Research Article |
Corresponding author: En-jing Tian ( tianenjing66@163.com ) Academic editor: María P. Martín
© 2024 Jia-hui Huang, Chun-yu Lei, Ya-lun Shen, En-jing Tian.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Huang J-h, Lei C-y, Shen Y-l, Tian E-j (2024) Two new species of Pholiota (Agaricales, Strophariaceae) from the southwest of China. MycoKeys 109: 171-185. https://doi.org/10.3897/mycokeys.109.133879
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Two new mushroom species from Southwest China, Pholiota cylindrospora and P. subterrestris, are described in this study. Pholiota cylindrospora is characterized by its dry pileus with slightly recurved and triangular scales, cylindrical basidiospores, and two types of pleurocystidia—leptocystidia and chrysocystidia—as well as its growth on soil. Pholiota subterrestris is identified by a brownish-orange pileus with numerous brown fibrillose scales; pale brown lamellae with even edges; a stipe covered with recurved fibrillose scales; elliptical spores with a distinct but small germ pore; and pleurocystidia containing typical amorphous refractive inclusions of chrysocystidia. Both species are described and illustrated, and a phylogenetic analysis of a multigene dataset (ITS + 28S) is presented. Morphological and phylogenetic analyses confirm that P. cylindrospora and P. subterrestris are distinct from the other Pholiota species, and both belong to the subgenus Pholiota. A key to the species of subgenus Pholiota from China is provided.
Chrysocystidia, morphology, phylogeny, Subgenus Pholiota, taxonomy
Pholiota (Fr.) P. Kumm. is distributed worldwide, especially in the northern temperate zone, and currently contains approximately 150–160 species (
Pholiota is an important and stable genus of the family Strophariaceae. Nevertheless, its previously proposed infrageneric classification systems vary owing to differing concepts of the genus, from as many as seven subgenera to three (
This study discovered two new species from the Southwest of China, belonging to subgenus Pholiota, based on morphological examination and phylogenetic analyses. The results are presented in the following sections.
Fresh specimens were collected from Southwest China and deposited at the
Herbarium of Mycology of Jilin Agricultural University (
Basidiospores were measured and examined according to the methods of
Abbreviations: “L” denotes the number of lamellae extending to the stipe; “I” indicates the number of lamellae between each adjacent pair; “Q” represents the length/breadth ratio of basidiospores; “Qm” refers to the mean value of “Q” measurements across all studied collections ± the sample standard deviation.
DNA was extracted from 10 to 20 mg of ground, dried basidiome tissue using a Plant Genomic DNA Kit (Tiangen Biotech Co., Ltd, Beijing). The internal transcribed spacer region (ITS) of Nuc rDNA was amplified and sequenced using the primers ITS1 (TCCGTAGGTGAACCTGCGG) and ITS4 (TCCTCCGCTTATTGATATGC) (
Species | Specimen-voucher (Herbarium) | Origin | GenBank accession numbers | |
---|---|---|---|---|
ITS | 28S | |||
Pholiota adiposa | ET37 (HMJAU37520) | China, Jilin | MN209721 | MN251112 |
P. agglutinata | AHS60691 (TENN-F-028806, isotype) | USA, Idaho | MN149356 | MN251113 |
P. aurivella | CBS 262.32 | Netherlands | MH855317 | – |
P. aurivella | SJ84131 | Sweden | AF195603 | – |
P. aurivella | ET27 (HMJAU37516) | China, Jilin | MN209728 | MN251116 |
