M. xing- shanensis belongs to subg. Globulares ser. Brunneospermi (Oliveira et al. 2020b; Oliveira et al. 2024). Within this series, only M. magnus lacks cheilocystidia. However, M. magnus has a larger pileus (31-122 mm), fulvous to rusty orange in color, a longer stipe (70-94 mm), and a pileipellis composed of Siccus-type broom cells (Magnago et al. 2016). Morphologically, among the known species of sect. Globulares, only M. riparius and M. ochraceus Berk. & Broome. share similar morphological characteristics with the new species M. xingshanensis, including a hymenidermal pileipellis com- posed of Globulares-type cells, well-developed pleurocystidia, and lack of cheilo- cystidia. However, M. riparius has a cinnamon pileus, with larger basidiospores (8.2-10x4.8-5.5 um), smaller pleurocystidia (40—44 x 8-9 um) (Singer and Digilio 1952); M. ochraceus has a larger pileus (30-80 mm), a longer stipe (60-110 mm) and smaller pleurocystidia (30-35 x 6-8 um) (Berkeley and Broome 1873). In addition, morphologically, M. pallidibrunneus J.S. Oliveira, and M. pinicola Jing Si, S.H. He & Hai J. Li have the aspect of M. xingshanensis with similarly basidioma size and colored pileus. However, M. pallidibrunneus has basidio- spores up to 9 um long, and presence of cheilocystidia (Oliveira et al. 2020b)); M. pinicola has smaller pleurocystidia (26-46 x 5-12 um), and presence of cheilocystidia (Li et al. 2023). Marasmius vulgaris J.Q. Yan & H. Chen, sp. nov. MycoBank No: 858720 Fig. 4 Etymology. “vulgaris” means “common” or “usual”, refers to the fact that many known species in this genus share similar macroscopic characteristics with the new species. Holotype. CHINA * ZheJiang Province, QingTian County, LiShui City, collected by Qin Na, Bin-Rong Ke, Zhi-Heng Zeng, 6 August 2021, HFJAU2875. Diagnosis. Marasmius vulgaris is mainly characterized by the rather small basidiomata, having a brownish orange, grayish orange pileus; with 3-5 tiers of lamellules; stipe white at apex, gradually darkening from the apex to the base, reddish brown towards base; basidiospores 13.5-16.0 x 3.0-4.0 um, pleurocystidia abundant, narrowly fusiform, lageniform, which frequently have capitate, papillate, or constricted moniliform structures at apices; pile- ipellis and cheilocystidia in the form of broom-cells of the Siccus-type. It dif- fers from M. confertus var. tenuicystidiatus Antonin by having bigger spores, which are up to 16.0 um in length. Description. Pileus 10-30 mm, convex when young, then broadly conical to plane, smooth, with or without slightly obtuse umbo, brownish orange (7C4- 5), grayish orange (5B3) at center, slightly paler to white towards margin, Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 397 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China striate up to 2/3 from the margin. Context thin. Lamellae 2.0-3.0 mm broad, adnexed, ventricose, moderately close, white, with 3-5 tiers of lamellules, edges even, concolorous. Stipe 25-65 mm long, 1.5-2.0 mm thick, central, cylindric, rarely twisted, equal, fibrous, hollow, white at apex, gradually dark- ening from the apex to the base, reddish brown towards base, smooth, and the base covered with white mycelium. Basidiospores (12.5)13.5-16.0(17.0) x 3.0-4.0(4.5) pm (av = 14.8 x 3.8 um), Q = (3.0)3.3-5.0(5.2), clavate or fusoid-clavate, often curved in profile, 3.0-4.0 um broad, clavate in face view, smooth, colorless, hyaline, inamyloid, thin-walled. Basidia 25.0-35.0 x 4.5-6.5 um, clavate, 4-spored. Pleurocystidia 33.0-58.0 x 5.0-11.0 um, narrowly fusiform, lageniform, frequently capitate, papillate, or constricted moniliform structures at api- ces. Cheilocystidia abundant, in form of Siccus-type broom cells, main body 13.0-26.0 x 4.0-11.0 