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Article

Morphological and Molecular Identification of Three New Macrofungal Species from Shenyang and Adjacent Areas, Northeast China

by
Zi-Qi You
1,2,
Lin-Jiang Zhou
1,2,
Hai-Sheng Yuan
1,* and
Hyang Burm Lee
3,*
1
CAS Key Laboratory of Forest Ecology and Silviculture, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110164, China
2
University of the Chinese Academy of Sciences, Beijing 100049, China
3
Environmental Microbiology Lab, Department of Agricultural Biological Chemistry, College of Agriculture and Life Sciences, Chonnam National University, Gwangju 61186, Republic of Korea
*
Authors to whom correspondence should be addressed.
J. Fungi 2026, 12(7), 491; https://doi.org/10.3390/jof12070491
Submission received: 22 May 2026 / Revised: 29 June 2026 / Accepted: 30 June 2026 / Published: 3 July 2026
(This article belongs to the Special Issue Taxonomy, Phylogeny, and Ecology of Fungi from Different Ecological Niches)
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Abstract

Macrofungal resources are abundant in Northeast China, but those from Shenyang and its surrounding areas remain insufficiently investigated. In this study, morphological and phylogenetic analyses were carried out on specimens collected from the vicinity of Shenyang, Liaoning Province. Phylogenetic trees were inferred using maximum likelihood and Bayesian inference methods based on sequence data from the nuclear ribosomal internal transcribed spacer (ITS) region and the nuclear ribosomal large subunit (LSU). Three new species, Descolea laevis (Bolbitiaceae), Leucocoprinus shenyangensis (Agaricaceae), and Tephrocybe umbonata (Lyophyllaceae), are described herein. Descolea laevis is characterized by pale yellow to light yellow basidiomata, a nearly smooth pileus surface, clavate to narrowly clavate cheilocystidia, fusiform to clavate pleurocystidia, and amygdaliform to limoniform, verrucose basidiospores. Leucocoprinus shenyangensis is distinguished by white to yellowish-white basidiomata, a pileus covered with light grey squamules, narrowly clavate to subcylindrical cheilocystidia, a hymenidermal pileipellis, and amygdaliform to limoniform basidiospores. Tephrocybe umbonata is characterized by orange-white to greyish-orange basidiomata, a pileus with a blunt umbo, a smooth or slightly finely fibrillose pileus surface, lageniform pleurocystidia with slightly acute apices, and amygdaliform to limoniform basidiospores. Detailed morphological descriptions, illustrations of microscopic structures, and phylogenetic evidence for the three new species are provided. The diagnostic characteristics separating the new taxa from their closely related species are also discussed.