P. aurivella | ET42 (HMJAU37521) | China, Jilin | MN209729 | MN251117 |
P. aurivella | ET19 (HMJAU37425) | China, Inner Mongolia | MN209771 | MN251154 |
P. avellaneifolia | AHS59589 (TENN-F-028809 isotype) | USA, Idaho | MN209731 | MN251120 |
P. baeosperma | TFB7383 (TENN-F-054431) | Chile | MG735312 | – |
P. baeosperma | RHP8315 (TENN-F-054993) | Argentina | KY559332 | – |
P. baptistii | ET542 TENN-F-028810 (isotype) | USA, Idaho | MN149364 | – |
P. brunnescens | AHS3525 (MICH 11657, holotype) | USA, Oregon | MG735292 | – |
P. brunnescens | PBM3057 (TENN-F-063855) | USA, California | MG735314 | – |
P. caespitosa | TENN-F-015908 (holotype) | USA, Tennessee | NR119908 | – |
P. carbonaria | AHS9500 (MICH 11663, holotype) | USA, California | MG735288 | – |
P. castanea | DPL7769 (TENN 071878) | USA, Texas | MH016952 | – |
P. castanea | TENN 020269 (holotype) | USA, Tennessee | HQ222025 | – |
P. chocenensis | PRM 895066 (holotype) | Czech Republic | NR_155622 | – |
P. conissans | SJ96017, FCUG1273 | Sweden | AF195606 | – |
P. conissans | CBS 175.47 | France | MH856205 | – |
P. conissans | CBS 243.50 | France | MH856603 | – |
P. conissans | voucher 395 | Italy | JF908575 | – |
P. conissans | voucher 6610 | Italy | JF908581 | – |
P. cylindrospora | ET-Ti2(holotype) | China, Yunnan | PQ013666 | PQ013732 |
P. cylindrospora | ET-yun2 | China, Yunnan | PQ013686 | PQ013733 |
P. decorata | AHS54770 (TENN-F-028816) | USA, Idaho | MN209734 | – |
P. ferrugineolutescens | TENN-F-028807 (isotype) | USA, California | HQ222026 | – |
P. flavescens | TENN-F-01309) (isotype) | USA, Tennessee | MN209735 | MN251124 |
P. fulviconica | LRH28818 (TENN-F-028818) | USA, Idaho | MN209738 | MN251126 |
P. fulviconica | AHS65898 (TENN-F-028820, isotype) | USA, Idaho | MN209739 | MN251127 |
P. fulvozonata | AHS73887 (MICH 5316, holotype) | USA, Idaho | MG735290 | – |
P. gallica | PRM 933232 | France | LN889967 | – |
P. gallica | MPU 3478 (holotype) | France | HG007988 | – |
P. gummos | SJ84095, FCUG1254 | Sweden | AF195605 | – |
P. gummos | CBS 216.39 | Portugal | MH855985 | – |
P. gummos | CBS 210.48 | Portugal | MH856313 | – |
P. highlandensis | NYS 1468.1 (isotype) | USA, New York | MH016956 | – |
P. highlandensis | PBM4085 (TENN-F-071544) | USA, Tennessee | MG735310 | – |
P. humii | AHS58633 (TENN-F-028822, isotype) | USA, Idaho | MN209740 | MN251128 |
P. jahnii | SJ83118, FCUG1061 | Sweden | AF195604 | – |
P. kodiakensis | TENN-F-028804 (isotype) | USA, Alaska | MN149360 | – |
P. lenta | ET33 (HMJAU37519) | China, Jilin | MN209742 | MN251130 |
P. lenta | PBM4233 (TENN-F-074640) | USA, N. Carolina | MN209743 | MN251131 |
P. limonella | ET28 (HMJAU37345) | China, Inner Mongolia | MN209747 | MN251135 |
P. limonella | MF68025 (TENN-F-074669) | USA, Tennessee | MN209748 | MN251136 |
P. limonella | ET11 (HMJAU37362) | China, Jilin | MN209741 | MN251129 |
P. limonella | ET13 (HMJAU37514) | China, Jilin | MN209746 | MN251134 |
P. lubrica | ET29 (HMJAU37517) | China, Jilin | MN209751 | MN251139 |
P. lubrica | ET4 (HMJAU22678) | China, Jilin | MN209753 | MN251141 |
P. lundbergii | LL950724 | Sweden | AF195607 | – |
P. lurida | AHS66386 (TENN-F- 028770, isotype) | USA, Michigan | MN209757 | – |
P. luteobadia | AHS43222 (MICH 11688, holotype) | USA, Michigan | MG735289 | – |
P. marangania | HLepp856 (CANB 574576) | Australia | MG735320 | – |
P. melliodora | AHS68780 (TENN-F-028861, paratype) | USA, Oregon | MN209758 | – |
P. mixta | SJ96022 | Sweden | AF195609 | – |
