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Research Article
New species, new records and common species of Pluteus sect. Celluloderma from northern China
expand article infoZheng-Xiang Qi, Ke-Qing Qian, Lei Yue, Li-Bo Wang, Di-Zhe Guo, Dong-Mei Wu§, Neng Gao§, Bo Zhang, Yu Li
‡ Jilin Agricultural University, Changchun, China
§ Biotechnology Research Institute, Xinjiang Academy of Agricultural and Reclamation Sciences, Shihezi, China
Open Access

Abstract

Wood-rotting fungi are organisms that can decompose wood substrates and extract nutrients from them to support their growth. They play a crucial role in the material cycle of forest ecosystems. The genus Pluteus plays a significant role in wood decomposition. In this study, the morphology and molecular systematics of the sect. Celluloderma of the genus Pluteus were carried out. Pluteus brunneodiscus was identified as a new species, along with the discovery of two new records, P. cystidiosus and P. chrysophlebius, and a common species, P. romellii. Pluteus brunneodiscus is characterized by the brown center of the pileus that transitions to white towards the margins, with the surface cracking to form irregular granules. It is typically found in Populus forests growing on decomposing twigs or wood chips. Line drawings, color photographs, and phylogenetic analyses of related species within the genus Pluteus accompany the descriptions of these four species. The analyses are based on ITS + TEF1-α sequence data. Finally, a key for the twenty species within the sect. Celluloderma of the genus Pluteus, which has been documented in China, is provided.

Key words

Line drawings, morphology, phylogeny, wood-rotting fungi

Introduction

The genus Pluteus Fr., which belongs to the Basidiomycota, Agaricomycetes, Agaricales, Pluteaceae, was established by Fries in 1863. The genus Pluteus is characterized by its free lamellae, pinkish spore print, inverse hymenophoral trama, smooth spherical to ellipsoidal basidiospores, various forms of pleurocystidia, and often cheilocystidia. It is predominantly found on decaying wood and has a global distribution (Vellinga and Schreurs 1985; Singer 1986; Justo et al. 2011a, 2011b).

The genus Pluteus was categorized into three sections based on the characteristics of the cystidia and pileipellis viz. (1) sect. Pluteus Fr is characterized by the existence of a cutis pileipellis and thick-walled pleurocystidia, (2) sect. Hispidoderma Fayod is characterized by a pileipellis that is a trichoderm composed of elongated cells and thin-walled pleurocystidia and (3) sect. Celluloderma Fayod is characterized by a pileipellis that is a hymeniderm or hymeniderm with cystidioid elements, comprising of clavate to spheropedunculate cells and thin-walled pleurocystidia (Lange 1917; Imai 1938; Singer 1956). Molecular phylogenetic analysis provides support for three sections (Pluteus Fr, Hispidoderma Fayod, and Celluloderma Fayod) (Menolli et al. 2010; Justo et al. 2011a, 2011b).

Singer further subdivided Pluteu s sect. Celluloderma into two subsections based on the composition of the pileipellis: subsect. Mixtini Singer, is characterized by elongated elements, and subsect. Eucellulodermini Singer is characterized by the absence of such elements (Singer 1956; Singer 1958). The molecular phylogenetic studies do not divide the Pluteus sect. Celluloderma into two subsections (Justo et al. 2011b). Some species belonged to the sect. Celluloderma (e.g., P. ephebeus (Fr.) Gillet and related species). Based on their characteristics, species composed of non-metuloid cystidia and a pileipellis as cutis should belong to the sect. Hispidoderma. This is not consistent with the phylogenetic results. Thus, the classification of the two subsections of sect. Celluloderma needs to be further justified.

Vellinga and Schreurs (1985) proposed a different taxonomic system to distinguish these species (e.g., P. ephebeus (Fr.) Gillet and related species), dividing the Pluteus sect. Celluloderma into three subsects, Mixtini, Eucelullodermini, and Hispidodermini (Fayod) Vellinga and Schreurs. The latter is characterized by a trichodermal pileipellis or a euhymeniderm consisting of cylindrical to fusiform elements, which are similar to some of the characteristics of the sect. Hispidoderma. Additionally, Schreurs and Vellinga proposed a new group sect. Villosi Schreurs and Vellinga, containing species with a cutis-like pileipellis and non-metuloid (Singer 1958; Singer 1986). The proposed new sections and subsections by Singer (1958, 1986), Vellinga, and Schreurs (1985) lack support from molecular systematic studies (Justo et al. 2011a; Justo et al. 2012).

Recent studies (Minnis et al. 2006; Menolli et al. 2010; Justo et al. 2011a, 2011b; Vizzini and Ercole 2011) have indicated that sect. Celluloderma includes species characterized by the presence of non-metuloid pleurocystidia and a pileipellis that is either euhymeniderm or epithelioid hymeniderm, composed of short elements, which may or may not be intermixed or not with elongate cystidioid elements (corresponding to Pluteus sect. Celluloderma as defined by Singer 1956, 1958, 1986), refers to species with a cutis-like pileipellis and non-metuloid cystidia (corresponding to Pluteus sect. Villosi or Hispidoderma sensu Singer p.p.).

In the current investigation, a new species, (P. brunneodiscus), two new records to China, (P. chrysophlebius and P. cystidiosus), and a common species, (P. romellii) are described. Detailed descriptions and illustrations are provided for the four species, along with clarification of the phylogenetic relationships of the identified species and related taxa from the genus Pluteus sect. Celluloderma.

Materials and methods

Morphology

In the field, photographs of fresh basidiomata were taken to scientifically and adequately reflect the growing environment and characteristics of the basidiomata, including the shape of the pileus, the color of the lamellae, and Munsell Soil Color Chart was followed for color codes (Munsell 2009). For fresh basidiomata, we promptly determined the size and recorded in detail the shape, size, color, odor, and other macroscopic characteristics of the basidiomata pileus, lamellae, and stipes. About 15 g of fresh context and lamellae were dried in a Ziplock bag with silica gel and returned to the lab for DNA extraction. Fresh basidiomata were dried at 40 ~ 45 °C (Hu et al. 2022), using a plant drying oven and preserved in the fungarium of Jilin Agricultural University (FJAU).

