Chinese scientific names

Yang, Kun L., Lin, Jia Y., Li, Guang-Mei & Yang, Zhu L., 2025, New and interesting fungi from Guangzhou City, South China, Phytotaxa 707 (1), pp. 1-95 : 6-42

Chinese scientific names

Chinese scientific names of nomenclatural novelties are designated in Supplement 2.

Taxonomy

Ascomycota Caval.-Sm.

Dothideomycetes O.E. Erikss. & Winka

Catinellales Ekanayaka, K.D. Hyde & H.A. Ariyansa

Atramentodiscaceae Kun L. Yang, Jia Y. Lin & Zhu L. Yang , fam. nov. ( Figs 1–2; Table 2; Supplements 2–3) Registration identifier FN572700

Etymology:— See the etymology of Atramentodiscus .

Type:— Atramentodiscus Kun L. Yang, Jia Y. Lin & Zhu L. Yang (see below)

Diagnosis:— Differing from the sister family Catinellaceae by the presence of croziers, smoother apothecial margin, and more uniformly pigmented ectal excipulum.

Description:— Anamorph: not observed. Teleomorph: Ascomata apothecial, sessile, blackish. Apothecium becoming nearly plane following maturity, with a margin typically not groovily striate. Hymenium surface smooth and shiny. External surface covered with abundant hyphae.

Ascospores unicellular, blackish, smooth under LM, but with verrucose ornamentation under SEM. Asci cylindrical, inoperculate, eight-spored, unitunicate, inamyloid, with an ocular chamber at the apex, and a basal part with a crozier. Hamathecia presenting as paraphyses, unbranched or branched, septate, with a gelatinized, usually inflated apex. Subhymenium composed of a textura angularis-globulosa. Medullary excipulum composed of a textura intricata. Ectal excipulum composed of a textura angularis-globulosa, becoming slightly darker towards the external surface. External hairs septate, unbranched.

Phylogeny:— According to the BLAST search of ITS and nrLSU sequences in GenBank ( NCBI 2025), our collections HKAS147739 and HTBM1993 represent a single species close to Catinellaceae and its nearby families in Dothideomycetes. According to recent studies on phylogeny of Dothideomycetes ( Hongsanan et al. 2020 a, Maharachchikumbura et al. 2021), Catinellaceae and eight other families, Aplosporellaceae , Botryosphaeriaceae , Holmiellaceae, Homortomycetaceae , Melanopsaceae , Phyllostictaceae , Planistromellaceae and Saccharataceae , tend to form a monophyletic group. Therefore, a phylogeny was inferred to classify our collections with collections of these nine families and our collections as the ingroup, and collections of a farther family Bezerromycetaceae ( Hongsanan et al. 2020a) as the outgroup, using two loci (ITS & nrLSU) available for most of the above collections ( Table 2; Supplement 3). In the result, the ML and BI topologies are almost identical, only with minimal variation in statistical support, so only the tree inferred from the ML analysis is displayed ( Fig. 1). Our collections form an undescribed family-level clade sister to Catinellaceae with significant support, here named Atramentodiscaceae. This family is monotypic for the moment.

Notes:— Historically, due to morphological similarity, many ascomycetes producing tiny, apothecial, darkly coloured and usually lignicolous ascomata were frequently assigned to the order Helotiales , under several genera especially Bulgariella P. Karst. , Catinella Boud. , Lagerheima Sacc. , Patinella Sacc. and Patellaria Fr. (e.g., Saccardo 1875, Boudier 1907, Durand 1922, Gamundí 1981). However, molecular phylogenetic studies confirmed this morphological assembly as rather heterogeneous, that it comprises taxa across many families (e.g., Catinellaceae , Holwayaceae), orders (e.g., Catinellales , Patellariales ), and even classes (e.g., Dothideomycetes, Leotiomycetes) ( Greif et al. 2007, Schoch et al. 2009, Yacharoen et al. 2015, Iturriaga et al. 2017, Hongsanan et al. 2020 a, b, Quijada et al. 2022). With molecular markers, our collections are recognized in the class Dothideomycetes, order Catinellales , representing a new family. The morphology of Atramentodiscaceae resembles Catinellaceae , but Atramentodiscaceae distinctly presents a crozier at ascus base that is not reported in Catinellaceae ( Durand 1922, Gamundí 1981, Greif et al. 2007).

According to exsiting relevant ITS sequences in GenBank, there are some more distinctly divergent lineages like DQ812139, MZ997311 and MZ997312 around Catinellaceae and Atramentodiscaceae in Catinellales awaiting descriptions, if no sequencing errors are included. Based on the recent scale on family delimitation in Dothideomycetes, possibly they all represent new families.

Atramentodiscus Kun L. Yang, Jia Y. Lin & Zhu L. Yang , gen. nov. ( Figs 1–2; Table 2; Supplements 2–3) Registration identifier FN572701

Etymology:— Referring to the blackish, discoid to nearly plane ascomata like inkstones.

Type:— Atramentodiscus nanshaensis Kun L. Yang, Jia Y. Lin & Zhu L. Yang (see below)

Description:— Anamorph: not observed. Teleomorph: Ascomata tiny, sessile, blackish. Apothecium discoid at first, becoming nearly plane, with a margin not groovily striate or very slightly groovily striate. Hymenium surface smooth and shiny. External surface covered with abundant hyphae anchored to the substrate. Context fleshy when fresh, becoming fragile after dried.

Ascospores unicellular, oblong, more or less amygdaliform, mostly biguttulate, blackish, smooth under LM, but with verrucose ornamentation under SEM. Asci cylindrical, inoperculate, eight-spored, unitunicate, inamyloid, with an ocular chamber at the apex, and a flexuous basal part with a crozier. Paraphyses subcylindrical, unbranched to branched, septate, with a gelatinized, usually inflated apex; apexes of paraphyses agglutinated together as a gelatinous pseudoepithecium covering the hymenium. Subhymenium composed of a dense textura angularis-globulosa. Medullary excipulum composed of a dense textura intricata. Ectal excipulum composed of a textura angularis-globulosa, becoming slightly darker towards the external surface. External hairs (anchoring hyphae) septate, unbranched.

Phylogeny:— See Atramentodiscaceae.

Notes:— See Atramentodiscaceae.

Atramentodiscus nanshaensis Kun L. Yang, Jia Y. Lin & Zhu L. Yang , sp. nov. ( Figs 1–2; Table 2; Supplements 2–3)

Registration identifier FN572702

Etymology:— Referring to the type locality.

Type:— CHINA. Guangdong Province: Guangzhou City, Nansha District, South Waterfront Cape (Nanbin Shuijiao) Park , 22°46’48”N, 113°30’51”E, elevation 1 m, April 27, 2024, Kun L . Yang & Jia Y . Lin, L24105 ( HKAS147739 , holotype ( ITS: PV620896 ; nrLSU: PV616964; rpb2: PV642296; tef-1α: PV642317 ); HTBM1992 , isotype) .