1. Introduction

Descolea Singer is a small genus in Bolbitiaceae, Agaricales, and is typified by D. antarctica Singer [1]. Species of the genus are typically characterized by epigeous agaricoid basidiomata, a dry to viscid pileus surface, a central stipe, and ornamented basidiospores. Descolea is phylogenetically closely related to Setchelliogaster, Descomyces, and Timgrovea. However, Setchelliogaster forms secotioid basidiomata, whereas the latter two genera produce fully gasteroid basidiomata [2]. Descolea is generally regarded as an ectomycorrhizal genus associated with woody plants in forest ecosystems [2]. Species of the genus have been reported mainly from Australasia and southern South America, where they are often associated with forests dominated by Nothofagaceae and Myrtaceae [2]. However, Asian records suggest that the host range of the genus may be broader, and D. quercina has been reported in association with Quercus in moist temperate forests of Pakistan [3]. Approximately 29 species of Descolea are accepted worldwide according to Index Fungorum and MycoBank. To date, records of Descolea from China remain scarce, and only two species, D. flavoannulata (Lj. N. Vassiljeva) E. Horak and D. pretiosa E. Horak, have been confirmed [4].
Leucocoprinus Pat., typified by L. cepistipes (Sowerby) Pat., is a lepiotaceous genus in Agaricaceae [5]. Traditionally, Leucocoprinus was separated from Leucoagaricus by a combination of characters, including striate to plicate pileus margins, metachromatic basidiospores, pseudoparaphyses among the basidia, and the general absence of clamp connections [6,7]. Molecular phylogenetic studies have shown, however, that these characters are not fully congruent with natural lineages, and that Leucocoprinus, Leucoagaricus, Micropsalliota, and related genera form a complex assemblage within Agaricaceae [8]. Recent treatments have differed in their interpretation of this complex, ranging from a broad circumscription of Leucocoprinus that transfers many taxa formerly placed in Leucoagaricus to Leucocoprinus [9], to classifications that retain Leucocoprinus and Leucoagaricus as separate genera and recognize segregate genera such as Candelolepiota and Macropsalliota [10]. More recent treatments have further recognized additional segregate genera, including Pulchrolepiota and Tristolepiota [11]. These alternative classifications indicate that the generic limits of Leucocoprinus remain unsettled and that traditional morphological features should be used cautiously for generic delimitation. Ecologically, species assigned to Leucocoprinus and allied genera are saprotrophic and occur on diverse organic substrates, including forest soil, litter, humus, rotten wood, compost, flowerpots, greenhouses, and other decomposing plant materials [6,12]. The group is most diverse in tropical and subtropical regions [13], but recent studies from temperate areas of China suggest that its diversity in northern regions remains incompletely documented [14,15].
The circumscription of Tephrocybe has undergone substantial revision in recent years. Molecular phylogenetic studies have shown that Tephrocybe in the traditional broad sense is paraphyletic, and several species formerly assigned to this genus have been transferred to Lyophyllum, Myochromella, Sagaranella, Sphagnurus, and other allied genera [16]. At present, Tephrocybe Donk is treated as a genus in Lyophyllaceae, and it is typified by T. rancida (Fr.) Donk [17]. The presence of siderophilous granules in the basidia, a diagnostic character of Lyophyllaceae, is also consistently observed in Tephrocybe [18]. Molecular phylogenetic studies have shown that the core lineage of Tephrocybe is placed in the termitomycetoid clade, and it is closely related to Termitomyces and Blastosporella [19]. In contrast to these allied groups, Tephrocybe has clamp connections, and it is not associated with termites [20]. Subsequent studies have further shown that some taxa superficially resembling Tephrocybe should be segregated into newly established genera such as Phaeotephrocybe and Praearthromyces, indicating that a tephrocyboid habit alone is insufficient to define the genus. Ecologically, species of Tephrocybe are mainly free-living terrestrial fungi, and the type species T. rancida occurs on soil in coniferous forests [19]. Approximately 62 species of Tephrocybe are currently accepted worldwide, and the known diversity of the genus is concentrated mainly in temperate regions of the Northern Hemisphere, especially in Europe, whereas Asia and North America also contain several records, and the Southern Hemisphere is clearly less represented [21]. Nine species have been recorded from China, mainly from northern or temperate forest habitats.
Taken together, the contrasting trophic modes and substrate preferences of these genera suggest that heterogeneous forest environments may provide multiple ecological niches for macrofungi and harbor overlooked taxonomic diversity. Northeastern China contains extensive temperate forest ecosystems with diverse dominant tree species, litter inputs, and forest-floor microhabitats. However, macrofungal diversity in Shenyang and adjacent areas remains insufficiently investigated compared with that of several other regions of China. Field surveys were therefore conducted to document poorly known macrofungal diversity in local forest ecosystems, especially agaric taxa occurring on the forest floor in different habitat contexts. During these surveys, collections referable to the three genera mentioned above were obtained and confirmed to represent three previously undescribed species based on morphological characteristics and phylogenetic analyses of ITS and LSU sequences. The aim of the present study is to clarify the taxonomic identities and phylogenetic positions of these new taxa and to provide detailed descriptions, illustrations, and ecological notes for them.

2. Materials and Methods

2.1. Specimen Collections

Specimens were collected from Qipanshan Forest Park, Shenyang City; Wangbin Township, Shenyang City; and Shenxiangu, Yongling Town, Xinbin County, Liaoning Province, northeastern China. For each collection, data on specimen locality, associated vegetation, ecological habit, collector, and collection date were recorded. When multiple specimens of the same species were collected from the same general area on the same date, their collecting points were separated by at least several hundred meters. Photographs of the basidiomata and their habitats were taken in the field. The specimens were subsequently dried without delay and preserved in sealed bags. All examined specimens were deposited in the herbarium of the Institute of Applied Ecology, Chinese Academy of Sciences (IFP).