The observation of microstructural features was based on dried specimens. The dry specimens were rehydrated in 94% ethanol for microscopic examination and then mounted in 3% potassium hydroxide (KOH), 1% Congo Red, and Melzer’s Reagent, using a light microscope (ZEISS, DM1000, Oberkochen, Germany). Specifically, the following symbols were used in the description: [n/m/p] indicates that ‘n’ randomly selected basidiospores from ‘m’ basidiomata of ‘p’ collections were measured, ‘avl’ means the average length of basidiospores, except for the extreme values, ‘avw’ refers to the average width of the basidiospores, except the extreme values, ‘Q’ represents the quotient of the length and width of a single basidiospore inside view, ‘Qm’ refers to the average Q value of all basidiospores ± standard deviation. The dimensions for basidiospores are given as (a)b–c(d). The range of b–c contains a minimum of 90% of the measured values. Extreme values (i.e., a and b) are given in parentheses (Qi et al. 2022).

Molecular phylogeny

DNA extraction, PCR amplification, and sequencing

According to the instructions, the total DNA of the specimens was extracted by the new plant genomic DNA extraction kit from Jiangsu Kangwei Century Biotechnology Limited Company, P.R. China. Subsequently, sequences of the internal transcribed spacer (ITS) region, and translation elongation factor 1-α (TEF1-α) were used for phylogenetic analyses. The amplification primers of the nr ITS: ITS1–5.8S–ITS2 regions were ITS1F and ITS4/ITS4B (White et al. 1990), and TEF1-α regions were EF1–983F and EF1–1567R (Rehner and Buckley 2005). The amplification reactions were carried out in a 25 µL system. The total amount of PCR mixed was as follows: dd H2O 13.5 µL, 10 × Taq Buffer 5 µL, 10 mM dNTPs 1 µL, 10 mM upstream primer 1 µL, 10 mM downstream primer 1 µL, DNA sample 2 µL, 2 U/mm Taq Polymerase 1.5 µL. The cycle parameters were as follows: 5 min at 98 °C; 30 s at 98 °C, 30 s at 55 °C, 1 min at 72 °C for 40 cycles; 7 min at 72 °C; storage at 4 °C (Ševcíková et al. 2022). The PCR product was subjected to 1% agarose gel electrophoresis. The purified PCR products were sent to Sangon Biotech Limited Company, P.R. China for sequencing using the Sanger method. The sequencing results were clipped with Seqman 7.1.0 (Swindell and Plasterer 1997) and subsequently deposited in GenBank (https://www.ncbi.nlm.nih.gov/genbank).

Data analysis

The species that were morphologically similar to new species, newly recorded species, and common species, and have high sequence similarity after blast were selected (Justo et al. 2011b, 2012; Menolli et al. 2015; Desjardin and Perry 2018; Hosen et al. 2019; Hosen et al. 2021; Ševčíková et al. 2022; Qi et al. 2022; Malysheva et al. 2023; Ševcíková et al. 2023; Xu et al. 2023), and details of the ITS and TEF1-α sequences of these species are shown in Table 1. The ITS and TEF1-α dataset comprised 134 representative sequences that exhibited the highest similarity to Pluteus spp., and two sequences of Volvopluteus michiganensis (A.H. Sm.) Justo and Minnis. as an outgroup.

Table 1.

Names, collection numbers, reported countries and corresponding GenBank accession numbers of the taxa used in this study.