Description:— Anamorph: not observed. Teleomorph: Ascomata tiny, sessile; odour indistinct; taste unknown. Apothecium 0.5–4.5 mm broad, discoid at first, becoming nearly plane, with a straight to flexuous margin not groovily striate or very slightly groovily striate. Hymenium surface smooth and shiny, dull black (#0B0C0E). External surface usually concolorous with hymenium surface , but sometimes lighter as flint brown (#736960), covered with abundant hyphae anchored to the substrate. Context fleshy when fresh, becoming fragile after dried.

Ascospores unicellular, oblong, more or less amygdaliform, mostly biguttulate, rarely uniguttulate, slightly thick-walled to thick-walled, blackish, smooth under LM, but with verrucose ornamentation under SEM, {40/4/2} 8.5–10.5 (11) [9.55 ± 0.62, 9.00] × (4) 4.5–5.5 [4.76 ± 0.34, 4.50] µm, Q = (1.70) 1.80–2.33 [2.01 ± 0.14, 2.00]. Asci cylindrical, inoperculate, eight-spored, unitunicate, nearly colourless to slightly brownish, inamyloid, 75–105 × 4.5–5.5 µm, with an ocular chamber at the apex, and a flexuous basal part with a crozier. Paraphyses subcylindrical, unbranched to branched, nearly colourless to slightly brownish, 60–105 × 0.5–3 µm, thin-walled, septate, with a gelatinized, usually inflated apex; apexes of paraphyses agglutinated together as a gelatinous pseudoepithecium covering the hymenium. Subhymenium up to 20 µm thick, slightly brownish to brownish, composed of a dense textura angularis-globulosa. Medullary excipulum up to 25 µm thick, brownish to blackish, composed of a dense textura intricata. Ectal excipulum up to 150 µm thick, composed of a textura angularis-globulosa, with cells slightly thick-walled to thick-walled, 10–40 µm in diameter, blackish, becoming slightly darker towards the external surface. External hairs (anchoring hyphae) cylindrical, up to 1 mm long, 8–10 µm wide, darkly brownish, thick-walled, septate, unbranched, with an obtuse apex.

Habit and distribution:— Gregarious, on estuarial deadwood. Currently known from China, and probably also from USA (e.g., MH281881 ).

Other collection examined:— CHINA. Guangdong Province: Guangzhou City, Nansha District, South Waterfront Cape (Nanbin Shuijiao) Park , 22°46’48”N, 113°30’51”E, elevation 1 m, April 27, 2024, Jia Y . Lin & Kun L . Yang, L24106 ( HTBM1993 ) .

Phylogeny:— See Atramentodiscaceae.

Notes:— In the close family Catinellaceae , an ontogenic study by Greif et al. (2007) found that the primordial ascomata of Catinella olivacea (Batsch) Boud. are globular with an inner cavity, and that the mature “apothecia” are resulted from apical rupture, and thus they considered the ascomata of C. olivacea as ascostromata better than incipient apothecia. In our collections of Atramentodiscus nanshaensis , the most immature ascomata that we observed are already apothecial ( Fig. 2b, yellow arrow). Whether they are also globular with an inner cavity earlier is undetermined at present.

Basidiomycota R.T. Moore

Agaricomycetes Doweld

Agaricales Underw.

Agaricaceae Chevall.

Agaricus cantonotraminus Kun L. Yang, Jia Y. Lin & Zhu L. Yang , sp. nov. ( Figs 3–4; Table 3; Supplements 2 and 4)

Registration identifier FN572703

Etymology:— Referring to the collection site in tram track area in Guangzhou City (Canton, an old name of Guangzhou City).

Diagnosis:— Differing from Agaricus braendlei by the narrower basidiospores, and from A. pampeanus by the gregarious basidiomata and longer basidiospores.

Type:— CHINA. Guangdong Province: Guangzhou City, Haizhu District, track area 40 m westward from Liede Bridge South Station of Guangzhou Tram Haizhu Line 1, 23°06’36”N, 113°19’46”E, elevation 5 m, March 9, 2025, Kun L . Yang, Jia Y . Lin & anonymous staffs of Guangzhou Metro , L25049 ( HKAS147740 , holotype ( ITS: PV620892 ; nrLSU: PV616968; rpb2: PV642295; tef-1α: PV642315 ); HTBM2331 , isotype) .

Description:— Basidiomata small to medium-sized; odour fungal; taste fungal and sweet. Pileus 23–82 mm in diameter, hemispherical at first, becoming plano-convex to plano-concave, ceramic white (#FEFEFA), merino white (#F9F5EC) to light brown (#B39966), becoming pinkish after soaked from lamellae with rain, smooth or forming fibrous, woolly to floccose squamules radially and horizontally; context ceramic white (#FEFEFA) to merino white (#F9F5EC), without a colour change after damaged. Lamellae adnexed to free, crowded, light peach red (#F8D3C9), cherrywood brown (#DAB697), beaver brown (#9D7B69) to coffee-bean brown (#483625), with an entire edge, interspersed with abundant lamellulae. Stipe 37–105 mm long, 6–14 mm thick, fusiform, clavate to subcylindrical, ceramic white (#FEFEFA) to merino white (#F9F5EC), covered with felty to fibrous squamules below the annulus; context ceramic white (#FEFEFA) to merino white (#F9F5EC), without a colour change after damaged. Annulus superior, membranous, easily broken, concolorous with the squamules on the stipe.

Basidiospores {40/4/2} 7–8.5 (9.5) [7.96 ± 0.56, 8.00] × (3.5) 4–5 (6) [4.63 ± 0.43, 4.50] µm, Q = (1.40) 1.50– 2.00 (2.14) [1.73 ± 0.15, 1.67], ellipsoid to oblong, more or less ovoid to amygdaliform, slightly thick-walled to thick-walled, smooth, brownish, with a small apiculus, with a germ pore indistinct to distinct. Basidia 16–26 × 7–9.5 μm, clavate, two- to four-spored, thin-walled, nearly colourless. Lamella trama subregular, composed of 2–8 µm wide, thin-walled, nearly colourless, compact, moderately to frequently branching hyphae. Cheilocystidia absent. Pleurocystidia absent. Pileus squamules composed of a rectocutis to tomentum by thin-walled, sometimes encrusted, nearly colourless hyphae 2–10 µm wide. Annulus composed of radially irregular, thin-walled to slightly thick-walled, nearly colourless, compact, moderately to frequently branching hyphae 3–7 µm wide. Clamp connections absent.

Habit and distribution:— Gregarious, on urban lawn of Zoysia japonica . Currently known from China and South Korea.

Other collections examined:— CHINA. Guangdong Province: Guangzhou City, Haizhu District, track area 40 m westward from Liede Bridge South Station of Guangzhou Tram Haizhu Line 1, 23°06’36”N, 113°19’46”E, elevation 5 m, April 1, 2023, Kun L . Yang, K2309 ( HTBM0524 ) ; Guangdong Province: Guangzhou City, Haizhu District, track area 35 m eastward from Liede Bridge South Station of Guangzhou Tram Haizhu Line 1, 23°06’38”N, 113°19’50”E, elevation 5 m, March 9, 2025, Kun L . Yang & Jia Y . Lin, L25048 ( HTBM2330 ) .