2.2. Morphological Studies

Macromorphological characters were observed under a stereo microscope (Nikon SMZ 645, Tokyo, Japan), and colour terms for basidiomata follow Kornerup and Wanscher [22]. Micromorphological characters were examined at 1000× magnification using a light microscope (Nikon Eclipse 80i, Tokyo, Japan). Hand-cut sections were mounted in Cotton Blue (CB), Melzer’s reagent (IKI), and 3% KOH. Line drawings of microscopic structures were made with the aid of a drawing tube. CB was used to determine cyanophily of the cell walls [23,24], IKI to test amyloid, inamyloid or dextrinoid reactions, and KOH to clear tissues, reveal pigments, and facilitate observation of tissue structures. Basidiospore length and width were measured excluding the apiculus, following common practice in fungal taxonomy. The following abbreviations are used: L = mean spore length, W = mean spore width, Q = L/W ratio, and n = number of basidiospores measured from a given number of specimens.

2.3. DNA Extraction, Amplification, and Sequencing

Total genomic DNA was extracted from dried specimens using the Rapid Fungal Genomic DNA Isolation Kit (Demeter Biotech Co., Ltd., Beijing, China) following the manufacturer’s instructions. The nuclear ribosomal internal transcribed spacer (ITS) region was amplified by polymerase chain reaction (PCR) using the primer pair ITS1 (5′-TCCGTAGGTGAACCTGCGG-3′) and ITS4 (5′-TCCTCCGCTTATTGATATGC-3′) [25,26]. The PCR program consisted of an initial denaturation at 94 °C for 5 min, followed by 34 cycles of 95 °C for 35 s, 55 °C for 55 s, and 72 °C for 45 s, with a final extension at 72 °C for 10 min. To further assess the phylogenetic placement of the collections, the nuclear ribosomal large subunit (LSU) region was amplified and sequenced using the primers LR0R (5′-ACCCGCTGAACTTAAGC-3′) and LR7 (5′-TACTACCACCAAGATCT-3′). The PCR program for LSU consisted of an initial denaturation at 94 °C for 2 min, followed by 35 cycles of 94 °C for 30 s, 48 °C for 1 min, and 72 °C for 1.5 min, with a final extension at 72 °C for 10 min [27]. PCR products were sequenced by the Beijing Genomics Institute (BGI). All newly generated sequences were assembled and manually edited in SeqMan v.7.1.0. Sequence similarity searches were conducted against GenBank using the web-based NCBI BLASTn program (National Center for Biotechnology Information, Bethesda, MD, USA; accessed on 10 March 2026) [28]. After base-calling quality was checked, the new sequences were submitted to GenBank.

2.4. Phylogenetic Analyses

Sequences used for phylogenetic comparisons were selected from GenBank based on BLAST similarity, taxonomic relevance, availability of voucher or specimen information, and representation of major lineages within each genus or allied group. Whenever available, sequences derived from type materials and reliably identified voucher specimens were given priority. The newly obtained sequences and selected GenBank sequences were aligned using MAFFT v.7 [29] (Table 1). All sequences were checked and adjusted to the same orientation prior to alignment, and the resulting alignments were manually refined in MEGA v.7.0 [30]. For the Descolea ITS + LSU combined dataset, Hebeloma plesiocistum and H. theobrominum were selected as outgroups following Kuhar et al. [2]. For the Leucocoprinus combined dataset, Agaricus bisporus was selected as the outgroup following Asif et al. [9]. For the Tephrocybe combined dataset, Rhizocybe alba and R. vermicularis were selected as outgroups following Khan et al. [31]. For each genus, concatenated alignments were generated from the combined ITS and LSU datasets and partitioned into four regions: ITS1, 5.8S, ITS2, and nLSU. The best-fit substitution model for each partition was selected using IQ-TREE v.2.4.0. For Descolea, the selected models were K2P+G4 for ITS1, JC for 5.8S, TPM2+G4 for ITS2, and K2P for nLSU. For Leucocoprinus, the selected models were TPM2u+F+I+G4 for ITS1, TIM2e+R2 for 5.8S, TVM+F+G4 for ITS2, and TIM3e+I+R2 for nLSU. For Tephrocybe, the selected models were HKY+F+G4 for ITS1, K2P+I for 5.8S, TPM2u+F+G4 for ITS2, and TN+F for nLSU. Phylogenetic analyses were performed for each dataset using Bayesian inference (BI) and maximum likelihood (ML) methods. BI analyses were conducted in MrBayes v.3.0 using a Markov chain Monte Carlo (MCMC) algorithm. Four chains were run from random starting trees for 10 million generations, with trees sampled every 1000 generations, until the average standard deviation of split frequencies fell below 0.01 [32]. The first 25% of sampled trees were discarded as burn-in, and the remaining trees were used to calculate Bayesian posterior probabilities (BPPs). ML analyses were performed using the same datasets in RAxML-HPC BlackBox v.8.2.12, and branch support was assessed with 1,000 non-parametric bootstrap replicates [33]. Phylogenetic trees were visualized using FigTree v.1.4.4 [34]. Branches with ML bootstrap support (MLBS) ≥ 70% and/or BPPs ≥ 0.95 were considered significantly supported. The alignments and trees were deposited in TreeBASE (No. S32652).