Taxon Collection Country GenBank No. Reference
ITS TEF1-α
Pluteus absconditus iNaturalist 112240775 USA (TN) OR229047 OR242143 Ševcíková et al. 2023
P. absconditus MO 136488 USA (TN) KM983689 OR242144 Ševcíková et al. 2023
P. aff. ephebeus BPI 882530 USA-Illinois JQ065025 Menolli et al. 2015
P. aff. ephebeus BPI 882531 USA-Illinois JQ065026 Menolli et al. 2015
P. aff. ephebeus HHB1213 USA-New Mexico KM983670 Menolli et al. 2015
P. aff. ephebeus AJ478 USA-Vigin Islands KM983675 Menolli et al. 2015
P. aff. ephebeus AJ535 Dominican Republic KM983676 Menolli et al. 2015
P. aletaiensis HMJAU 60207 China OM991943 OP573273 Qi et al. 2022
P. aletaiensis HMJAU 60208 China OM992247 OP573274 Qi et al. 2022
P. aurantiorugosus GDGM41547 China MK791275 Ševcíková et al. 2022
P. aurantiorugosus LE 312815 Russia (Europe) ON864103 ON813296 Ševcíková et al. 2022
P. austrofulvus AJ 857 USA, Arkansas KM983701 ON813290 Ševcíková et al. 2022
P. austrofulvus AJ 860 USA, Arkansas KM983699 ON813288 Ševcíková et al. 2022
P. brunneidiscus HMJAU 60206 China OM991893 Qi et al. 2022
P. brunneidiscus HMJAU 60210 China OM943513 Qi et al. 2022
P. cervinus REG 13641 USA HM562152 Qi et al. 2022
P. cf. nanus LE 213093 Russia FJ774081 Justo et al. 2011
P. cf. ephebeus LOU15198 Spain KM983671 Menolli et al. 2015
P. cf. ephebeus Shaffer4673 France HM562080 Menolli et al. 2015
P. cf. ephebeus Pearson sn England HM562198 Menolli et al. 2015
P. cf. ephebeus 9823 Italy JF908620 Menolli et al. 2015
P. cf. ephebeus 10151 Italy JF908621 Menolli et al. 2015
P. cf. fastigiatus NKI12 Brazil KM983678 Menolli et al. 2015
P. cf. fuliginosus FK2158 Brazil KM983677 Menolli et al. 2015
P. chrysophlebius TNSF12383 Japan HM562125 Justo et al. 2011a
P. chrysophlebius SF10 (BPI) USA (IL) HM562180 Justo et al. 2011a
P. chrysophlebius TNSF12388 Japan HM562088 Justo et al. 2011a
P. chrysophlebius SF12 (BPI) USA (IL) HM562182 Justo et al. 2011a
P. chrysophlebius SF11 (SIU) USA (IL) HM562181 Justo et al. 2011a
P. chrysophlebius FJAU66561 China OR994065 PP062824 This study
P. cutefractus BRNM825872 Spain OR229050 OR242162 Ševcíková et al. 2023
P. cutefractus GM 3458 Spain OR229048 OR242165 Ševcíková et al. 2023
P. cutefractus FG 26092015 Slovenia OR229053 OR242164 Ševcíková et al. 2023
P. cystidiosus LE 312852 Russia (Far East) OR229063 OR242175 Ševcíková et al. 2023
P. cystidiosus LE 313335 Russia (Far East) OR229062 OR242174 Ševcíková et al. 2023
P. cystidiosus AJ 782 (NBM-F-009790) USA (MA) KM983687 OR242171 Ševcíková et al. 2023
P. cystidiosus AJ 617 (NBM-F-009788) USA (NY) KM983686 OR242173 Ševcíková et al. 2023
P. cystidiosus FJAU66556 China OR994068 PP062825 This study
P. cystidiosus FJAU66557 China PP002166 PP062826 This study
P. diptychocystis NMJ184 Brazil KM983674 Menolli et al. 2015
P. ephebeus AJ234 Spain HM562044 Menolli et al. 2015
P. fenzlii TNSF12376 Japan HM562091 Menolli et al. 2015
P. fenzlii F1020647 Slovakia HM562111 Menolli et al. 2015
P. fenzlii LE 246083 Russia FJ774082 Holec et al. 2017
P. fulvibadius AJ 815 USA, California KM983698 ON813285 Ševcíková et al. 2022
P. fulvibadius HRL3391 Canada, Québec ON864094 ON813287 Ševcíková et al. 2022
P. gausapatus BRNM817745 South Korea OR229067 OR242177 Ševcíková et al. 2023
P. gausapatus BRNM817745 South Korea OR229067 OR242177 Ševcíková et al. 2023
P. halonatus FK2084 Brazil KM983680 Menolli et al. 2015
P. halonatus NKI17 Brazil KM983679 Menolli et al. 2015
P. heteromarginatus AJ172 USA HM562058 Hosen et al.2019
P. hirtellus SFSU:DED 8259 West Africa MG968804 Desjardin and Perry 2018
P. inconspicuus PDD 72485 New Zealand MN738614 Ševcíková et al. 2023
P. inflatus BRNM817761 Czech Republic OR229033 OR242136 Ševcíková et al. 2023
P. inflatus BRNM825836 Czech Republic OR229035 OR242132 Ševcíková et al. 2023
P. inflatus BRNM825837 Czech Republic OR229036 OR242133 Ševcíková et al. 2023
P. insidiosus 15120 Italy JF908626 Justo et al. 2012
P. longistriatus Minnis309203 USA HM562082 Hosen et al.2019
P. lucidus LE F-347426 Russia OQ732746 Malysheva et al. 2023
P. mammillatus Singer244A USA-Florida HM562120 Holec et al. 2017
P. mammillatus Minnis309202 USA-Missouri HM562086 Holec et al. 2017
P. mammillatus ASM7916 USA-Missouri HM562119 Holec et al. 2017
P. brunneodiscus FJAU66132 China PP002168 PP062821 This study
P. brunneodiscus FJAU66133 China PP002169 PP062822 This study
P. brunneodiscus FJAU66134 China PP002167 PP062823 This study
P. parvisporus AJ 855 USA, Arkansas ON864099 ON813295 Ševcíková et al. 2022
P. parvisporus iNaturalist 112236342 USA, Tennessee ON864098 ON813294 Ševcíková et al. 2022
P. phlebophorus AJ 81(NBM-F-009110) Spain HM562039 ON133554 Ševcíková et al. 2023
P. phlebophorus AJ228 (LOU) Spain HM562138 Justo et al. 2011a
P. phlebophorus AJ194 (LOU) Spain HM562137 Justo et al. 2011a
P. phlebophorus AJ193 (LOU) Spain HM562144 Justo et al. 2011a
P. plautus P59 USA-California KF306016 Menolli et al. 2015
P. podospileus LE 303682 Russia (South Siberia) KX216331 OR242169 Ševcíková et al. 2023
P. podospileus LE 303687 Russia (South Siberia) KX216332 OR242168 Ševcíková et al. 2023
P. podospileus LE 313589 Russia (South Siberia) OR229060 OR242167 Ševcíková et al. 2023
P. riberaltensis var. conquistensis FK1043 Brazil HM562162 Menolli et al. 2015
P. romellii AJ 232 Spain HM562062 ON813280 Ševcíková et al. 2022
P. romellii BRNM 761731 Czech Republic ON864065 ON813278 Ševcíková et al. 2022
P. romellii BRNM 816205 Czech Republic ON864063 ON813276 Ševcíková et al. 2022
P. romellii BRNM 825845 Slovakia ON864070 ON813281 Ševcíková et al. 2022
P. romellii FJAU66558 China OR994057 PP062827 This study
P. romellii FJAU66559 China OR994061 PP062828 This study
P. rugosidiscus BRNM761706 Slovakia MH010876 LT991752 Ševcíková et al. 2023
P. rugosidiscus Homola109 (MICH) USA (MI) HM562079 Justo et al. 2011a
Pluteus sp. SP394389 USA HM562161 Justo et al. 2012
Pluteus sp. iNaturalist 27406926 (NBM-F-009806) USA (IN) ON006984 OR242176 Ševcíková et al. 2023
P. squarrosus GDGM 42320 China MK791274 Hosen et al.2019
P. squarrosus GDGM 42302 China MK791273 Hosen et al.2019
P. thomsonii LE 303662 Russia KX216329 Justo et al. 2012
P. tomentosulus MO163564 USA-Pennsylvania KM983673 Menolli et al. 2015
P. tomentosulus MO93719 USA-Oregon KM983672 Menolli et al. 2015
V. michiganensis HMJAU-CR45 China MW242665 Qi et al. 2022
Volvopluteus michiganensis HMJAU-CR43 China MW242664 Qi et al. 2022