Phylogeny:— According to the BLAST search of ITS sequences in GenBank ( NCBI 2025), our collections HKAS147740, HTBM0524 and HTBM2330 represent a single species of Agaricus sect. Agaricus . Therefore, a phylogeny was inferred to classify our collections with representative collections of known species of Agaricus sect. Agaricus and our collections as the ingroup, and a collection of Agaricus sect. Rubricosi as the outgroup according to a recent study on phylogeny of Agaricus ( Zhao et al. 2016) , using the ITS locus available for most of the above collections ( Table 3; Supplement 4). In the result, the ML and BI topologies are almost identical, only with a minimal variation in statistical support, so only the tree inferred from the ML analysis is displayed ( Fig. 3). Our collections form an undescribed clade with two other collections misidenfitied as “ A. campestris ” from China or South Korea with significant support, and this clade is sister to another undescribed clade from Australia with significant support.

Notes:— As compared with other sections in the same genus, Agaricus sect. Agaricus is typically characterized by nonspecial odour, indistinct colour changed after touched or damaged, absence of cheilocystidia, and negative KOH and Schäffer’s reactions ( Zhao et al. 2016). In this section, the Agaricus campestris complex is well-known for the difficulty in species identification.

Species of Agaricus campestris complex are often nearly indistinguishable on macromorphology. Previously, the well-known species A. campestris was frequently reported in China, but as revealed by molecular phylogeny, these reports are probably misidentifications of several closely related species like A. aristocratus and A. braendlei , and true A. campestris has not been proven as distributed in China with molecular evidence ( Liu et al. 2020). Agaricus cantonotraminus here described is one more new species recognized in the A. campestris complex. Phylogenetically its closest known species are A. braendlei and A. pampeanus , but the shape and size of their basidiospores are significantly different ( Spegazzini 1880, Peck 1899, Liu et al. 2020). It may be an edible species, and possibly many citizens have eating experience on it without knowing its correct name (e.g., Chen & Zhang 2019).

In Guangzhou City, A. cantonotraminus frequently produces remarkable fairy rings on grass-covered track areas of tram in every spring, but after entering summer it rarely emerges.

Hymenagaricus conghuaensis Kun L. Yang, Jia Y. Lin & Zhu L. Yang , sp. nov. ( Figs 5–6; Table 4; Supplements 2 and 5)

Registration identifier FN572704

Etymology:— Referring to the type locality.

Diagnosis:— Differing from other Asian Hymenagaricus species by a combination of purplish brown pileus squamules flaky and slightly felty at center, becoming fibrous towards margin, and ellipsoid basidiospores sized 4.5–5 × 3–3.5 µm.

Type:— CHINA. Guangdong Province: Guangzhou City, Conghua District, Liuxi River National Forest Park , on a moss land, 23°45’17”N, 113°46’56”E, elevation 200 m, June 20, 2024, Kun L . Yang & Jia Y . Lin, K24041 ( HKAS147741 , holotype ( ITS: PV620889 ; nrLSU: PV616969; rpb1: PV642282; rpb2: PV642294; tef-1α: PV642311 ); HTBM2031 , isotype) .

Description:— Basidioma small; odour fungal; taste unknown. Pileus 11 mm in diameter, plano-convex; background ceramic white (#FEFEFA); squamules flaky and slightly felty at center, becoming fibrous towards margin, merino white (#F9F5EC), meat brown (#D7B19D) to beaver brown (#9D7B69), becoming lighter towards margin. Lamellae adnexed to free, nearly crowded, medium grey (#E0DEDD), muddy grey (#938F89) to flint brown (#736960), with a slightly serrate edge, interspersed with abundant lamellulae. Stipe 18 mm long, 1.5 mm thick, cylindrical, slightly curved; background merino white (#F9F5EC) to mist brown (#DCD8C9); squamules fibrous, merino white (#F9F5EC); context turning darker and reddish after damaged. Annulus superior, fibrous and fugacious, concolorous with the squamules on the stipe.

Basidiospores {40/1/1} (4) 4.5–5 [4.63 ± 0.27, 4.50] × 3–3.5 [3.23 ± 0.25, 3.00] µm, Q = 1.29–1.67 [1.44 ± 0.13, 1.50], ellipsoid, slightly thick-walled, smooth, tinged cotton brown (#D2CAAD) to dark heather brown (#9D917B), with a small apiculus, with an indistinct germ pore. Basidia 16–18.5 × 5–7 μm, clavate, one-, two-, three- or four-spored, thin-walled, nearly colourless. Lamella trama regular to subregular, composed of 3–11 µm wide, thin-walled, nearly colourless, compact, moderately to frequently branching hyphae. Cheilocystidia abundant, 12–26 × 3–6 µm, clavate to subcylindrical, sometimes flexuous, thin-walled, smooth, nearly colourless. Pleurocystidia absent. Pileus squamules composed of a hymeniderm to trichoderm at center, becoming an entire trichoderm towards margin, by slightly thick-walled and encrusted, interwoven hyphae 2–8 µm wide issued from a lower layer of clavate to subcylindrical cells measured 13.5–24 × 6.5–11 µm standing on subglobose cells measured 11–17 × 10–17 µm, tinged blush red (#D9B7B4). Stipe squamules composed of a trichoderm by 2.5–4 µm wide, thin- to slightly thick-walled and encrusted hyphae, tinged light cardamon yellow (#E9E9AD). Clamp connections occasionally present in the pileus squamules, absent in the other tissues.

Habit and distribution:— Solitary, on soil, in southern subtropical monsoon forests. Currently known from China.

Phylogeny:— According to the BLAST search of ITS sequence in GenBank ( NCBI 2025), our collection HKAS147741 is a species of Hymenagaricus . Therefore, a phylogeny was inferred to classify our collections with representative collections of known species of Hymenagaricus and HKAS147741 as the ingroup, and two collections of Coniolepiota as the outgroup according to our recent study on phylogeny of Agaricaceae ( Yang et al. 2024a) , using two loci (ITS and nrLSU) available for most species in this group ( Table 4; Supplement 5). In the result, the ML and BI topologies are almost identical, only with a minimal variation in statistical support, so only the tree inferred from the ML analysis is displayed ( Fig. 5). The genus Hymenagaricus is resolved into five distinct groups, and our collection HKAS147741 forms an undescribed clade sister to Hy. splendidissimus with significant support.

Notes:— The genus Hymenagaricus is typically characterized by brownish basidiospores and pileus squamules with a more or less hymeniform structure ( Heinemann 1981, Heinemann 1985, Heinemann & Little Flower 1984). Watling (1999) proposed a new genus Heinemannomyces to accommodate a species similar to Hymenagaricus but with floccose squamules, called He. splendidissimus . Yang et al. (2024b) found hymeniform structure beneath the floccose squamules, explained He. splendidissimus as a modified Hymenagaricus species with outgrowing hyphae on squamules, and synonymized Heinemannomyces to the earlier name Hymenagaricus with the new combination Hymenagaricus splendidissimus .

According to the current phylogeny ( Fig. 5), the new species Hy. conghuaensis proposed here, is a species most closely related to Hy. splendidissimus ever known. It also exhibits outgrowing hyphae on the central squamule on pileus, generally similar to Hy. splendidissimus , but the squamules are fewer and sparser, the basidioma size is smaller, and the greyish brown lamellae and brownish basidiospores are different.

in this study is in bold. Unavailable items are indicated with -. Holotypes and paratypes are indicated by HT and PT,

respectively; note that the paratypes of a species are only indicated when its holotype is not present.