um, cylindrical to clavate, hyaline, thin-walled; apical setulae 4.0-13.0 x 1.0-2.0 um, cylindrical to conical, subacute, yellow to pale yellow, thick-walled. Pileipellis hymeniform, mottled, composed of Sic- cus-type broom cells; main body 14.0-27.0 x 5.5-9.0 um, clavate or pyri- form, hyaline to pale yellow, thin- to thick-walled; apical setulae 4.0-10.5 x 0.8-1.5 um, crowded, cylindrical, subacute, thick-walled. Lamellae trama in- terwoven, with hyphae 7.0-10.0 um in diam, hyaline, dextrinoid, thin-walled. Stipitipellis a cutis composed of cylindrical hyphae 4.5-11.0 um wide, par- allel, smooth. Caulocystidia absent. Clamp connections present. Habitat. Scattered on soil in broad-leaved forest or mixed coniferous and broad-leaved forests. Additional specimens examined. CHINA * JiangXi Province, Lushan Na- tional Nature Reserve, collected by Jing-Cheng Wu, Hong-Zhao Pan, 30 June 2019, HFJAU0904; ZheJiang Province, SuiChang County, collected by Jun- Qing Yan, Yan-Liu Chen, 12 July 2020, HFJAU1901; collected by Yu-Peng Ge, Bin-Rong Ke, Zhi-Heng Zeng, 14 July 2020, HFJAU1976; ZheJiang Province, QingTian County, LiShui City, collected by Jun-Qing Yan, Ze-Wei Liu, 5 Au- gust 2021, HFJAU2719, HFJAU2748. Note. Based on molecular and morphological evidence, M. vulgaris belongs to subg. Globulares ser. Conferti (Oliveira et al. 2020b; Oliveira et al. 2024). With- in this series, M. vulgaris is morphologically similar to M. confertus var. tenui- cystidiatus, M. bondoi Wannathes, Desjardin & Lumyong, and M. graminipes. However, M. confertus var. tenuicystidiatus has smaller basidiospores (10-14 x 4.5—5.7 um), a pruinose stipe apex, sometimes rostrate pleurocystidia (Antonin et al. 2011); M. bondoi has a pruinose pileus surface, pleurocystidia with lobed apices, and the absence of papillate structures (Wannathes et al. 2009a); M. graminipes has larger basidiospores (18-21 x 4 um) and the presence of cau- locystidia (Yang et al. 2013). Morphologically, M. eyssartieri Antonin & Buyck and M. subtangerinus Antonin, Ryoo & H.D. Shin share similar morphological characteristics with the new species M. vulgaris, including basidiospores measuring 13-16 um in length, the presence of pleurocystidia and pileipellis and cheilocystidia in the form of broom cells of the Siccus-type. However, M. eyssartieri differs in having a smaller pileus (5 mm), 0-1 tier of lamellulae, and clavate to occasionally ros- trate pleurocystidia (Antonin and Buyck 2006), while M. subtangerinus has 0-1 tier of lamellulae and clavate to rostrate pleurocystidia (Antonin et al. 2011). Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 328 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China tidia; N-T. Cheilocystidia. All microscopic structures were observed in 5% KOH, structures of E-O and S-T were stained by 1% Congo red. Scale bars: 20 mm (A-D); 20 um (E-T). Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 329 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China In addition, morphologically, M. rongklaensis Wannathes, M. subabundans Chun Y. Deng & T.H. Lim, and M. pseudoconfertus T.H. Li & Chun Y. Deng. have the aspect of M. vulgaris with similarly colored pileus and stipe. However, M. rongklaensis has smaller basidiospores (9-11 x 4.5—5.5 um) and presence of cheilosetae and pileosetae (Wannathes et al. 2019); M. subabundans has smaller basidiospores (7-9 x 3—4.5 um) and absence of pleurocystidia (Deng et al. 2012); M. pseudoconfertus has the slightly wider basidiospores (4-5 pm) and absence of pleurocystidia (Deng et al. 2011). Marasmius subpurpureostriatus J.Q. Yan & H. Chen, sp. nov. MycoBank No: 858721 Figs Etymology. “subpurpureostriatus” refers to its macroscopic morphology, which is similar to