3. Results

3.1. Phylogeny

BLAST searches of the newly generated ITS sequences recovered several close GenBank matches, including unpublished, directly submitted, isolate-derived, and environmental sequences. These results indicate that the corresponding lineages, or closely related lineages, may have previously been detected in GenBank. However, BLAST parameters were used only to identify closely matching sequences and were not treated as evidence of conspecificity by themselves. Because these accessions lack formal taxonomic treatment and/or associated basidiomatal morphological data, they were not used as primary evidence for species delimitation.
The combined ITS and LSU dataset was used to infer the phylogenetic positions of the three new species. Because these taxa belong to different genera, three separate phylogenetic trees were reconstructed. The Descolea dataset comprised 1476 characters, including 602 from ITS and 874 from LSU. Of these, 78 were constant, 1195 were parsimony-uninformative variable, and 203 were parsimony-informative. The dataset included two sequences from the new species, 28 sequences of Descolea [36,37,38,39,40], and two outgroup sequences, viz. H. plesiocistum and H. theobrominum [41,42]. The Leucocoprinus dataset comprised 1539 characters, including 729 from ITS and 810 from LSU. Of these, 841 were constant, 204 were parsimony-uninformative variable, and 494 were parsimony-informative. The dataset included two sequences from the new species, 102 sequences of Leucocoprinus [43,44,45,46,47,48,49,50,51,52,53,54,55,56,57,58,59,60,61,62,63,64,65,66,67,68,69,70,71,72,73,74,75,76,77,78], and two sequences of A. bisporus as the outgroup [35]. Because the generic limits of Leucocoprinus have recently undergone substantial revision, a conservative treatment was adopted for the Leucocoprinus dataset. The dataset included sequences identified or formally treated as Leucocoprinus in GenBank and the cited taxonomic studies, because BLAST searches placed the new species among Leucocoprinus sequences and the collections showed morphological affinity with the traditional concept of the genus. This treatment was used to evaluate the phylogenetic placement of the new species and does not represent a broader revision of Leucocoprinus, Leucoagaricus, or allied genera. The Tephrocybe dataset comprised 1490 characters, including 629 from ITS and 861 from LSU. Of these, 298 were constant, 945 were parsimony-uninformative variable, and 247 were parsimony-informative. The dataset included two sequences from the new species, 21 sequences of Tephrocybe [81,82,83], and two outgroup sequences, R. alba and R. vermicularis [79,80]. A 50% majority-rule consensus phylogram was generated.
Maximum likelihood and Bayesian analyses recovered similar topologies; therefore, only the ML trees are presented, with MLBS and BPPs shown at the nodes. In the phylogenetic tree inferred from the combined ITS and LSU dataset, the two collections of D. laevis formed a distinct lineage with strong support (ML = 91%, BPP = 1.00), and this lineage was grouped with D. flavoannulata and D. indoquercina (Figure 1). The two collections of L. shenyangensis formed a distinct lineage with full support (ML = 100%, BPP = 1.00) (Figure 2). The two collections of T. umbonata also formed a distinct, fully supported lineage (ML = 100%, BPP = 1.00), which was resolved as sister to the clade comprising T. coracina, T. fibrosipes, and T. ochraceobrunnea (ML = 100%, BPP = 1.00) (Figure 3). These results supported the taxonomic placement of the three new species.