Leucocoprinus phantasmaticus Kun L. Yang, Jia Y. Lin & Zhu L. Yang , sp. nov. ( Figs 7–8; Table 5; Supplements 2 and 6)

Registration identifier FN572705

Etymology:— Referring to the basidiomata hanging like ghosts following senescence.

Diagnosis:— Differing from other Asian Leucocoprinus species by a combination of typically lignicolous habit, whitish fragile basidiomata, and pileus squamules composed of inflated cells.

Type:— CHINA. Guangdong Province: Guangzhou City, Tianhe District, Furnace Mountain (Huolushan) Forest Park, 23°11’07”N, 113°23’05”E, elevation 200 m, June 13, 2023, Kun L . Yang , K23282 ( HKAS147742 , holotype ( ITS: PV620893 ; nrLSU: PV616971; rpb1: PV642280; rpb2: PV642293; tef-1α: PV642312 ); HTBM0797 , isotype) .

Description:— Basidiomata fragile, tiny to small, erect on substrate at first, becoming hanging following senescence, without a distinct colour change after touched, damaged or dried; odour fungal; taste unknown. Pileus 5–16 mm in diameter, conical to paraboloid at first, becoming hemispherical to plano-convex, more or less umbonate at center; background pure white (#FFFFFF), ceramic white (#FEFEFA), merino white (#F9F5EC), star white (#FBFBEB) to cream orange (#FEF3CE); squamules furfuraceous, concolorous with or lighter than the background. Lamellae free, crowded, pure white (#FFFFFF), ceramic white (#FEFEFA), merino white (#F9F5EC), star white (#FBFBEB) to cream orange (#FEF3CE), with a serrate edge, interspersed with abundant lamellulae. Stipe 21–28 mm long, 1–2 mm thick, cylindrical or tapering upwards, slightly curved, bulbous at the base; background concolorous with the pileus background; squamules fibrous to furfuraceous, concolorous with the pileus squamules. Annulus superior, easily broken and fugacious, concolorous with the squamules on the stipe.

Basidiospores {40/3/3} (8) 8.5–10.5 (11) [9.63 ± 0.64, 10.00] × (5) 5.5–6.5 (7) [6.04 ± 0.48, 6.50] µm, Q = (1.45) 1.46–1.80 (1.82) [1.60 ± 0.10, 1.54], ellipsoid to oblong, more or less ovoid to amygdaliform, slightly thick-walled to thick-walled, smooth, nearly colourless but containing slightly yellowish to greenish granules, dextrinoid, with a small apiculus, with a distinct germ pore. Basidia 15–18 × 8.5–10 μm, clavate, two- to four-spored, thin-walled, nearly colourless, each surrounded by 4–5 hymenial physalides sized 15–21 × 11.5–17 μm. Subhymenium poorly developed. Lamella trama composed of 2.5–6 µm wide, thin-walled, nearly colourless, compact, moderately to frequently branching hyphae. Cheilocystidia abundant, 20–65 × 12–21 µm, clavate, thin-walled, smooth, nearly colourless. Pleurocystidia absent. Pileus squamules composed of subglobose, ellipsoid to elongate or clavate, slightly thick-walled, nearly colourless inflated cells sized 21–50 × 12–16 µm. Stipe squamules composed of a trichoderm by 5–11 µm wide, thin- to slightly thick-walled, nearly colourless hyphae. Clamp connections absent.

Habit and distribution:— Usually solitary on deadwood, occasionally on rocky soil, in southern subtropical monsoon forests. Currently known from China.

Other collections examined:— CHINA. Guangdong Province: Guangzhou City, Tianhe District , South China Agricultural University, Arboretum , 23°09’27”N, 113°21’22”E, elevation 40 m, May 12, 2023, Jia Y . Lin & Kun L . Yang , L2338 ( HTBM0871 ); same location, June 12, 2023, Kun L . Yang , K23269 ( HTBM0784 ); same location, August 30, 2023, Jia Y . Lin, L23351 ( HTBM1332 ). Guangdong Province: Guangzhou City, Huangpu District , Jiangdong Village , Boluoshan Hill , 23°11’36”N, 113°32’25”E, elevation 150 m, October 10, 2023, Zhen-Chao Liu, Jia Y . Lin & Kun L . Yang, S23634 ( HTBM1745 ). Collection of other species sequenced for phylogenetic analyses: Macropsalliota meleagris ( CHINA. Guangdong Province: Guangzhou City , Tianhe District , South China Botanical Garden , 23°11’22”N, 113°21’32”E, elevation 50 m, November 2, 2024, Fei Bao, Kun L. Yang & Jia Y. Lin, L24283 ( HTBM2268 )) .

Phylogeny:— According to the BLAST search of ITS sequences in GenBank ( NCBI 2025), our collections HKAS147742, HTBM0784, HTBM0871 and HTBM1745 represent a single species of Leucocoprinus s. l. Therefore, a phylogeny was inferred to classify our collections with representative collections of known species of Leucocoprinus s. l. and our collections as the ingroup, and a collection of Chlorophyllum as the outgroup according to our recent study on phylogeny of Agaricaceae ( Yang et al. 2024a) , using four loci (ITS, nrLSU, rpb2 & tef-1α) necessary for resolving this group ( Table 5; Supplement 6). In the result, the ML and BI topologies are almost identical, only with a minimal variation in statistical support, so only the tree inferred from the ML analysis is displayed ( Fig. 7). Our collections HKAS147742, HTBM0784, HTBM0871 and HTBM1745 form an undescribed clade close to Leucocoprinus birnbaumii and its allies with significant support, and this clade is further split into two significantly supported subclades with minimal pairwise variation.

Notes:— The basidiomata of this species are so fragile that our attempts to observe its natural structure of lamella trama and pileus squamules were failed.

The genus Leucocoprinus is similar to Leucoagaricus , but can be distinguished by the pileus with variform squamules composed of perpendicular to interwoven hyphae, frequent presence of colour changes after touched, damaged or dried, usually thicker basidiospore wall, monomorphous to polymorphous basidia, sometimes differentiated hymenial physalides, cheilocystidia without crystals on the apex, and the habit of occurring in various habitats and occasionally symbiotic with attine ant species ( Yang et al. 2024a). Leucocoprinus phantasmaticus conforms the morphological concept of Leucocoprinus .

In the current phylogeny ( Fig. 7), with more rpb2 and tef-1α sequences contributed by Li et al. (2025) and this study, the monophyly of Leucoagaricus plus Leucocoprinus s. str., Leucocoprinus s. str. itself, and Macropsalliota, all gained stronger support as compared with the earlier study Yang et al. (2024a). Notably, Leucoagaricus and Leucocoprinus s. str. still each form a distinct clade and have morphological characteristics to be recognized as separate genera, and the statistical support values for Leucoagaricus itself and Leucocoprinus s. str. itself are both greater than that for the common threshold of Leucoagaricus plus Leucocoprinus s. str.