that of Marasmius purpureostriatus. Holotype. CHINA + JiangXi Province, Nanchang Botanical Garden, 26 April 2023, collected by Xiao-Lin Yuan, HFJAU4740. Diagnosis. Marasmius subpurpureostriatus is mainly characterized by the rather small basidiomata, grayish-green pileus with deep violet sulcate; the stipe surface is densely pruinose; basidiospores clavate to fusoid-clavate, 17.5-22.0 x 4.0-5.5 pm; pleurocystidia absent; cheilocystidia clavate to broad- ly clavate, occasionally appearing subfusiform, apex obtuse, rarely with short papilla. It differs from M. purpureostriatus by having smaller spores, which are shorter than 22 um. Description. Pileus 20-32 mm, plano-convex to plane, with obtuse umbo at center, smooth, with dark sulcate up to center from the margin, grayish-green (28B4), light green (28A4—5) sulcate dark violet (18F4). Context thin. Lamellae 1.0-2.5 mm broad, adnexed, ventricose, distant, with 0-1 tier of lamellulae, yel- lowish gray (4B2) with white, edges even, concolorous. Stipe 53-70 mm long, 1.5-3.0 mm thick, central, cylindrical with a subbulbous base, pruinose, hollow, apex dark violet (15F5), gradually paler toward the base, base reddish brown (8E4). The base covered by white mycelium. Basidiospores (16.5)17.5—22.0(23.5) x 4.0—5.5(6.0) um (av = 20.0 x 5.0 um), Q = (3.0)3.5-4.5(5.0), clavate to fusoid-clavate, often curved in profile, 4.0— 6.5 um broad, clavate in face view, smooth, colorless, hyaline, inamyloid, thin- walled. Basidia 40.0-50.0 x 8.0-14.0 um, clavate, 4-spored. Pleurocystidia absent. Cheilocystidia 13.0-31.0 x 6.0-13.0 um, irregularly clavate to broadly clavate, rarely subfusiform, apex obtuse, rarely with short papilla, smooth, thin- walled. Pileipellis a hymeniderm composed of cells 16.5-34.0 x 9.0-16.0 um, pyriform or broadly clavate, smooth, hyaline, thin-walled. Lamellae trama in- terwoven, with hyphae 4.0-10.0 um in diam, hyaline, dextrinoid, thin-walled, non-gelatinous. Stipitipellis a cutis composed of cylindrical hyphae 4.0-7.5 um wide, parallel, smooth. Caulocystidia absent. Clamp connections present. Habitat. Scattered on soil in broad-leaved forest. Note. Based on molecular and morphological evidence, M. subpurpureostria- tus belongs to subg. Globulares ser. Purpureostriati (Oliveira et al. 2020b; Oliveira et al. 2024). Within this series, M. subpurpureostriatus is morphologically simi- lar to M. purpureostriatus, M. pseudopurpureostriatus Wannathes, Desjardin & Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 330 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China Figure 5. Morphological structures of M. subpurpureostriatus. A-C. Basidiomata; D. Basidiospores; E. Pileipellis; F-K. Cheilocystidia. All microscopic structures were observed in 5% KOH, structures of E-K were stained by 1% Congo red. Scale bars: 30 mm (A-C); 20 um (D-K). Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 331 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China Lumyong, and M. albopurpureus T.H. Li&C.Q. Wang. However, M. purpureostriatus is distinguished by a long and narrow stipe (52-103 x 0.5-1.5 mm), larger ba- sidiospores (up to 28 um long), and cheilocystidia that are exclusively cylindri- cal, broadly clavate, or pyriform (Desjardin and Horak 1997; Wannathes et al. 2009a). M. pseudopurpureostriatus has a glabrous stipe surface, larger basidio- spores (up to 25.0 um long), and cheilocystidia restricted to clavate or broadly clavate forms (Wannathes et al. 2009a); M. albopurpureus is distinguished by a strongly rugulose to sulcate pileus (white to purple), cream to purple lamellae, and cheilocystidia that are solely clavate to broadly clavate (Wang et al. 2015). In addition, morphologically, only M. bekolacongoli Beeli and