3.2. Taxonomy

Descolea laevis Z.Q. You & H.S. Yuan, sp. nov. (Figure 4 and Figure 5).
Fungal Names: FN 573717.
Diagnosis. Similar to Descolea indoquercina, but distinguished by smaller basidiospores (10.7–12.4 × 6.5–8.5 μm), shorter basidia (24.8–45.2 × 8.1–12.3 μm), the presence of cheilocystidia and pleurocystidia, a smooth pileus lacking scales or warts, and a thick, immovable annulus.
Type. CHINA. Liaoning Province, Shenyang City, Qipanshan Forest Park, on the ground in mixed Quercus mongolica-Pinus koraiensis forests, 18 September 2024, Yuan 21,385 (holotype IFP 020246).
Etymology. laevis (Lat.), referring to the nearly smooth pileus surface lacking scales or warts.
Description. Basidiomes: medium-sized, solitary to scattered. Pileus: convex to plano-convex when young, becoming plane with a shallow central depression at maturity; surface nearly smooth, lacking scales or warts; disc greyish brown to reddish brown (9D4–9D5), margin pale yellow to light yellow (4A3–4A4); margin incurved when young, becoming straight to slightly upturned at maturity, with conspicuous irregular striations; when dry, pileus markedly shrunken, at first with a brown to dark brown centre (6E5–6F5) and a yellowish brown to linoleum brown surface (5E7–5E8), later with a greyish brown to dark brown centre (7F3–7F4) and a light brown to golden brown margin (5D6–5D7). Context: thin, slightly thicker at the centre, light orange to greyish orange (5A4–5B4). Lamellae: adnexed to adnate, rather crowded (≥14 L + l/cm), relatively broad, with lamellulae; edge uneven and slightly undulate, reddish white to pinkish white (7A2–8A2); when dry, distinctly undulate, wrinkled, and twisted, becoming tan to cognac brown (6E6–6E7). Stipe: central, cylindrical, slightly enlarged towards the base; surface dry, with conspicuous longitudinal fibrils; upper part pale, yellowish white to pale yellow (4A2–4A3), lower part light orange (5A4–5A5); basal mycelium white. Annulus: rather thick, immovable, median to superior, persistent, slightly flaring, pale yellow to pastel yellow (3A3–3A4). Odour: mild, not distinctive.
Basidiospores: (10.1–)10.7–12.4(–13.3) × (3.8–)6.5–8.5(–9.2) μm, L = 11.35 μm, W = 7.21 μm, Q = 1.25–1.85 (n = 60/2); amygdaliform to limoniform, guttulate, verrucose, slightly thick-walled, with a prominent papilla and a broad, smooth apiculus, dextrinoid, acyanophilous.
Basidia: 24.8–45.2 × 8.1–12.3 μm, narrowly clavate to subcylindrical, hyaline, thin-walled, 4-sterigmate; sterigmata slender, 3.8–6.1 × 0.8–2.2 μm; contents granular to guttulate; basal septum simple.
Cheilocystidia: 18.7–34.2 × 4.8–7.3 μm, clavate to narrowly clavate, sometimes subcylindrical, thin-walled.
Pleurocystidia: 27.6–45.3 × 4.8–12.2 μm, fusiform to narrowly fusiform, sometimes clavate, apex occasionally attenuated, thin-walled.
Pileipellis: epicutis epithelium-like, 60–166 µm thick, composed of inflated, fusiform to subglobose elements, 7.8–19.1 × 3.7–11.2 µm, surface heavily encrusted with rusty-brown pigment; subcutis composed of cylindrical, thin-walled hyphae, 2.8–10.3 µm wide, with encrusting pigment; clamp connections present.
Annulus: composed of cylindrical, smooth, hyaline, thin-walled, interwoven hyphae, 1–12 μm wide, with clamp connections.