The phylogenetic analysis by Li et al. (2025) using the same four loci as those in this study resolved Candelolepiota and Macropsalliota into a single clade, and Leucocoprinus s. str. into two separate clades. Such results could not be reproduced in our study, and are likely caused by overtrimmed alignments and insufficient four-locus data for Candelolepiota and Macropsalliota.

Amanitaceae E.-J. Gilbert

Amanita huangpuensis Kun L. Yang, Jia Y. Lin, Zhen-Chao Liu & Zhu L. Yang , sp. nov. ( Figs 9–10; Table 6; Supplements 2 and 7)

Registration identifier FN572706

Etymology:— Referring to the type locality.

Diagnosis:— Differing from Amanita elephas , A. indica and other similar species in A. pseudoporphyria complex by a combination of small to medium-sized basidiomata producing non-greyish stipe with a superior annulus and whitish to greyish volval remnants around stipe base, with volval remnants usually retained on pileus, and subglobose, broadly ellipsoid to ellipsoid basidiospores sized 6–7.5 × 5–6 µm.

Type:— CHINA. Guangdong Province: Guangzhou City, Huangpu District, Jiangdong Village , Boluoshan Hill , 23°11’36”N, 113°32’25”E, elevation 150 m, May 14, 2023, Kun L . Yang, Zhen-Chao Liu & Jia Y . Lin, K23215 ( HKAS147743 , holotype (nrLSU: PV616958; rpb2: PV642288; tef-1α: PV642306 ); HTBM0730 , isotype) .

Description:— Basidiomata small to medium-sized; odour fungal; taste unknown. Pileus 20–52 mm in diameter, hemispherical, plano-convex to applanate, sometimes slightly umbonate at center, light grey (#F1F0F0), dark heather brown (#9D917B), soy-sauce brown (#695B50) to duck blood red (#564843), sometimes with pale meat brown (#E4C8BB) tinges, radially forming a mottled pattern; volval remnants usually retained, flaky, pure white (#FFFFFF), ceramic white (#FEFEFA) to merino white (#F9F5EC); margin not striate, appendiculate with tiny fibers. Lamellae emarginate, adnexed to free, crowded, pure white (#FFFFFF), ceramic white (#FEFEFA) to cream orange (#FEF3CE), with a serrate edge, interspersed with abundant, attenuate lamellulae. Stipe 26–89 mm long, 5–8 mm thick, tapering upwards but with apex slightly expanded, clavately bulbous at base, pure white (#FFFFFF), ceramic white (#FEFEFA) to alabaster white (#F9F9F9), covered with fibrous squamules more abundant below the annulus; volval remnants membranous, with free limb up to 14 mm high, pure white (#FFFFFF), ceramic white (#FEFEFA), merino white (#F9F5EC) to light grey (#F1F0F0), with both sides concolorous. Annulus superior, membranous, easily broken and fugacious, concolorous with the stipe. Context pure white (#FFFFFF), ceramic white (#FEFEFA) to merino white (#F9F5EC), without a colour change after damaged.

Basidiospores {40/3/2} 6–7.5 [6.83 ± 0.40, 7.00] × 5–6 [5.46 ± 0.38, 5.50] µm, Q = 1.08–1.40 (1.50) [1.26 ± 0.12, 1.27], subglobose, broadly ellipsoid to ellipsoid, thin-walled, smooth, nearly colourless, amyloid, with a small apiculus. Basidia 29–40 × 9.5–11 μm, clavate, mostly four-spored, rarely two- or three-spored, thin-walled, nearly colourless. Subhymenium composed of 1–3 layers of subglobose to ellipsoid, thin-walled, nearly colourless inflated cells sized 5.5–15 × 5–10 µm. Pleurocystidia absent. Lamella trama bilateral, compact, composed of a mediostratum by 2–10 µm wide, thin-walled to slightly thick-walled, nearly colourless, sometimes thromboplerous and slightly yellowish, moderately branching hyphae, and two lateral strata each diverging at an angle of about 30 to 65 degree by abundant hyphae 2–9 µm wide, thin-walled to slightly thick-walled, nearly colourless, sometimes thromboplerous and slightly yellowish, moderately to frequently branching, and abundant inflated cells sized 75–100 × 10–15 µm, elongate to clavate, interjacent or terminal, thin-walled to slightly thick-walled, nearly colourless. Inflated cells on lamella edge sized 18–40 × 10–22 µm, subglobose, ellipsoid to elongate or clavate, thin-walled, nearly colourless. Volval remnants on pileus radially subregular to irregular, not to slightly gelatinized in trama, strongly gelatinized at the bottom, composed of abundant hyphae 2–12 µm wide, thin-walled to slightly thick-walled, nearly colourless, moderately compact to compact, moderately to frequently branching, and scarce inflated cells sized 33–80 × 20–65 µm, subglobose to ellipsoid, interjacent or terminal, slightly thick-walled, nearly colourless. Pileipellis radially subregular and moderately gelatinized at the base, becoming more irregular and strongly gelatinized towards the surface, composed of 2–11 µm wide, thin-walled, slightly brownish, moderately compact to compact, moderately branching hyphae without distinct layer differentiation. Volval remnants around stipe base radially subregular, moderately gelatinized at the inner surface, becoming not gelatinized towards the outer surface, without distinct layer differentiation, composed of abundant hyphae 1.5–12 µm wide, thin-walled, nearly colourless, sometimes thromboplerous and slightly yellowish, moderately compact to compact, moderately branching, and scarce inflated cells sized 50–62 × 24–38 µm, ellipsoid to elongate or clavate, interjacent or terminal, thin-walled to slightly thick-walled, nearly colourless. Annulus composed of radially irregular, abundant inflated cells sized 9–35 × 7–30 µm, subglobose to ellipsoid, slightly thick-walled, nearly colourless, loose to moderately compact, and scarce to moderately abundant hyphae 2–9 µm wide, thin-walled to slightly thick-walled, nearly colourless, moderately to frequently branching. Clamp connections absent.

Habit and distribution:— Solitary to gregarious, on soil, in southern subtropical monsoon forests dominated by plants of Fagaceae . Currently known from China.