M. violaceoi- des Antonin share similar morphological characteristics with the new species M. subpurpureostriatus, including a hymenidermal pileipellis composed of Globulares-type cells, the basidiospores measuring 16.0-24.0 um in length, and absence of pleurocystidia. However, M. bekolacongoli has a larger pileus (30-67 mm) with a yellow colored or tinged striae, and a much longer stipe (50-150 mm) with pale yellow and light brown downward color (Beeli 1928; Douanla-Meli and Langer 2008; Antonin et al. 2010a); M. violaceoides exhib- its a distinctly campanulate, violaceous pileus, a very long and glabrous stipe (110-125 mm) (Antonin 2004). Discussion This study strongly supports the phylogenetic findings of Oliveira et al. (2024). In the phylogenetic tree (Fig. 1), the four new species were each assigned to clade /brunneospermus, /confertus and /purpureostriatus (Oliveira et al. 2020b; Oliveira et al. 2024). Specifically, Oliveira et al. (2020b) proposed the series Brunneospermi based on the clade /brunneospermus, which is charac- terized by a hygrophanous pileus; 2-3 (rarely 4—5) tiers of lamellulae; present elongate pleurocystidia; pileipellis mostly composed of Globulares-type cells, rarely composed of Siccus-type cells; the series Conferti based on the clade / confertus, which is characterized by an orange to ochraceous, or olivaceous pil- eus; 2—5 (or more) tiers of lamellulae; present versiform pleurocystidia; pileipel- lis consisting of Siccus-type cells; and the series Purpureostriati based on the clade /purpureostriatus, which is characterized by a markedly sulcate pileus; distant lamellae, 0-2 tiers of lamellulae; absent pleurocystidia; pileipellis con- sisting only of Globulares-type cells (Oliveira et al. 2020b). Four new species also respectively conform to these characteristics. In the phylogenetic tree (Fig. 1), M. blandus groups together with M. brun- neospermus and M. fusicystidiosus; M. vulgaris is closely related to M. gr- aminipes; and M. subpurpureostriatus is closely related to M. purpureostri- atus. The distinctions among these species have been discussed in the section of note. In addition, M. xingshanensis groups together with M. mac- rocystidiosus. However, M. macrocystidiosus can be distinguished from M. xingshanensis by having larger basidiospores (6.9-10.5 x 3.3-4.4 um) and presence of cheilocystidia (Kiyashko et al. 2014). In conclusion, the phylogenetic tree strongly supports new species distinct clade, suggesting that Marasmius may contain undiscovered biodiversity. Further studies are needed to elucidate its evolutionary history and ecological roles, thereby enhancing our understanding. Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 332 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China Key to related species 15 16 Pileipellis is composed of Siccus-type CelIS...........cccccescceseeceeeeeeeeeeeeeeees 2 Pileipellis is composed of Globulares-type CellS ...........cccccecceesseeesseeeeees 11 Cheilosetae and pileosetae present...............ccseceesseeeees M. rongklaensis Cheilosetae and pileosetae ADSENT.......... cece cccccessecceceessseeeceeesssseeeeesenaaeee 3 Basidiospores mainly shorter than 10.0 UM..............:::ccccsssccsssseceeesseeeesseees 4 Basidiospores mainly longer than 10.0 UM .......... cee ecesseceesrteeeeeteeeeseeeeees 5 Cheilocystidia: PreSe mt ccrs.c: verv.ceduseese, cise ieee odectirae: cede eeuts M. subabundans CME GCY SULA SA SIN sii ck, te Rea ah Ae oe ve cee hea soy cheshire M. magnus PICU CYSUGIA ADSE Ms coscc vee dedeceur recy peverecsmeatien snot: M. pseudoconfertus PICUROCYSTIGral DIESEMNiler wes 5, eee ccc tne pen Oe Sear eee ee ee 6 Basidiospores mainly longer than 18.0 UM............... eee M. graminipes Basidiospores mainly shorter than 18.0 UM.............::ccccesscccseseceeeereeesseeeees 7 Pileus up to 65 mm wide, stipe with pruinose at apex............ceeeeeeseeeeeees Woks: emplenR caakGeenueaeente eamcesarane «tee bestscaetessreteens M. confertus var. tenuicystidiatus PHéusimainiy: 330 Mim diameter. 