For obtaining ITS + TEF1-α datasets of related species, sequence alignment was initially performed for ITS and TEF1-α using the “automatic” strategy and normal alignment mode of MACSE V2.03 (Ranwez et al. 2018) and MAFFT (Katoh and Standley 2013), respectively. Subsequently, the alignments were manually adjusted in BioEdit v7.1.3 (Hall 1999). Afterward, ITS and TEF1-α sequences were aligned and combined using Phylosuit V1.2.2 (Zhang et al. 2020). Then, ModelFinder (Kalyaanamoorthy et al. 2017) was used to select the best-fit models using the Bayesian information criterion (BIC). In this case, the Maximum likelihood (ML) analyses were performed in IQTree 1.6.8 (Nguyen et al. 2015), and the Bayesian inference phylogenies were performed in MrBayes 3.2.6 (Ronquist et al. 2012) (two parallel runs, 2,000,000 generations), in which the initial 25% of sampled data were discarded as burn-in. The above software was integrated into PhyloSuite 1.2.2 (Zhang et al. 2020). The ML phylogenetic tree was evaluated using the bootstrap method with a bootstrap value of 1,000 replicates; BI determined that the analysis reached smoothness with a variance of less than 0.01 and terminated the calculation. Finally, the evolutionary tree was followed up with Figtree v1.4.

Results

Phylogenetic analyses

This study’s nrITS dataset comprises 93 sequences and 650 characters (gaps included). The TEF1-α dataset comprises 41 sequences and 530 characters (gaps included). The combined nrITS + TEF1-α dataset consists of 134 sequences and 1180 characters, including gaps. Of these, 16 sequences (8 nrITS and 8 TEF1-α) were newly generated in this study (Table 1). The overall topologies of the ML and BI trees were nearly identical for all datasets.

For clarity and brevity, we use the term “strongly supported” for a clade/relation that receives a bootstrap (BS) 90 and a posterior probability (PP) = 1, and “well supported” if it receives a BS 70 and a PP of 0.95. The individual support values are shown in Fig. 1.

Figure 1. 

Phylogenetic tree of the sect. Celluloderma of the genus Pluteus. The best tree from the ML and BI analysis of the nrITS + TEF1-α dataset. The two values of internal nodes respectively represent the maximum likelihood bootstrap (MLBP)/Bayesian posterior probability (BIPP). This study species is in bold and red font.

Within the sect. Celluloderma, six strongly supported clades are recovered in the combined nrITS + TEF1-α dataset:

  1. Clade I: This includes the clade we consider to represent P. mammillatus (Longyear) Minnis, Sundb. & Methven from the USA, P. fenzlii (Schulzer) Corriol & P.-A. Moreau from Japan, Slovakia, and Russia, P. halonatus from Brazil.
  2. Clade II: Includes only the newly described P. brunneodiscus from China. This also includes the clade we consider to represent P. squarrosus Hosen & T.H. Li from China, P. hirtellus Desjardin & B.A. Perry from West Africa, P. plautus (Weinm.) Gillet from the USA, P. tomentosulus Peck from the USA, P. diptychocystis Singer from Brazil, and P. riberaltensis var. conquistensis from Brazil, while P. ephebeus from Spain, France, England, and Italy (P. cf. ephebeus and P. aff. ephebeus), P. fuliginosus Murrill from Brazil (P. cf. fuliginosus), P. fastigiatus Singer from Brazil (P. cf. fastigiatus).
  3. Clade III: Includes the newly described P. cystidiosus (China). This clade also includes the clade we consider to represent P. podospileus Sacc. & Cub. (Russia), P. cutefractus Ferisin, Dovana & Justo (Spain, Slovenia), P. inflatus Velen (Czech Republic), P. inconspicuus E. Horak (New Zealand); three recently described species, P. cystidiosus (Russia, USA), P. absconditus Justo, Kalichman & S.D. Russell (USA), and P. gausapatus Ševčíková & Antonín (South Korea), and one likely undescribed species from the USA (iNaturalist 27406926).
  4. Clade IV: Includes the newly described P. romellii (China). It also includes P. fulvibadius Murrill (USA and Canada), P. aurantiorugosus (Trog) Sacc (China and Russia). Three recently described species, P. austrofulvus Justo, Minnis, S.D. Russell & Kalichman (USA), P. parvisporus Justo, Kalichman & S.D. Russell (USA) and P. aletaiensis Z.X. Qi, B. Zhang and Yu Li (China).
  5. Clade V: Includes the newly described P. chrysophlebius (China). This clade also includes the clade we consider to represent P. chrysophlebius (Japan, USA, Japan), P. phlebophorus (Ditmar) P. Kumm (Spain), and P. rugosidiscus Murrill (Slovakia, USA).
  6. Clade VI: This clade includes the clade that we consider to represent P. insidiosus Vellinga & Schreurs (Italy) and P. thomsonii (Berk. & Broome) Dennis (Russia).

Taxonomy

Pluteus brunneodiscus Z.X. QI, B. Zhang & Y. Li, sp. nov.