P. mixta | PBM2499 (TENN-F-062357) | USA, Mass. | MH016953 | – |
P. molesta | AHS65008 (TENN-F-028830, isotype) | USA, Idaho | MG735296 | – |
P. molesta | JFA9246 (WTU 10719) | USA, Washington | MG735297 | – |
P. multicingulata | PBM3587 (TENN-F-066655) | Australia | MN209760 | – |
P. multicingulata | ET23 (HMJAU37414) | China, Yunnan | MN209761 | MN251146 |
P. nameko | ET7 (HMJAU37512 / TENN-F-074767) | China, Jilin | MN209762 | MN251147 |
P. nameko | ET10 (HMJAU22620 as Pholiota “microspora”) | China, Jilin | MN209759 | MN251145 |
P. occidentalis | AHS58470 (TENN-F-028874, paratype) | USA, Idaho | MN209765 | MN251150 |
P. olivaceodisca | TENN-F-017778 (holotype) | USA, Tennessee | NR119909 | – |
P. olivaceophylla | MICH 290502 (holotype) | USA, CA | KF878381 | – |
P. parvula | AHS42234 (TENN-F-028834, isotype) | USA, Michigan | MN149362 | – |
P. polychroa | PBM2866 (TENN-F-062649) | USA, Louisiana | MG735317 | – |
P. prolixa | AHS5027 (TENN-F-028838, isotype) | USA, Michigan | MN209766 | – |
P. rubronigra | AHS56192 (TENN-F-028840, isotype) | USA, California | MH016955 | – |
P. rufodisca | B925 (TENN-F-028869, paratype) | USA, New Mexico | MN209767 | – |
P. sp. | ET38 (HMJAU22691) | China, Jilin | MN209737 | MN251125 |
P. spumosa | ET12 (HMJAU37513) | China, Jilin | MN209776 | MN251159 |
P. spumosa | LRH12950 (TENN-F-012950 | USA, Tennessee | MN149361 | – |
P. squarrosa | PBM2735 (TENN-F-062547) | USA, Colorado | DQ494683 | DQ470818 |
P. squarrosa | AH-48188 | Spain | MF345959 | – |
P. squarrosa | SFC20140912-I01 | South Korea | KX773886 | – |
P. squarrosa | Voucher 28 | Germany | FR686575 | – |
P. squarrosa | ET26 (HMJAU37515) | China, Jilin | MN209777 | MN251160 |
P. squarrosa | ET15 (HMJAU37366) | China, Jilin | MN209778 | MN251161 |
P. squarrosoides | TENN-F-061728 | USA, Tennessee | FJ596877 | – |
P. squarrosoides | SFC20120814-45 | South Korea | KX773887 | – |
P. squarrosoides | JACA-MICO-00132 | Spain | MF345957 | – |
P. squarrosoides | TENN-F-061692 | USA, N. Carolina | FJ596860 | – |
P. squarrosoides | AH-48186 | France | MF345958 | – |
P. stratosa | AHS64684 (TENN-F-028845, isotype) | USA, Michigan | MN209779 | – |
P. subsaponacea | AHS74095 (MICH 5332, holotype) | USA, Idaho | MG735287 | – |
P. subterrestris | ET-072240 (holotype) | China, Guizhou | PQ013700 | PQ013730 |
P. subterrestris | ET-gui2 | China, Guizhou | PQ013701 | PQ013731 |
P. tennesseensis | TENN-F-018848 (holotype) | USA, Tennessee | NR_119910 | – |
P. terrestris | RAS371 (TENN-F-074807) | USA, Tennessee | MN209781 | – |
P. terrestris | iNat62825743 | USA,New York | OP681747 | – |
P. terrestris | iNat62849835 | USA,New York | MW644574 | – |
P. tetonensis | WGS3763 (TENN-F-028849, isotype) | USA, Wyoming | MN149367 | – |
P. velaglutinosa | AHS9285 (TENN-F-028851, isotype) | USA, Oregon | MH016954 | – |
P. virescens | HMJAU22498 (holotype) | China, Jilin | JF961378 | – |
P. virescentifolia | TENN-F-020591 (holotype) | USA, Tennessee | NR_119911 | – |
P. virgata | B763 (TENN-F-028832, paratype) | USA, New Mexico | MN209782 | – |
Stropharia acanthostipitata | CA01222011 (TENN-F-071898) | Dominican Republic | MG735313 | – |
S. acanthostipitata | JBSD 127401(holotype) | Dominican Republic | NR_156637 | MF882994 |
S. rugosoannulata | X-31 | USA | KC176328 | – |
Bayesian Inference (BI) analysis was conducted using MrBayes 3.2.7 (
Eight new sequences (four ITS and four 28S) from the two species were produced in this study (Table
In the phylogram (Fig.