Other collections examined:— CHINA. Guangdong Province: Guangzhou City, Huangpu District, Jiangdong Village, Boluoshan Hill, 23°11’36”N, 113°32’25”E, elevation 150 m, August 21, 2023, Zhen-Chao Liu, Jia Y. Lin & Kun L. Yang, S23475 (HTBM1586); same location, August 23, 2023, Zhen-Chao Liu, Jia Y. Lin & Kun L. Yang, L23294 (HTBM1275). Collections of other species sequenced for phylogenetic analyses: Amanita atrobrunnea ( CHINA. Guangdong Province: Guangzhou City, Huangpu District, Jiangdong Village, Boluoshan Hill, 23°11’36”N, 113°32’25”E, elevation 150 m, June 24, 2023, Zhen-Chao Liu, Jia Y. Lin & Kun L. Yang, S23217 (HTBM0460)); A. caojizong ( CHINA. Zhejiang Province: Hangzhou City, West Lake District, Hangzhou Botanical Garden, exact location unknown, July 10, 2023, Two Crisp Hash Browns & Kun L. Yang, L23165 (HTBM1117); Hunan Province: Changsha City, exact location unknown, July 10, 2023, Wu-Ping Luo & Kun L. Yang, L23167 (HTBM1119); Boundary of Guangdong and Fujian Provinces, exact location unknown, May 5, 2023, Kun L. Yang, kindly donated by anonymous net friends, S2353 (HTBM1641); Zhejiang Province: exact location unknown, July 7, Bin Chen & Kun L. Yang, S23297 (HTBM1008); Zhejiang Province: Ningbo City, Fenghua District, exact location unknown, July 21, 2023, Bin Chen & Kun L. Yang, S23316 (HTBM1027); Zhejiang Province: exact location unknown, June 30, 2023, Bin Chen & Kun L. Yang, S23257 (HTBM0500)); A. manginiana ( CHINA. Zhejiang Province: exact location unknown, July 7, 2023, Bin Chen & Kun L. Yang, S23295 (HTBM1006) & S23296 (HTBM1007)); A. modesta ( CHINA. Guangdong Province: Guangzhou City, Huangpu District, Jiangdong Village, Boluoshan Hill, 23°11’36”N, 113°32’25”E, elevation 150 m, August 21, 2023, Zhen-Chao Liu, Jia Y. Lin & Kun L. Yang, S23495 (HTBM1606)); A. pseudoporphyria ( CHINA. Zhejiang Province: Hangzhou City, West Lake District, Hangzhou Botanical Garden, exact location unknown, July 10, 2023, Two Crisp Hash Browns & Kun L. Yang, L23166 (HTBM1118); Zhejiang Province: Ningbo City, Jiangbei District, Jianghu Lake Reservoir, exact location unknown, July 21, 2023, Bin Chen & Kun L. Yang, S23315 (HTBM1026)).

Phylogeny:— According to the BLAST search of nrLSU sequences in GenBank ( NCBI 2025), our collections HKAS147743, HTBM1275 and HTBM1586 represent a single species of Amanita sect. Roanokenses . Therefore, a phylogeny was inferred to classify our collections with representative collections of known species of Amanita sect. Roanokenses and our collections as the ingroup, and a collection of Amanita sect. Amidella as the outgroup according to a recent study on phylogeny of Amanita ( Cui et al. 2018) , using five loci (ITS, nrLSU, rpb2, tef-1α & β-tub) necessary for resolving the phylogeny of this group ( Table 6; Supplement 7). In the result, the ML and BI topologies are almost identical, only with a minimal variation in statistical support, so only the tree inferred from the ML analysis is displayed ( Fig. 9). Our collections HKAS147743, HTBM1275 and HTBM1586 form an undescribed clade nested within the Amanita pseudoporphyria species complex with significant support.

Notes:— Species of the Amanita pseudoporphyria complex are very similar to each other, mostly exhibiting a radially mottled greyish brown pileus with whitish flaky volval remnants (when present), whitish lamella, whitish annulus, and whitish stipe with whitish volval remnants forming a distinct limb at the base. Many of these species probably have similar ecological niches and we frequently observe several different species occurring together ( Fig. 10). Although a species in this complex, namely A. caojizong , is known as edible for the moment ( Cui et al. 2018), we suggest not to consider it for food due to the great difficulty in distinguishing poisonous species with similar morphology. For example, the collection HTBM1117 identified as A. caojizong and HTBM1118 identified as A. pseudoporphyria (causing acute kidney injury) were found from two sites just over 10 m apart, and were indistinguishable in macromorphology ( Fig. 10).

In the current phylogeny ( Fig. 9), A. huangpuensis nests with two species already known as poisonous, A. neoovoidea and A. pseudoporphyria , both causing acute kidney injury, in a significantly supported subclade, and thus A. huangpuensis may also be poisonous. Interestingly, A. caojizong known as edible separates outside this subclade, with its exact position unresolved.

in bold. Unavailable items are indicated with -. Holotypes are indicated by HT.

Entolomataceae Kotl. & Pouzar

Clitopilus zoysiae Kun L. Yang, Jia Y. Lin & Zhu L. Yang , sp. nov. ( Figs 11–12; Table 7; Supplements 2 and 8) Registration identifier FN572707

Etymology:— Referring to the host.

Diagnosis:— Differing from other species of Clitopilus sect. Scyphoides by the habit of growing on living or dead stems of Zoysia plants.

Type:— CHINA. Guangdong Province: Guangzhou City, Haizhu District, track area in Pazhou Pagoda Station of Guangzhou Tram Haizhu Line 1, 23°06’06”N, 113°22’17”E, elevation 5 m, August 20, 2024, Jia Y . Lin & Kun L . Yang, L23287 ( HKAS147744 , holotype ( ITS: PV620874 ; nrLSU: PV616939; rpb2: PV642287; tef-1α: PV642305 ); HTBM1268 , isotype) .

Description:— Basidiomata tiny, without a distinct colour change after damaged; odour indistinct; taste unknown. Pileus reniform, 4–9 mm wide, extending 3–6 mm outwards the lateral stipe, pure white (#FFFFFF) to ceramic white (#FEFEFA), smooth to slightly felty. Lamellae subdistant to nearly crowded, ceramic white (#FEFEFA) to light cream orange (#FFF5E0), with an entire edge, interspersed with abundant lamellulae. Stipe lateral and reduced.

Basidiospores {40/4/2} 5.5–6.5 [6.06 ± 0.32, 6.00] × 3–3.5 [3.30 ± 0.24, 3.50] µm, Q = 1.71–2.00 [1.84 ± 0.11, 1.71], oblong and angled with 8–12 inconspicuous longitudinal ridges under LM, also with minute transverse ridges under SEM but invisible under LM, thin-walled, smooth, nearly colourless, with a small apiculus. Basidia 14–21 × 4.5–6 μm, clavate, four-spored, thin-walled, nearly colourless. Lamella trama subregular to irregular, composed of 3–6 µm wide, thin-walled, nearly colourless, compact, moderately branching hyphae. Cheilocystidia absent. Pleurocystidia absent. Pileipellis composed of a rectocutis to tomentum by 2–3.5 µm wide, thin-walled, nearly colourless, compact to moderately compact, moderately branching hyphae. Clamp connections absent.

Habit and distribution:— Gregarious, on living or dead stems of lawngrass ( Zoysia japonica ), in urban areas. Currently known from China.

Other collections examined:— CHINA. Guangdong Province: Guangzhou City, Haizhu District, Pazhou Pagoda Park , 23°06’05”N, 113°22’16”E, elevation 5 m, August 20, 2024, Jia Y . Lin & Kun L . Yang, L23284 ( HTBM1265 ) & L23286 ( HTBM1267 ) .

Phylogeny:— According to the BLAST search of ITS and nrLSU sequences in GenBank ( NCBI 2025), our collections HKAS147744, HTBM1265 and HTBM1267 represent a single species of Clitopilus sect. Scyphoides . Therefore, a phylogeny was inferred to classify our collections with representative collections of known species of Clitopilus sect. Scyphoides and our collections as the ingroup, and a collection of Clitopilus sect. Clitopilus as the outgroup according to a recent study on phylogeny of Clitopilus ( Jian et al. 2020) , using four loci (ITS, nrLSU, rpb2 & tef-1α) necessary for species identification in this group ( Table 7; Supplement 8). In the result, the ML and BI topologies are almost identical, only with a minimal variation in statistical support, so only the tree inferred from the ML analysis is displayed ( Fig. 11). Our collections HKAS147744, HTBM1265 and HTBM1267 form an undescribed clade with significant support, and this clade is sister to the lectotype of C. giovanellae with significant support.