MycoBank No: 851479
Figs 2A–B, 3

Typification

China. Xinjiang Uygur Autonomous Region, Ili Kazakh Autonomous Prefecture, Tekes County, Aktamu Wetland, 43°15'22.61"N, 81°75'90.21"E, alt. 1243 m, 6 July 2022, Z.X. Qi (FJAU 66134, holotype!).

Sequences holotype

ITS: PP002167, TEF1-α: PP062823.

Etymology

“brunneo-”: brown, “-discus”: pileus disc. The species epithet “brunneodiscus” (Lat.) refers to the brown of the middle part of the pileus disc.

Diagnosis

Pluteus brunneodiscus differs from P. tomentosulus by its brown pileus in the middle, transitioning to white toward the margins, and the surface cracks to form irregular granules. It grows in poplar forests (Populus alba var. pyramidalis Bge) with decaying wood branches or chips.

Description

Basidiomata medium to large. Pileus 39–71 mm in diam, initially compressed hemispherical, surface with dense brown irregular granules (5.0YR 5/2), dirty white (5.0YR 9/2), middle brown (5.0YR 4/4), margin entire, gradually spreading at maturity, pileus middle dark brown (5.0YR 3/6), margin irregularly dehiscent at maturity or after hygrophanous. Context whitish (5.0YR 9/2), odorless, 3–6 mm thick. Lamellae initially dirty white (5.0YR 9/2), becoming flesh-brown to earth-brown at maturity (5.0YR 8/4- 5.0YR 6/4), free, dense, thick, unequal, slightly ventricose, 6–7 mm wide. Stipe 37–55 mm long, 8–11 mm wide, dirty white (5.0YR 9/2), cylindrical, slightly thicker at the base, fibrous, with white longitudinal stripes on the surface. Odorless. Spore prints pink.

Basidiospores [120, 12, 3] (–6.5) 7.0–7.5 (–8.0) × 5.0–6.0 (–6.5) µm, avL × avW = 7.0 × 6.0 µm, Q = 1.16–1.30–1.45 µm, avQ = 1.16 µm, globose, subglobose, slightly pink, smooth, thin-walled, non-dextrinoid, partially containing one droplet or irregular inclusions. Basidia 25–32 × 7–11 μm, fusiform to clavate, thin-walled, 4–sterigmate, and hyaline in KOH. Pleurocystidia abundant, scattered, 55–102 × 22–36 μm, vesicular to narrowly vesicular, or clavate, thin-walled, smooth, and hyaline in KOH. Cheilocystidia abundant, clustered, 41–79 × 18–29 μm, subfusiform to fusiform, or ventrally bulbous, apically broadly digitate 15–23 μm long, thin-walled, hyaline. Lamellar trama divergent. Pileipellis a cutis to trichodermium, hyphae 4–10 µm diam, cylindrical, hyaline, non-gelatinous; terminal cells inflated, 62–91 × 22–31 μm, obtusely rounded or pointed apically, thin-walled, with brown cytoplasmic pigments. Stipitipellis a cutis, hyphae 5–9 µm diam, cylindrical, hyaline, non-incrusted, non-gelatinous, thin-walled. Caulocystidia absent. Clamp connections absent in all tissues.

Figure 2. 

Basidiomata features A–B Pluteus brunneodiscus C–D P. cystidiosus E–F P. chrysophlebius G–I P. romellii. Photos by Zheng-xiang Qi (A–B, G–I). Photos by Di-zhe Guo (C–F). Scale bars: 1 cm.

Ecology and distribution

Solitary to scattered on the ground in the broad-leaved forests (Populus alba var. pyramidalis Bge) with decaying wood branches or wood chips. Known from Xinjiang Uygur Autonomous Region of China.

Additional specimens examined

China. Xinjiang Uygur Autonomous Region, Ili Kazakh Autonomous Prefecture, Tekes County, Aktamu Wetland, 43°15'22.61"N, 81°75'90.21"E, alt. 1243 m, 6 July 2022, Z.X. Qi, D.M. Wu, N. Gao and B.K. Cui, FJAU 66132 (ITS: PP002168, TEF1-α: PP062821). China. Xinjiang Uygur Autonomous Region, Ili Kazakh Autonomous Prefecture, Tekes County, Aktamu Wetland, 43°15'22.61"N, 81°75'90.21"E, alt. 1243 m, 6 July 2022, Z.X. Qi, FJAU 66133 (ITS: PP002169, TEF1-α: PP062822).

Notes

Morphologically, Pluteus brunneodiscus is very similar to P. tomentosulus in having a white pileus. The difference lies in the surface texture, as P. tomentosulus has a very finely granular-tomentose surface that becomes bald at maturity, while P. brunneodiscus features a brown center of the pileus, transitioning to white toward the margins, with the surface cracking to form irregular granules (Vellinga and Schreurs 1985; Orton 1986; Vellinga 1990; Desjardin and Perry 2018).

In phylogenetic analyses, P. brunneodiscus clusters in the ephebeus clade as a sister species to P. aff. ephebeus, and has a support ratio of 1/100. However, the pileus of P. aff. ephebeus are sooty, shield-shaped fruiting bodies with pubescent or downy surfaces. They grow on rotting wood or stumps and are widely distributed in Britain and Ireland (Orton 1986; Justo et al. 2011a; Menolli et al. 2015). These characteristics distinguish P. brunneodiscus from P. aff. ephebeus.

Figure 3. 

A Macroscopic characteristics of Pluteus brunneodiscus B basidiospores C pleurocystidia D basidia E pileipellis terminal cells F cheilocystidia. Scale bars: 1 cm (A); 10 µm (B–F).

Pluteus cystidiosus (Minnis and Sundb.) Justo, Malysheva & Lebeuf, in Ševčíková et al., Journal of Fungi 9(9, no. 898): 34 (2023)

Figs 2C–D, 4

Pluteus seticeps var. cystidiosus Minnis and Sundberg N. Amer. Fung. 5(1): 13 (2010). Syn.