18. orale tect oadihanennepters Meaplatanti nee: 8 Besta cre IU Foun 24 10% 0 Real De 1\ =| cota eam N nn OER IP ee TEU ROTU RS ER, ere NUE EA AOE Gr PrN aT A 9 Deniers eS ates Solel ge mtants pets seh ee is mms cious a as enc d ana hone ae 10 Lamellae pale yellowish white to pale cream.................. M. subtangerinus Lamellae brown, pubescent €dGeé..............c:ccccssssccsssseeessteeeees M. eyssartieri Pileus with pruinose, pleurocystidia with lobed apices, and the absence of Pai AVSS UNCLES acs BSc Rete tice ts tA Fox rane eae eal oars See M. bondoi Pileus glabrous, pleurocystidia apices with capitate, papillate and con- stricted into MonilifOrmM StrUCTUFES ........... cee ceececeseeeeseeeeseeeeees M. vulgaris Basidiospores mainly longer than 10.0 UM ..............:::cccessceessseceseseeeeeseeeees 12 Basidiospores mainly shorter than 10.0 UM...............ccccssscecsesseecsessseesensees 17 Pileus grayish green and light green, sulcate dark violet...............0.eeeeeee aie Peaks dee Ne deak reat tee teens LAER, a8 anaes See M. subpurpureostriatus PIREUS DROW TO: FEGLOF- VIG TEL cit nears eRe Bae dao sec ew tee girs ee Bes 13 Pileus with a yellow colored or tinged striae and stipe with pale yellow and light: browndownward Colornt 0c. Mca aks M. bekolacongoli NOIR AS AD OVER secsec cee. euinnvmecited nat cane kn ems oecere at tue manatee cc teatneneeey ceetts lenient 14 Basidiospores 15.5-22.0 um long, stipe 110-125 x 2.5-3.5 mm.............. toss | hunted as! Las itinraca is Tihs, cheatin CRONDe i RN CEU ORM, WS M. violaceoides Basidiospores mainly longer than 22.0 um, stipe mainly < 100 mm long....15 Pileus larger, up to 38 mm, stipe apex grayish magenta ..............::ceeeeeeees Patras abarcaicotomicea eat de tes ut seas aie c mary use lacng tee ee M. pseudopurpureostriatus Pileus rather small, only up to 20 mm, stipe apex Violet...............ceeee 16 Pileus with dark violet ridges, stipe 0.5-1.5 mm thick, basidiospores up to 2 OMANI LONG iets. eerste etree Oe ere Siyet rae ear tere Me ttsses OTC M. purpureostriatus Pileus with white to violet white ridges, stipe 1.5-3 mm thick, basidio- spores Shorter than 25 UM ONG .............::ccecessecceessseeeeenees M. albopurpureus CHENSCYSIIGIE-PRES SMM e205 scan een Ban AR a, Stan seuet Me, Pear ten 18 CUHEIOCYSUCA-AU SOM ay. a2. a3 lode newton hne et eieed eat al rhea: 27 BaSidiOSpOres: Mainly Se OMIM AWIGTED 2.0. hcos ctu aut esce tangs ededbonecuaneters 19 Basidiospores mainly < 3.5 UM Wit ..........cecccesscccsssscecssseeecsesseeesesaees 23 Pleurocystidia mainly shorter than 50 UM..............cc:cccessceeeneees M. pinicola Pleurocystidia mainly longer than 50 UM ............ccesececsssceeeesseeeeesteeeeeaees 20 Mycokeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 333 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China 20 | ‘Cheiloeystidia: Mainly SHOrter Wan SOVUIM ste eced cree. maceccrersraccenensuepceuncepess 21 - Cheilocystidia mainly longer than 30 UM................ccecssseesssteesssteeessteeeeees 22 21 Basidiospores 8.5-10.0 x 3.5—4.0 um, stipe 110 x 3 mm... fusicystidiosus - Basidiospores 6.2-7.9 x 3.5-4.2 um, stipe 50-70 x 3-6 mmM............... Seausetesdlt Abia oats tulecarsikel is eeactants eatin maruscei tea Rh Done: M. goossensiae ZAP SUP SSGnayts Mi: OLANG Get ser d e Seaton hs es dete ee eed M. macrocystidiosus =" Stipe WwhitesOr ‘Qrayisti WIIte: 2, <,.5,cccesele weevodenstececes added M. pallidibrunneus 23 Pleurocystidia subfusiform, narrowly utriform, apex with short or long papilla, surface rarely with NOCUIOSE..............cccceescceseeceeeeeeeees M. blandus = WMNOTEAS FACES 2228s cer chores vecees tenes ciers Bele aah ccematee sae euociene Wl Sn watoevantiwl. 