Differs from other Pholiota species by dry pileus with slightly recurved and triangular scales, cylindrical basidiospores, and two types of pleurocystidia, including leptocystidia and chrysocystidia, as well as growing on soil.
China. • Yunnan, Baoshan City, Gaoligong Mountains, Baihualing; elev. 910 m; 25°03'33′′N, 98°49'18′′E; scattered on soil in broad-leaved forest; 30 November 2015; Shi-liang Liu 37432 (holotype:
Referring to the cylindric basidiospores.
Pileus 50–70 mm in diameter, convex to broadly convex, becoming nearly plane, with a low obtuse umbo; surface dry, pale orange to apricot (5A3–5B6), paler towards the margin, decorated with concentric, slightly recurved, triangular, light brown (6D7) scales. Context pale (2A2), odor and taste mild. Lamellae adnate, broad, moderately close, L = 38–42, I = 2–6, pallid at first, becoming brown (6D6), the edges waved. Stipe 45–60 mm long, 8–11 mm thick, central, equal to tapered towards the base, solid, ground color pallid, smooth to silky above an annular zone, towards the base with yellowish brown to brown (5D6–6D6) multizonate squamules or sometimes with scattered brown (6D5) fibrils, with white mycelium at the base.
Basidiospores (6.5–)7.0–9.5(–10.0) × 3.0–4.0 µm, Q = 1.75–2.83, Qm = 2.30, in face view oblong to cylindric, sometimes crooked, in profile cylindric to slightly inequilateral, wall smooth and thin, germ pore very minute to not evident, pale rusty to yellowish brown (6E8–5E8) in KOH, slightly paler in Melzer’s reagent. Basidia 19–27 × 4.9–6.5 µm, 4-spored, clavate, hyaline in KOH. Pleurocystidia of two types: 1) leptocystidia, 35–55 × 8.5–10 µm, clavate to subfusoid, thin-walled, smooth, content homogeneous, hyaline to pale yellowish brown to brownish (5D5–5E7) in KOH; 2) chrysocystidia, 39–60 × 9.5–12.5 µm, lageniform to clavate with a rostrate to mucronate apex, rarely forked near the apex, with an amorphous highly refractive inclusion, pale rusty to yellowish brown (6E8–5D5) in KOH. Cheilocystidia 23–39 × 6–11 µm, fusiform, clavate to subcapitate at apex, wall thin and smooth, content homogeneous, hyaline, pale yellow to yellow brownish (4A3–5C7) in KOH. Caulocystidia not observed. Gill trama of parallel hyaline to yellowish white (4A2) hyphae in KOH and with smooth walls, the cells inflated, up to 23 µm in diam. Pileipellis a cutis of brownish yellow to light brown (5C8–5D5) hyphae 4.5–11 µm in diam., thin-walled, slightly incrusted to asperulate. Clamp connections present in all the tissues.
Scattered on soil in broad-leaved forest in late autumn.
China. • Yunnan: Tengchong City, Yunfeng Mountain, on soil; 15 November 2019; Xiao-ming Zhang 37433 (
Cylindrical basidiospores are uncommon in the genus Pholiota, making this species easily distinguishable from the others. In addition, it is readily recognized because of its dry pileus with slightly recurved and triangular scales and two types of pleurocystidia, including leptocystidia and chrysocystidia, as well as its growth on soil.