Description

Basidiomata medium to large. Pileus 25–41 mm in diam, compressed hemispherical, surface spreading when young, surface with longitudinal vein-like folds from middle to margin when mature, margin mostly transverse folds, light brown to dark brown (5.0YR 5/6-5.0YR 4/12), margin entire. Context dirty white (2.5YR 9/4), odorless, 5–8 mm thick. Lamellae dirty white (2.5YR 9/4), free, dense, thick, unequal, ventricose, 15–18 mm wide. Stipe 30–41 mm long, 12–17 mm wide, cylindrical, slightly thicker at the base, hollow, fibrous, with brown serpentine or crumbly scales on the surface (2.5YR 9/2). Odorless. Spore prints pink.

Basidiospores [200, 10, 2] (–5.0) 5.5–6.0 (–6.5) × (–4.5) 5.0–5.5 μm, avL × avW = 6.0 × 5.0 µm, Q = 1.10–1.20–1.30 μm, avQ = 1.20 μm, spherical, subglobose, slightly pink, smooth, thin-walled, non-dextrinoid, partially containing one droplet or irregular inclusions. Basidia 23–31 × 7–10 μm, clavate, thin-walled, 4-sterigmate, and hyaline in KOH. Pleurocystidia abundant, scattered, 55–102 × 22–36 μm, rod-shaped or subpyriform, vesicular, thin-walled, smooth, and hyaline in KOH. Cheilocystidia abundant, clustered, 37–60 × 15–22 μm, clavate, fusiform or vesicular, thin-walled. Lamellar trama divergent. Pileipellis a hymeniderm or epithelioid hymeniderm, made up of two types of elements; spheropedunculate or pyriform, 27–55 × 24–34 μm; broadly fusiform, inflated-fusiform, lanceolate, narrowly utriform, often mucronate, 56–105 × 11–23 μm; all elements with brown intracellular pigment, often aggregated in spots, slightly thick-walled. Stipitipellis a cutis of cylindrical, hyphae 8–11 μm wide, with pale brown pigment. Caulocystidia common, often in clusters, 36–112 × 9–20 μm, cylindrical, narrowly clavate, narrowly fusiform, spheropedunculate, with brown or yellow-brown pigment. Clamp connections absent in all studied tissues.

Ecology

Scattered on decaying wood in mixed coniferous forests (Pinus koraiensis Siebold and Zucc).

Distribution

Canada, the USA, Japan, Russian Far East.

Additional specimens examined

China. Heilongjiang Province, Liangshui National Nature Reserve. 47°11'22.24"N, 128°47'89.11"E, 23 June 2019, D.Z. Guo, FJAU 66556 (ITS: OR994068, TEF1-α: PP062825). China. Heilongjiang Province, Liangshui National Nature Reserve. 47°11'22.24"N, 128°47'89.11"E, 28 June 2019, D.Z. Guo, FJAU 66557 (ITS: PP002166, TEF1-α: PP062826).

Note. Ševcíková et al. (2023) elevated Pluteus seticeps var. cystidiosus to P. cystidiosus based on specimens from the USA, Canada, Japan, and Russia. The present study reports P. cystidiosus as a new record in China. There was almost complete overlap in morphological variation between those reported in the present study and the holotype specimen. Both grow in temperate/cold-temperate forests. However, the basidiospores of the species in the present study were slightly larger, measuring (–5.0) 5.5–6.0 (–6.5) × (–4.5) 5.0–5.5 µm, while those of the holotype specimen were smaller, measuring 4.5–5.5 (–6.2) × 3.5–5.0 µm.

Figure 4. 

A Macroscopic characteristics of Pluteus cystidiosus B caulocystidia C basidiospores D pleurocystidia E cheilocystidia F basidia G pileipellis. Scale bars: 1 cm (A); 10 µm (B–G).

The phylogenetic tree also supports the results of our morphological study, showing that our specimens are clustered in the same branch as those from the USA and Russia, with a support ratio of 1/100.

Pluteus chrysophlebius (Berk. & M.A. Curtis) Sacc., Syll. fung. (Abellini) 5: 678 (1887)

Figs 2E–F, 5

Agaricus chrysophlebius Berk. and M.A. Curtis 1859. Syn.

Description

Basidiomata medium. Pileus 15–22 mm in diameter, surface not spreading, irregularly pitted, smooth, central part umbo, wrinkled or veined, yellow to bright yellow (5.0Y 9/12-5.0Y 9/20), with a hyaline stripe in the central part 3/4 of the way toward the margin, margin entire. Context yellowish (5.0Y 9/8), odor inconspicuous. Lamellae yellow to brownish yellow (5.0Y 9/6- 5.0Y 9/8), free, dense, thick, unequal, ventricose, 6–8 mm wide. Stipe 25–42 mm long, 4–6 mm wide, cylindrical, slightly thicker at the base, fibrous, bright yellow to yellow (5.0Y 9/10-5.0Y 9/18), smooth, with white tomentose dense cilia at the base. Odorless. Spore prints pink.

Basidiospores [90, 3, 1] 5.5–6.0 × (–4.5) 5.0–5.5 μm, avL × avW = 6.0 × 5.0 µm, Q = 1.09–1.20–1.33 μm, avQ = 1.20 μm, globose, subglobose, slightly pinkish, smooth, thinly walled, non-dextrinoid, partially containing one droplet or irregular inclusions. Basidia 23–34 × 7–11 μm, clavate, thin-walled, 4-sterigmate, and hyaline in KOH. Pleurocystidia scattered, 52–78 × 15–24 μm, broad and long-necked vase-like, partly with a long neck, neck with inclusions, thin-walled, smooth, and hyaline in KOH. Chilocystidia abundant, clustered, smaller, 45–66 × 14–21 μm, similar to pleurocystidia, long-necked vase-shaped to fusiform, thin-walled. Lamellar trama divergent. Pileipellis an euhymeniderm of spheropedunculate and subglobose elements 28–67 × 18–41 μm, with brown or light brown, at the center brown to dark brown. Stipitipellis a cutis, hyphae 5–9 μm wide, hyaline, non-gelatinous, thin-walled. Caulocystidia absent. Clamp connections absent in all tissues.