Additional specimen (paratype) examined. CHINA. Liaoning Province, Shenyang City, Qipanshan Forest Park, on the ground in mixed Quercus mongolica-Pinus koraiensis forests, 18 September 2024, Yuan 21,391 (IFP 020247).
Leucocoprinus shenyangensis Z.Q. You & H.S. Yuan, sp. nov. (Figure 6 and Figure 7).
Fungal Names: FN 573718.
Diagnosis. Morphologically similar to Leucocoprinus dacrytus, but distinguished by basidiospores lacking a germ pore, thin-walled cheilocystidia, and a hymenidermal epicutis of the pileipellis composed of subglobose to short ellipsoid cells with narrowly clavate to clavate terminal elements, gradually passing into an interwoven hyphal layer below.
Type. CHINA. Liaoning Province, Shenyang City, Wangbin Township, on the ground in mixed forests, 6 August 2024, Yuan 20,026 (holotype IFP 020248).
Etymology. shenyangensis (Lat.), named after the collection site of the type specimen, Shenyang City.
Description. Basidiomes: small, solitary to scattered. Pileus: ovoid-campanulate to campanulate when young, expanding to convex to nearly plane, with a low but distinct obtuse umbo at the centre; surface exuding droplets of various sizes, light yellow to orange (4A4–5A7), scattered with minute light grey (8C1–8D1) granulose to squamulose elements; disc greyish brown to reddish brown (8D3–8D4), margin paler, white to yellowish white (3A1–3A2); margin distinctly striate at maturity, the striations extending over much of the pileus radius; when dry, pileus markedly shrunken and radially plicate-striate, with the disc darker, sunburn to tan brown (6D5–6E5), and the margin yellowish white to pale yellow (4A2–4A3). Context: thin, white to orange white (5A1–5A2). Lamellae: free, rather crowded, subequal in length, with entire edges, white to yellowish white (1A1–1A2); when dry, slightly crisped, light yellow to champagne yellow (4A4–4B4). Stipe: central, cylindrical, slender, slightly enlarged towards the base; surface smooth to finely fibrillose, smoother above the annulus; lower part pale yellow, becoming white upwards (4A3–4A1); basal mycelium white. Annulus: thin, membranous, median to superior, persistent, orange white to pale orange (5A2–5A3); surface sometimes exuding light yellow to orange (4A4–5A7) droplets. Odour: not recorded.
Basidiospores: (3.1–)3.5–7.2(–9.0) × (2.5–)2.8–4.5(–5.5) μm, L = 6.14 μm, W = 3.79 μm, Q = 1.01–2.40 (n = 60/2); ovoid to ellipsoid-oblong in frontal view, amygdaliform to limoniform in side view, smooth, hyaline, without a distinct germ pore, slightly thick-walled, dextrinoid, cyanophilous.
Basidia: 15.8–26.3 × 6.7–9.2 μm, narrowly clavate to subcylindrical, hyaline, thin-walled, usually 4-sterigmate; basal septum simple.
Cheilocystidia: 23.2–36.8 × 4.8–8.3 μm, narrowly clavate to subcylindrical, often slightly curved, apex rounded to slightly acute, base gradually narrowed, hyaline, thin-walled.
Pleurocystidia: absent.
Pileipellis: epicutis hymenidermal, composed of rather regularly arranged, subglobose to short ellipsoid cells, with terminal elements narrowly clavate to clavate, suberect to erect, (14.2–)17.1–19.8(–22.9) × (2.7–)5.1–6.2(–8.4) μm, apex rounded to slightly acute, sometimes containing pale brown to brown intracellular pigment; subcutis composed of interwoven, subcylindrical, smooth, hyaline, slightly thick-walled hyphae, 2–10 μm wide, gradually passing into the context; clamp connections not observed.