The stipe with multizonate squamules of this species reminds one of Pholiota multicingulata Horak, but the latter has smaller basidiospores (6.5–8 × 4.5–5 µm) and lacks chrysocystidia (
Pholiota cylindrospora is similar to P. squarrosa (Vahl) P. Kumm. Both species have a dry pileus with obvious scales and two types of pleurocystidia: leptocystidia and chrysocystidia. However, the latter has smaller (6–7.5 × 3.8–4.5 µm) and elliptic basidiospores with a distinct germ pore and is wood-inhabiting (
Pholiota subterrestris is distinguished from the other species of the genus Pholiota by brownish orange pileus with numerous brown fibrillose scales, pale brown and moderately broad lamellae with even edges, stipe covered with recurved fibrillose scales, evanescent fibrillose annulus; elliptic spores with distinct but small germ pore, fusoid ventricose pleurocystidia with typical amorphous refractive inclusion of chrysocystidia.
China. • Guizhou: Bijie City, Nayong County, Dapingqing National Wetland Park; elev. 1990 m; 26°41'10′′N, 105°27'30′′E; scattered on soil at base of stump; 22 July 2020; En-jing Tian 37434 (holotype:
The epithet subterrestris refers to the similarity to Pholiota terrestris.
Pileus 10–35 mm in diameter, subhemispherical when young, with involute margin, slightly spread when mature, surface slightly viscid when wet, greyish orange to brownish orange (5B4–6C6), center dark, becoming paler towards the margin, covered with brown (6E7) fibrillose scales. Context thick, white to yellowish white (1A1–2A2), mild odor and taste. Lamellae decurrent to adnate or adnexed, moderately broad, close, L = 39–44, I = 3–7, pale yellow to light brown (4A3–5D6), edges even. Stipe 18–30 mm long, 4–7 mm thick, central, equal, solid, brownish orange (5C4) above, brown (6E5) below, covered with light brown (6D6) fibrillose squamulose, often compressed at the base
Basidiospores 5.4–6.1 × 3.6–4.2 µm, Q = 1.30–1.56, Qm = 1.47, smooth, in face view broadly elliptic to elliptic, in profile a little bean-shaped, apical pore distinct but small, containing oil-bearing droplet or irregular inclusions, greyish yellow (4B5) in KOH and light yellow (4A5) in Melzer’s reagent. Basidia 17.1–21.8 (–24.0) × 5.0–6.3 µm, 4-spored, clavate, hyaline in KOH, pale yellow (4A3) in Melzer’s reagent. Pleurocystidia of two types: 1) 33.3–46.7 × 5.5–9.5 µm, subclavate to fusoid ventricose with a subacute apex, wall thin and smooth, content homogeneous, blond to dark yellow (4C4–4C8) to hyaline in KOH; 2) chrysocystidia, 34.2–58 × 8.3–10.4 µm, fusoid to fusoid-ventricose, the neck often drawn out to a crooked filamentose projection with a subacute apex (up to 19.6 µm long), walls thin, smooth, with a refractive-amorphous inclusion, hyaline in KOH. Cheilocystidia abundant, (20.0–) 22.9–39.2 (–48.3) × 3.9–12.7 µm, clavate to clavate-capitate, wall thin, at times somewhat thickened in apex (about 1 µm), hyaline in KOH. Caulocystidia none observed. Gill trama of parallel hyphae, the cells inflated up to 27.5 µm, thin-walled, smooth, hyaline in KOH. Pileipellis hyphae hyaline to light brown (6D7), 3.7–10 µm diam., with encrusted walls. Content hyphae hyaline, 10–37.5 (–50) µm diam., cells inflated, smooth and thin-walled. Clamp connections present in all the tissues.
Scattered or gregarious on soil at the base of a stump or on buried wood in summer.