Figure 5. 

A macroscopic characteristics of Pluteus chrysophlebius B basidiospores C basidia D pleurocystidia E cheilocystidia F pileipellis. Scale bars: 1 cm (A); 10 µm (B–G).

Ecology

Solitary on decaying wood in mixed coniferous forests.

Distribution

North America, South America.

Additional specimens examined

China. Heilongjiang Province, Liangshui National Nature Reserve. 47°11'22.24"N, 128°47'89.11"E, 24 June 2019, D.Z. Guo, FJAU 66561 (ITS: OR994065, TEF1-α: PP062824).

Note

Pluteus chrysophlebius was first reported in China. It can be distinguished from other yellow-pileus species such as P. admirabilis (Peck) Peck, P. aurantiacus Murrill, P. melleus Murrill, and P. rugosidiscus Murrill by its yellowish pileus and stipe, as well as its bald pileus texture (Minnis and Sundberg 2010; Malysheva et al. 2016). The phylogenetic analysis also supports the differentiation of species.

In the phylogenetic tree, P. chrysophlebius formed a cluster with TNSF12383 and TNSF12388 in Asia and was sister to SF10-SF12 in the United States, with strong support for both clades.

Pluteus romellii (Britzelm.) Lapl., Dict. iconogr. champ. sup. (Paris): 533 (1894)

Figs 2G–I, 6

Agaricus romellii Britzelm., Hymenomyceten aus Südbayern VIII: 5 (1891). Syn.

Description

Basidiomata medium to large. Pileus 20–56 mm broad, compressed hemispherical to spreading, surface with vein-like projections extending to the pileus margin, often with striated dehiscence, with a greasy or almost waxy texture, brown to yellowish-brown (7.5YR 8/8-7.5YR 6/12), margins wavy dehiscence with translucent-striate. Context light yellow (7.5YR 8/12), odorless, 2–3 mm thick. Lamellae yellowish (10.0YR 8/10), free, medium dense, unequal, entire, ventricose, 5–7 mm wide. Stipe 26–41 mm long and 4–8 mm wide, cylindrical, slightly thicker at the base, fibrous, upper part of the stipe white to yellowish (10.0YR 9/8-10.0YR 7/12), smooth, lower part of the stipe with white tomentum, yellow to yellow-brown (10.0YR 8/8-10.0YR 8/12). Odorless. Spore print pale pink.

Basidiospores [120, 4, 2] 7.0–7.5 (–8.0) × 6.0–6.5 µm, avL × avW = 7.0 × 6.0 µm, Q = 1.07–1.25~1.33 µm, avQ = 1.16 µm, globose, subglobose to ellipsoid, transparent to slightly pinkish, smooth, and thin-walled, non-dextrinoid, partially containing one droplet or irregular inclusions. Basidia 27–32 × 8–10 μm, clavate, thin-walled, 4-sterigmate, and hyaline in KOH. Pleurocystidia abundant, scattered, 55–102 × 22–36 μm, rod-shaped or subcylindrical, fusiform, with neck and apical part broader and obtuse, thinly walled, smooth, and hyaline in KOH. Cheliocystidia abundant, clustered, 41–79 × 18–29 μm, pyriform or similarly pleurocystidia shape, thin-walled. Lamellar trama divergent. Pileipellis an euhymeniderm of spheropedunculate and subglobose elements 25–48 × 23–35 μm, with brown or light brown, at the center brown to dark brown. Stipitipellis a cutis, hyphae 6–10 μm wide, hyaline, non-gelatinous, thin-walled. Caulocystidia absent. Clamp connections absent in all tissues.

Figure 6. 

A macroscopic characteristics of Pluteus romellii B basidiospores C pleurocystidia D basidia E cheilocystidia F pileipellis. Scale bars: 1 cm (A); 10 µm (B–E); 20 µm (F).

Ecology

Solitary to scattered on decaying wood in coniferous forests (Picea schrenkiana Fisch.).

Distribution

Europe, Americas, East Asia, Africa.

Additional specimens examined

China. Xinjiang Uygur Autonomous Region, Ili Kazakh Autonomous Prefecture, Tekes County, Jongkushtai Village, 43°12'26.61"N, 81°91'97.21"E, alt. 2139 m, 10 July 2022, Z.X. Qi, J.J. Hu, and B. Zhang, FJAU 66558 (ITS: OR994057, TEF1-α: PP062827). China. Xinjiang Uygur Autonomous Region, Ili Kazakh Autonomous Prefecture, Tekes County, Jongkushtai Village, 43°15'22.61"N, 81°75'90.21"E, alt. 2147 m, 11 July 2022, Z.X. Qi, J.J. Hu, and B. Zhang, FJAU 66559 (ITS: OR994061, TEF1-α: PP062828).

Note

Initially, the description of Pluteus romellii was rather vague (Britzelmayr 1891), stating that P. romellii was similar to P. nanus (Pers.) P. Kumm, with spores measuring 6–7 μm, and found growing in the soil of Bavaria. It is now widely acknowledged that P. romellii is characterized by a brown pileus, yellow stipe, and the absence of elongated elements in the pileipellis. This species is placed on the phylogenetic tree in subsect. Eucellulodermini under sect. Celluloderma (Orton 1986; Vellinga 1990; Ševcíková et al. 2023). Here, our description of the P. romellii is consistent with the commonly accepted characterization. Phylogenetic analysis shows that it clustered with the epitype (BRNM 761731) with strongly supported (99/0.98).