Annulus: composed of cylindrical, smooth, hyaline, occasionally branched hyphae, 1.2–13.5 µm wide, with occasional rusty brown to reddish-brown intracellular pigment, without clamp connections.
Additional specimens (paratype) examined. CHINA. Liaoning Province, Shenyang City, Qipanshan Forest Park, on the ground in Populus forest, 6 August 2024, Yuan 19,598 (IFP 020249).
Tephrocybe umbonata Z.Q. You & H.S. Yuan, sp. nov. (Figure 8 and Figure 9).
Fungal Names: FN 573719.
Diagnosis. Similar to Tephrocybe platypus, but distinguished by a distinct obtuse umbo at the centre of the pileus, basidia usually 4-spored, and pleurocystidia with slightly acute apices.
Type. CHINA. Liaoning Province, Xinbin County, Yongling Town, Shenxiangu, on the ground in Larix kaempferi forest, 4 October 2024, Yuan 21,497 (holotype IFP 020250).
Etymology. umbonata (Lat.), referring to the distinct umbo at the centre of the pileus.
Description. Basidiomes: small, solitary to subgregarious. Pileus: hemispherical to convex when young, becoming nearly plane with a distinct obtuse umbo to low papilla at the centre; surface dry, smooth or slightly finely fibrillose; disc reddish white to reddish grey (8A2–8B2), margin paler, orange white to greyish orange (5A2–5B4); margin with conspicuous wavy translucent striations, sometimes shallowly cracked; when dry, pileus shrunken, with a distinct central depression and an oak brown to linoleum brown (5D6–5E7) surface. Context: thin, white to reddish white (7A1–7A2). Lamellae: rather distant to moderately crowded, with nearly entire edges, orange white to pale orange (5A2–5A3); when dry, becoming dark blonde to mustard brown (5D4–5E6). Stipe: central, subcylindrical, slender, often curved, fibrous; surface smooth at the base, slightly roughened and covered with white granulose remnants towards the apex, white to pale yellow (4A1–4A3); basal mycelium white. Annulus: absent. Odour: not recorded.
Basidiospores: (4.2–)4.7–7.5(–8.1) × (2.0–)2.8–4.1(–4.3) μm, L = 6.64 μm, W = 3.49 μm, Q = 1.13–2.67 (n = 60/2); ellipsoid to ovoid in frontal view, amygdaliform to limoniform in side view, smooth, hyaline, guttulate, moderately thick-walled, with a distinct lateral apiculus, inamyloid, cyanophilous.
Basidia: 19.8–31.2 × 4.9–9.3 μm, clavate to narrowly clavate, rarely subcylindrical, slightly curved, hyaline, thin-walled, 4-sterigmate; contents granular, non-metachromatic; basal clamp connections present.
Cheilocystidia: absent.
Pleurocystidia: 14.7–39.3 × 2.0–5.1 μm, lageniform to narrowly lageniform, sometimes fusiform, rarely subcylindrical, apex slightly acute, base gradually narrowed, thin-walled.
Marginal cells: 16.7–27.3 × 3.8–7.2 μm, resembling pseudocystidia, narrowly clavate to cylindrical, sometimes elongated-cylindrical, occasionally irregularly curved.
Pileipellis: epicutis a non-gelatinized cutis, rather compact, composed of slender, repent, parallel to subparallel hyphae, 0.5–5.1 μm wide, locally slightly interwoven, without evident erect terminal elements; subcutis indistinct, composed of loosely arranged hyphae, 4.2–12.3 μm wide, gradually passing into the context; clamp connections present.
Annulus: absent.
Additional specimens (paratype) examined. CHINA. Liaoning Province, Xinbin County, Yongling Town, Shenxiangu, on the ground in Larix kaempferi forest, 4 October 2024, Yuan 21,507a (IFP 020251).