China. • Guizhou: Bijie City, Qixingguan District, Baima Mountain, gregarious on soil or on buried wood, 15 August 2022, Guang-cheng Cao 37435 (
This species is characterized by a brownish orange pileus with numerous brown fibrillose scales, pale brown lamellae with even edges, a stipe covered with recurved fibrillose scales, an evanescent fibrillose annulus, elliptic spores with a distinct but small germ pore, and fusoid ventricose pleurocystidia with typical amorphous refractive inclusions of chrysocystidia.
Pholiota subterrestris is similar to P. terrestris Overh., especially in terms of micro-characteristics, such as pleurocystidia, cheilocystidia, and basidiospores. However, it is easy to differentiate between the two species based on the macro characteristics. P. terrestris has a larger pileus (20–80 mm broad) without an orange tone, narrow and crowded lamellae with uneven edges, and a longer stipe (30–80 mm long) (
1 | Pileus dry | 2 |
– | Pileus viscid | 4 |
2 | Spores cylindric, germ pore very minute to not evident | Pholiota cylindrospora sp.nov. |
– | Spores elliptic, germ pore distinct | 3 |
3 | Spores 6–7.5 × 3.8–4.5 µm, germ pore apex not truncate | P. squarrosa |
– | Spores 5–6 × 3–3.5 µm, germ pore apex truncate | P. kodiakensis |
4 | Pileus ground color white to whitish | P. squarrosoides |
– | Pileus ground color not white to whitish | 5 |
5 | Spores 5–8 µm wide | 6 |
– | Spores 2.5–5 µm wide | 7 |
6 | Spores 6–8 µm wide, germ pore apex truncate | P. aurivelloides |
– | Spores 5–6 µm wide, germ pore apex not truncate | P. aurivella |
7 | Pileus with fibrillose scales | 8 |
– | Pileus with spot-like or triangular scales | 10 |
8 | Pileus and stipe brilliant yellow, spores smaller (4–5 × 2.5–3 µm) | P. flammans |
– | Pileus darker, spores bigger (> 4.5 µm long, > 3.5 µm wide) | 9 |
9 | Pileus 10–35 mm broad with orange tone, lamellae with even edges, stipe 18–30 mm long | P. subterrestris sp.nov. |
– | Pileus 20–80 mm broad without orange tone, lamellae with uneven edges, stipe 30–80 mm long | P. terrestris |
10 | Spores 5–6 µm long | P. adiposa |
– | Spores 6–8 µm long | 11 |
11 | Lamellae whitish when young and becoming ferruginous | P. limonella |
– | Lamellae yellow or pallid brownish when young | 12 |
12 | Lamellae pallid brownish when young and becoming cinnamon, germ pore distinct | P. abietis |
– | Lamellae yellow when young, germ pore minute or inconspicuous | 13 |
13 | Stipe with appressed fibrillose scales, annulus persistent | P. filamentosa |
– | Stipe with recured floccose scales, annulus evanescent | P. squarrosoadiposa |
We thank Shi-liang Liu, Guang-cheng Cao, and Xiao-ming Zhang for their contributions to the collection and photography of the specimens.
The authors have declared that no competing interests exist.
No ethical statement was reported.
This research was funded by the National Natural Science Foundation of China (No. 31300017) and National Key Research and Development Program of China (2019YFC1604703).
En-jing Tian: conceptualization, resources, supervision, project administration, methodology, and funding acquisition. En-jing Tian, Jia-hui Huang, Chun-yu Lei, and Ya-lun Shen: investigation. Jia-hui Huang and Chun-yu Lei: software. Jia-hui Huang, Chun-yu Lei, and Ya-lun Shen: validation. Jia-hui Huang and Ya-lun Shen: data curation. En-jing Tian, Jia-hui Huang and Chun-yu Lei: visualization. En-jing Tian and Jia-hui Huang: formal analysis, writing-original draft preparation, and review and editing.
Jia-hui Huang https://orcid.org/0009-0001-0323-3446
Chun-yu Lei https://orcid.org/0009-0001-4275-6813
Ya-lun Shen https://orcid.org/0009-0000-7394-4457
En-jing Tian https://orcid.org/0000-0001-9742-7569
The research data used in this study can be found in GenBank (https://www.ncbi.nlm.nih.gov) and MycoBank (https://www.mycobank.org).