Key to the reported species of Pluteus sect. Celluloderma in China

1 Pileipellis consists of spheropedunculate cells and elongated cystidioid elements 2
Pileipellis consists of spheropedunculate cells without elongated cystidioid elements 7
2 With caulocystidia 3
Without caulocystidia 6
3 With pleurocystidia 4
Without pleurocystidia Pluteus cinnabarinus
4 Cheilocystidia with short to long mucronate at the apex Pluteus aurantioruber
Cheilocystidia without short to long mucronate at the apex 5
5 Pleurocystidia larger, measuring 35–73 (–82) × 11–31 µm Pluteus cystidiosus
Pleurocystidia smaller, measuring 36–51 × 13.4–24 µm Pluteus podospileus
6 Pileus middle reticulate elevated, radially rugose Pluteus thomsonii
Pileus brown with stripes extending to the margins Pluteus striatus
7 Pileipellis consists of globular, obpyriform, or spheropedunculate cells 8
Pileipellis consists of without globular, obpyriform, or spheropedunculate cells 16
8 Grows on rotting wood 9
Grows on non-rotting wood Pluteus aletaiensis
9 Pileus, stipe bright-colored 10
Pileus, stipe not bright-colored 13
10 Pileus middle folded, groove-like striate Pluteus chrysophaeus
Pileus middle non-folded, groove-like striate 11
11 Pileus bright red or orange-red Pluteus aurantiorugosus
Pileus non-bright red to orange-red 12
12 Pileus smooth, widely distributed in North America Pluteus chrysophlebius
Pileus goose-yellow, margin striate Pluteus admirabilis
13 Basidiomata small Pluteus nanus
Basidiomata non-small 14
14 Lamellae edged with a powdery creamy material Pluteus pulverulentus
Lamellae edged without a powdery creamy material 15
15 Pleurocystidia with neck and broad, blunt apex Pluteus romellii
Pileus teal brown, dark cinnamon-colored, with black ribbed veins or wrinkles Pluteus phlebophorus
16 Grows on rotting wood 17
Grows on non-rotting wood Pluteus brunneodiscus
17 Pileus margin with hyaline stripes 18
Pileus margin without hyaline stripes 19
18 Cheilocystidia with mucronate at the apex Pluteus pallidus
Cheilocystidia without mucronate at the apex Pluteus brunneoalbus
19 Pileus with dark brown frosting powder, radially dehiscent to margins Pluteus diettrichii
Pileus surface squarrose, stipe with surface covered by caulocystidia elements Pluteus squarrosus

Discussion

Singer (1986) and Vellinga and Schreurs (1985) classified sect. Celluloderma into two subsections: subsect. Celluloderma and subsect. Mixtini. However, subsequent systematic analyses of sect. Celluloderma did not have a high level of support from internal topology analysis, leading to the conclusion subsect. Eucellulodermini and subsect. Mixtini should not conform to natural taxonomy. Singer (1986) proposed that species with non-metuloid cystidia, a cutis, and trichodermal pileipellis should be classified in the sect. Hispidoderma. Vellinga and Schreurs (1985) proposed sect. Villosi on the basis of a cutis-like pileipellis and non-metuloid cystidia. However, in the ephebeus clade, there are P. ephebeus from Europe and P. riberaltensis var. conquistensis from the USA. These species should be placed in sect. Hispidoderma and classified based on the pileipellis, but molecular results indicate that it belongs to sect. Celluloderma. In the phylogenetic tree, it is the sister group to P. fenzlii, P. mammillatus, and some species have a partial veil. P. brunneodiscus in the ephebeus clade in the present study, which has non-metuloid cystidia and pileipellis as a cutis, shares their views with Vellinga and Schreurs (1985). The phylogenetic tree also exhibits a high level of support. Further research is needed to restore these species to sect. Villosi.

The presence of a partial veil in P. aurantiorugosus, P. aurantiorugosus var. aurantiovelatus, P. fenzlii, and P. mammillatus suggests that the occurrence or nonoccurrence/ lack of the partial veil in the evolutionary history of Pluteus occurred independently. As stated by Singer states (Singer 1958; Minnis and Sundberg 2010; Justo et al. 2011a, 2011b; Vizzini and Ercole 2011), this characteristic is homoplasic and unsuitable for the natural classification of these fungi at the supraspecific rank.

Acknowledgments

We are very thankful to Mr. Yong-Lan Tuo, Ya-Jie Liu, and Yang Wang (Engineering Research Center of Edible and Medicinal Fungi, Ministry of Education, Jilin Agricultural University, China) for their kind help in molecular studies. We thank the associate editor and the reviewers for their useful feedback that improved this paper.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

The study was supported by the Research on the Creation of Excellent Edible Mushroom Resources and High Quality and efficient Ecological Cultivation Technology in Jiangxi Province (20212BBF61002); the Scientific and Technological Tackling Plan for the Key Fields of Xinjiang Production and Construction Corps (no. 2021AB004); the National Key R and D of Ministry of Science and Technology (2023YFD1201601); the Modern Agroindustry Technology Research System (CARS20) 6the “111” program (D17014).

Author contributions

Zheng-xiang Qi: conceptualization, writing - original draft and review and editing, data curation, formal analysis, investigation, methodology and visualization. Ke-qing Qian: Writing - review and editing. Lei Yue: formal analysis. Li-bo Wang: investigation. Di-zhe Guo: investigation. Dong-mei Wu: methodology and visualization. Neng Gao: methodology and visualization. Bo Zhang: project administration, resources and supervision. Yu Li: writing - review and editing, formal analysis.

Author ORCIDs

Zheng-Xiang Qi https://orcid.org/0000-0002-0037-9407

Ke-Qing Qian https://orcid.org/0000-0002-4627-5240

Bo Zhang https://orcid.org/0000-0001-9508-8188

Data availability

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

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