Notes:— Clitopilus sect. Scyphoides is typically characterized by small-sized, omphaloid to pleurotoid basidiomata, basidiospores with more than five, distinct or obscure longitudinal ridges and minute transverse ridges, and the habit of growing on soil or deadwood ( Jian et al. 2020). Clitopilus zoysiae generally fits its morphological concept, but is distinct in the habit of growing on living or dead stems of Zoysia japonica .

Another species in sect. Scyphoides known to have a similar habit, is the recently described C. parasiticus ( Jian et al. 2025) . However, C. parasiticus looks polymorphic on nutritional modes, exhibiting both parasitic behavior on leaves of Dryopteris and Oplismenus , and saprotrophic growth in soil environments. Jian et al. (2025) further noticed that the basidiospores from saprotrophic basidiomata were larger than those from parasitic ones, implying that C. parasiticus may be better suited to a soil habitat.

In the current phylogeny ( Fig. 11), C. zoysiae is sister to the lectotype of C. giovanellae with significant support. However, C. giovanellae processes omphaloid basidiomata saprotrophic on sandy soils, and has never been reported as parasitic on plants ( Bresadola 1881, Singer 1942, Moreno et al. 2007).

In our morphological studies, we frequently observed the conidiospores of Curvularia species in mountings of Clitopilus zoysiae tissues ( Fig. 12h–i). Curvularia species are known as pathogens of Poaceae plants ( Tan et al. 2018), and the interactions among Clitopilus , Curvularia and Zoysia may be worth further studying.

in bold. Unavailable items are indicated with -. Holotypes and lectotypes are indicated by HT and LT, respectively.

Lycoperdaceae F. Bercht. & J. Presl

Calvatia scandens Kun L. Yang, Jia Y. Lin, Z. H. Zhang, Zhen-Chao Liu & Zhu L. Yang , sp. nov. ( Figs 13–14; Table 8; Supplements 2 and 9)

Registration identifier FN572708

Etymology:— Referring to its habit of clinging on stones, sticks or bricks by rhizomorphs.

Diagnosis:— Differing from other known Calvatia species by having tiny basidiomata with a habit of clinging on stones, sticks or bricks by rhizomorphs, ellipsoid basidiospores, and capillitia strongly encrusted with granules when mature.

Type:— CHINA. Zhejiang Province: Lishui City, Suichang County, Jiulongshan Naure Reserve , 28°22’31”N, 118°51’03”E, elevation 613 m, July 27, 2023, Z. H . Zhang, Qian-Qian Liu & Zhan-Sheng Tang, ZhangZH98- S23614 ( HKAS147745 , holotype ( ITS: PV620869 ; nrLSU: PV616926 ); HTBM1725 , isotype) .

Description:— Basidiomata tiny, 3–12 mm in diameter, 3–7 mm in height, sessile, obpyriform, turbinate, subglobose to tuberiform, each issued by a single rhizomorph, without a distinct colour change after damaged or dried; odour fungal; taste unknown. Exoperidium very thin, pure white (#FFFFFF), ceramic white (#FEFEFA), zircon white (#F3F5FD) to star white (#FBFBEB), nearly smooth, finely granular to felty. Endoperidium very thin, concolorous with or lighter than exoperidium. Gleba cottony, concolorous with peridium at first, becoming straw brown (#C4B179) to dull olive brown (#736938).

Basidiospores {40/4/2} (4.5) 5–5.5 [5.14 ± 0.27, 7.00] × 3–4 [3.45 ± 0.27, 3.50] µm including echinulate ornamentation about 0.5 µm high, Q = 1.38–1.67 [1.49 ± 0.10, 1.43], mostly ellipsoid, rarely oblong, thick-walled, yellowish to brownish, with a small apiculus. Immature basidia without sterigmata 4–9 × 3–4 μm, clavate to cylindrical, thin-walled, nearly colourless. Mature basidia with sterigmata not observed. Gleba trama composed of generative hyphae 1.5–4.5 μm wide, thin-walled, nearly colourless, moderately compact to compact, moderately to frequently septate, moderately to frequently branching, subregularly arranged, becoming dissolved following maturity, and capillitia 2.5–5 μm wide, Lycoperdon - type to Calvatia - type, rarely septate, rarely branching, thick-walled, nearly colourless and smooth at first, becoming yellowish to brownish and strongly encrusted with granules following maturity. Exoperidium composed of a spherocystoderm to conioderm by inflated cells sized 8–29 × 9.5–32 µm, subglobose to oblate ellipsoid, thin-walled to slightly thick-walled, nearly colourless. Endoperidium radially subregular to irregular, composed of 2.5–5 μm wide, thin-walled, nearly colourless, compact, moderately to frequently branching hyphae without clamp connections.

Habit and distribution:— Gregarious, clinging on mossy stones, sticks or bricks by strong rhizomorphs, in subtropical forests or urban areas. Currently known from China.