24 24 Basidiospores mainly longer than 6.5 UM.............ccccecessseessstseesesteeesseneees 25 - Basidiospores mainly shorter than 6.5 UM..............cccessccsssseceseneeessneeeees 26 25 Stipe pale ochraceous towards the base or pale whitish-brown or gray- ish-brown, pileocystidia aDSent...............ccesceessteeeeees M. brunneospermus - Stipe yellowish white, turning brownish orange to light brown on bruising, DIEOCYVSTICIADLESEM avec cetecees aces naeenegeemcuns eumerseederhnaactenssmmrne M. desjardinii 26 ‘Plewroeystidia Only fUSO cco ic.n csc ntered ee areendsenetinonietes M. phlebodiscus - Pleurocystidia cylindrical, clavate, fusoid, often subrostrate ...................0. Ey CR trae ese SRE, Coe kt een EAs ean ee rt lay M. muramwyanensis 27 Basidiospores mainly longer than 8.0 UM............ccceceeeseeeentees M. riparius = —Basidiosporessmainly-shorter than: 7:0) WM ies cc. erecta betcrer dedicate. 28 28 Pileus mainly > 30 mm in diameter, stipe 60-110 x 4-7 um, pileocystidia SOS SCO Gs HIM ee herent ari cruaten eae tere eave ear M. ochraceus — Pileus mainly < 30 mm in diameter, stipe 20-41 x 1.5-3 um, pileocystidia Bo SOX 7 165 Wn eat eeceee eee memnsiere Stivs ceevnaseccusevedgr ees M. xingshanensis Acknowledgments The authors are very grateful for the assistance of Lin-Gen Chen, Cheng-Feng Nie, Ling Ding, Jing-Cheng Wu, Yan-Liu Chen, Yu-Peng Ge, Bin-Rong Ke, Zhi- Heng Zeng, Ze-Wei Liu, Xiao-Lin Yuan in the field specimen collection and the anonymous reviewers of the manuscript. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Use of Al No use of Al was reported. Funding This work was financed by the National Natural Science Foundation of China (32460326), Jiangxi Provincial Natural Science Foundation (20224BAB205003), and the Project of FAAS (XTCXGC2021007). MycoKkeys 120: 317-338 (2025), DOI: 10.3897/mycokeys.120.157997 334 Hong Chen et al.: Four new species of Marasmius subgenus Globulares from China Author contributions Conceptualization, Jun-Qing Yan; methodology, Jun-Qing Yan and Sheng-Nan Wang; software, Hong Chen, and Yu-Qin Xu; formal analysis, Hui Zeng, Ya-Ping Hu, Jun-Qing Yan, and Sheng-Nan Wang; investigation, Hong Chen, Yu-Qin Xu and Jun-Qing Yan; resources, Hui Zeng, Ya-Ping Hu and Jun-Qing Yan; writing - original draft, Hong Chen; writing - review and editing, Jun-Qing Yan; visualization, Jun-Qing Yan and Sheng-Nan Wang; supervision, Jun-Qing Yan; project administration, Jun-Qing Yan; funding acquisition, Jun-Qing Yan. All authors have read and agreed to the published version of the manuscript. Author ORCIDs Hong Chen ® https://orcid.org/0009-0004-0107-3962 Yu-Qin Xu © https://orcid.org/0009-0002-0735-8323 Hui Zeng © https://orcid.org/0000-0003-2025-844X Ya-Ping Hu © https://orcid.org/0000-0003-1242-1139 Sheng-Nan Wang ® https://orcid.org/0000-0003-0648-271X Jun-Qing Yan © https://orcid.org/0000-0003-1128-5171 Data availability All of the data that support the findings of this study are available in the main text. References Antonin V (2003) New species of Marasmius (Basidiomycetes, Tricholomataceae) from tropical Africa—l. Sect. Epiphylli, Fusicystides, Globulares, Hygrometrici and Neoses- siles. Mycotaxon 85: 109-130. Antonin V (2004) New species of marasmioid genera (Basidiomycetes, Tricholoma- taceae) from tropical Africa—V. Marasmius violaceoides, a new species based on M. violaceus Henn. in the sense of Singer. 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