Other collections examined:— CHINA. Zhejiang Province: Lishui City, Suichang County, Jiulongshan Naure Reserve, 28°25’15”N, 118°49’18”E, elevation 600 m, July 27, 2023, Z. H. Zhang & Zhan-Sheng Tang, ZhangZH101- S23615 (HTBM1726). Guangdong Province: Guangzhou City, Huangpu District, Jiangdong Village, Boluoshan Hill, 23°11’36”N, 113°32’25”E, elevation 150 m, August 26, 2024, Zhen-Chao Liu, Jia Y. Lin & Kun L. Yang, S24033 (HTBM2113) & S24034 (HTBM2114). Collections of other species sequenced for phylogenetic analyses: Calvatia boninensis ( CHINA. Guangdong Province: Guangzhou City, Huangpu District, Muqiang Reservoir, 23°13’N, 113°28’E, elevation 46 m, May 27, 2023, Yu-Rong Liang, S2370-ACE55 (HTBM0313); Guangdong Province: Shenzhen City, Nanshan District, exact location unknown, June 19, 2023, Li-Ping Tang & Kun L. Yang, S23163 (HTBM0406)); C. cyathiformis ( CHINA. Jiangxi Province: Shangrao City, Qianshan County, exact location unknown, June 10, 2023, Dan-Ying Liu & Kun L. Yang, S23116 (HTBM0359); Jiangsu Province: Nanjing City, Jiangning District, exact location unknown, June 26, 2023, Ze-En Du & Kun L. Yang, S23209 (HTBM0452)); C. fenzlii ( CHINA. Jiangsu Province: Nanjing City, Xuanwu District, Zijinshan Mountain, 32°04’58”N, 118°50’13”E, elevation 39 m, June 29, 2023, Xia Chen, Jia Y. Lin & Kun L. Yang, S23360 -Chenxia469 (HTBM1471)); C. gigantea ( CHINA. Hebei Province: Chengde City, Chengde County, Xiabancheng Town, June 16, 2023, Su-Min Liu, Chun-Ling Ma & Kun L. Yang, S23160 (HTBM0403)); C. holothurioides ( CHINA. Guangdong Province: Guangzhou City, Tianhe District, South China Agricultural University, Arboretum, 23°09’27”N, 113°21’22”E, elevation 40 m, June 8, 2023, Jia Y. Lin & Kun L. Yang, L2373 (HTBM0906); same location, September 12, 2023, Jia Y. Lin, L23420 (HTBM1401); Guangdong Province,:Guangzhou City, Baiyun District, 23°13’N, 113°18’E, elevation 85 m, June 20, 2023, Yu-Rong Liang, S23177 -ACE65 (HTBM0420) & S23178 -ACE69 (HTBM0421); Guangdong Province: Guangzhou City, Huangpu District, Jiangdong Village, Boluoshan Hill, 23°11’36”N, 113°32’25”E, elevation 150 m, June 13, 2023, Zhen-Chao Liu & Kun L. Yang, S23137 (HTBM0380); same location, June 24, 2023, Zhen-Chao Liu & Kun L. Yang, S23193 (HTBM0436); Jiangsu Province: Nanjing City, Xuanwu District, Zijinshan Mountain, 32°04’53”N, 118°50’15”E, elevation 56 m, July 12, 2023, Xia Chen, Jia Y. Lin & Kun L. Yang, S23371 -Chenxia545 (HTBM1482)); C. nipponica ( CHINA. Henan Province: Xinxiang City, Yanjin County, Shipogu Town, June 8, 2023, Meng Yu & Kun L. Yang, S23114 (HTBM0357)); C. pyriformis ( CHINA. Guangdong Province: Guangzhou City, Tianhe District, South China Agricultural University, Stadium of Taishan District, 23°09’22”N, 113°21’33”E, elevation 40 m, June 11, 2023, Jia Y. Lin & Kun L. Yang, L2379 (HTBM0912); Guangdong Province: Guangzhou City, Panyu District, South China University of Technology, exact location unknown, June 28, 2023, Jia Y. Lin & Kun L. Yang, K23316 (HTBM0831); same location, June 8, 2023, Fei-Yang Chen & Kun L. Yang, kindly donated by anonymous net friends, S23112 (HTBM0355); Guangdong Province: Foshan City, Nanhai District, Foshan University (Xianxi Campus), exact location unknown, June 10, 2023, Shi-Ting Liang & Kun L. Yang, S23118 (HTBM0361); Guangdong Province: Guangzhou City, Conghua District, Guangzhou Nanfang College, exact location unknown, June 14, 2023, Jing Qiu & Kun L. Yang, S23155 (HTBM0398)); C. shennongjiaensis ( CHINA. Chongqing Municipality, Beibei District, Southwest University, Love Hill besides No. 1 Stadium, June 8, 2023, Xie Xie & Kun L. Yang, S2399 (HTBM0342); Hunan Province: Changsha City, exact location unknown, July 10, 2023, Wu-Ping Luo, Jia Y. Lin & Kun L. Yang, L23171 (HTBM1123); Sichuan Province: Dazhou City, Tongchuan District, Pujia Town, exact location unknown, June 12, 2023, Xian-Mei Yang & Kun L. Yang, S23132 (HTBM0375); Guizhou Province: Guiyang City, Guanshanhu District, Baihuahu Forest Park, exact location unknown, June 12, 2023, June 25, 2023, Antistar & Kun L. Yang, S23207 (HTBM0450); Anhui Province: exact location unknown, June 30, 2023, Xin-Tong Gao & Kun L. Yang, S23264 (HTBM0507); Jiangsu Province: Nanjing City, Xuanwu District, Zijinshan Mountain, 32°03’23”N, 118°51’52”E, elevation 73 m, June 12, 2023, Xia Chen & Kun L. Yang, S23170 -ChenXia334 (HTBM0413); Jiangsu Province: Nanjing City, Xuanwu District, Zijinshan Mountain, 32°05’00”N, 118°50’23”E, elevation 37 m, June 29, 2023, Xia Chen, Jia Y. Lin & Kun L. Yang, S23359 -ChenXia460 (HTBM1470); Jiangsu Province: Nanjing City, Xuanwu District, Zijinshan Mountain, 32°03’24”N, 118°49’51”E, elevation 39 m, July 2, 2023, Xia Chen, Jia Y. Lin & Kun L. Yang, S23361 -ChenXia487 (HTBM1472); Jiangsu Province: Nanjing City, Xuanwu District, Zijinshan Mountain, 32°03’05”N, 118°50’08”E, elevation 36 m, July 6, 2023, Xia Chen, Jia Y. Lin & Kun L. Yang, S23364 - ChenXia514 (HTBM1475); Jiangsu Province: Nanjing City, Xuanwu District, Zijinshan Mountain, 32°03’19”N, 118°49’51”E, elevation 41 m, July 6, 2023, Xia Chen, Jia Y. Lin & Kun L. Yang, S23365 -ChenXia522 (HTBM1476); Jiangsu Province: Nanjing City, Xuanwu District, Zijinshan Mountain, 32°03’05”N, 118°50’04”E, elevation 35 m, July 13, 2023, Xia Chen, Jia Y. Lin & Kun L. Yang, S23372 -ChenXia548 (HTBM1483)).

Phylogeny:— According to the BLAST search of nrLSU sequences in GenBank ( NCBI 2025), our collections HKAS147745, HTBM1726, HTBM2113 and HTBM2114 represent a single species of Calvatia . Therefore, a phylogeny was inferred to classify our collections with representative collections of known species of Calvatia and our collections as the ingroup, and a collection of Disciseda as the outgroup according to a recent study on phylogeny of Lycoperdaceae ( Li et al. 2024a) , using the ITS locus available for most collections of this group ( Table 8; Supplement 9). In the result, the ML and BI topologies are almost identical, only with a minimal variation in statistical support, so only the tree inferred from the ML analysis is displayed ( Fig. 13). Our collections HKAS147745, HTBM1726, HTBM2113 and HTBM2114 form an undescribed clade in sect. Calvatia .

Notes:— Five sections proposed in the genus Calvatia in previous studies based on morphology, namely sect. Calvatia , sect. Cretacea , sect. Hippoperdon , sect. Langermannia and sect. Sculpta , are recovered in our phylogeny ( Fig. 13). The new species C. scandens described here is recognized in sect. Calvatia , a section characterized by furfuraceous exoperidum and olivaceous mature gleba ( Kreisel 1992). It generally fits the morphological concept of this section, but is quite distinct in the tiny size of basidiomata, the habit of clinging on substrate by rhizomorphs and the capillitia strongly encrusted with granules that differ from any other taxa previously known in this section.

Only two of our collections, HKAS147745 and HTBM2113, contain immature basidiomata of C. scandens . However, all these immature basidiomata processes very immature basidia without sterigmata (e.g., Fig. 14g), and no recognizable basidia could be found in more mature basidiomata. As a result, the characteristics from mature basidia of this species are unknown.

in this study are in bold. Unavailable items are indicated with -. Holotypes are indicated by HT.

Lyophyllaceae Jülich