4. Discussion

In this study, three new species belonging to Descolea, Leucocoprinus, and Tephrocybe were confirmed from the areas surrounding Shenyang, Liaoning Province, China, based on morphological observations and molecular phylogenetic analyses. Phylogenetic results support these taxa as independent lineages within their respective genera, and morphological comparisons further show that they can be clearly distinguished from their allied species.
In the phylogenetic tree, the two collections of Descolea laevis were placed within a clade including D. flavoannulata, D. indoquercina, D. quercina, and D. pretiosa (ML = 100%, BPP = 1.00). These species share annulate basidiomata, amygdaliform to limoniform basidiospores bearing verrucose ornamentation, a conspicuous papilla, and a smooth apiculus. D. laevis differs from D. indoquercina by its smaller basidiospores and the presence of both pleurocystidia and cheilocystidia. D. laevis differs from D. flavoannulata in possessing pleurocystidia and a thick, immovable annulus, although the basidia are similar in size [84]. D. quercina is distinguished by more coarsely verrucose basidiospores and by having 2-spored basidia with clamp connections at the base [3]. D. pretiosa differs from D. laevis in having a pileus densely covered with ochraceous brown to pale ochraceous yellow scales, lacking cheilocystidia, and having an epithelium of clavate cells in the pileipellis [1].
Leucocoprinus shenyangensis is readily recognized by its slender stipe, overall pale yellow to white basidiomata, and pileus with a greyish brown to reddish brown blunt umbo. Phylogenetically, L. shenyangensis formed a distinct and fully supported lineage in the combined ITS and LSU analyses. It was placed within a well-supported clade including L. dacrytus, L. brunneodiscus, L. margaritifer, L. silvestris, L. glaber, L. karjaticus, L. flavirobustus, and L. tangerinus (BPP = 0.97), with L. dacrytus recovered as its closest relative. The two species are similar in basidiospore size and shape, but L. dacrytus differs in having slightly larger, thick-walled cheilocystidia and a cutis-like pileus covering [80]. L. brunneodiscus differs in its orange basidiomata, 2-spored basidia, clavate cheilocystidia, and subcutis- to cutis-like pileus covering [49]. L. margaritifer is distinguished by its brownish pink pileus, larger, branched cheilocystidia, and trichodermal pileus covering [43]. L. silvestris differs in having rosy to pink-ocher squamules on the pileus, a pinkish stipe base, branched cheilocystidia, and a trichodermal pileus covering [43]. L. glaber, L. karjaticus, and L. flavirobustus form a small allied subclade, and share 2-spored basidia, cylindrical to capitate cheilocystidia composed of cell chains, and a pileus covering composed of cylindrical element chains. However, L. glaber has a light orange pileus margin [52], L. karjaticus has larger basidiomata and yellowish brown to reddish brown squamules [65]. L. flavirobustus has a darker reddish brown pileus centre [52]. L. tangerinus is readily distinguished by the presence of pleurocystidia [85]. The exudation of colored droplets is not unique to L. shenyangensis, as similar golden yellow to brown, orange-white, or amber droplets have also been reported in some allied taxa, including L. dacrytus, L. brunneodiscus, and L. margaritifer [43,49,80]. However, taxa reported to exude colored droplets were not recovered as a separate, well-supported monophyletic group in the present ITS and LSU phylogeny, but were placed within a broader clade that also included species with colorless droplets or without reported droplet exudation. Thus, this feature is not interpreted as evidence for an independent infrageneric lineage.
Tephrocybe umbonata formed a fully supported clade with T. ochraceobrunnea, T. fibrosipes, and T. coracina, which was sister to T. rancida. These allied taxa share several characters, including 4-spored basidia with conspicuous siderophilous granules, absence of cheilocystidia, and a pileipellis composed of filamentous hyphae. Nevertheless, T. umbonata differs from the other similar species in having pleurocystidia with slightly pointed apices. Although T. ochraceobrunnea and T. fibrosipes resemble T. umbonata in their relatively pale basidiomata, T. ochraceobrunnea has broader basidiospores (4.0–5.0 μm wide), whereas T. fibrosipes has larger basidiospores (7–9 × 4–5 μm) [86,87]. By contrast, T. coracina and T. rancida have darker basidiomata, from deep sooty-brown to nearly black. T. umbonata further differs from T. coracina in having narrower basidiospores and from T. rancida in lacking a pseudorrhiza at the stipe base [88].
The three new species were collected from distinct forest-floor habitats around Shenyang, including mixed Quercus mongolica-Pinus koraiensis forests, Populus forests or mixed forests, and Larix kaempferi forests. Descolea laevis was collected on the ground in a mixed forest of Q. mongolica and P. koraiensis. BLAST results suggest that this species, or a closely related lineage, may have been detected previously in forest soils from Anhui Province, China (GenBank accession no. PX601745), and from Changbai Mountain, Jilin Province, China (e.g., GenBank accession no. JQ666800). D. indoquercina, D. flavoannulata, and D. quercina have all been reported from Himalayan forests dominated by Quercus spp. [84]. This pattern suggests that closely related taxa of Descolea may occur across a relatively broad range of forest habitats, including Quercus-dominated and mixed forests. Leucocoprinus shenyangensis and its allied taxa are saprotrophic and mainly occur on the forest floor and other organic-rich substrates [43,65,85]. BLAST results indicate that this species, or a closely related lineage, may be more widely distributed in China (e.g., GenBank accession no. PX513137). L. shenyangensis was collected on the ground in a Populus and a mixed forest, whereas L. dacrytus occurs on decayed wood of Quercus rubra in deciduous forest [89], indicating differentiation in substrate use despite a shared forest habitat background. Lyophyllaceae includes ecologically diverse taxa, and multigene phylogenetic studies have suggested repeated transitions from free-living saprotrophic to parasitic or mutualistic lifestyles within the family [19]. Several lyophylloid taxa occupy specialized substrates, including pyrophilous habitats, as shown by Tephrocybe anthracophila among fungi fruiting after fire [90], and nitrogen-rich substrates, as shown by the ammonia fungus T. tesquorum in urea-treated forest soil [91]. A blue reaction with p-dimethylaminobenzaldehyde (PDAB) has also been discussed as a macroscopic chemical character in lyophylloid fungi [92]. In contrast to these specialized ecological patterns, several species of Tephrocybe have been reported from relatively general forest-floor habitats, including soil, humus, litter layers, and adjacent woodland ground [21,88]. T. umbonata was collected on the ground in a Larix kaempferi forest. BLAST results suggest that this species, or a closely related lineage, may have been detected previously in Virginia, USA (e.g., GenBank accession no. PX925887), and in beech-maple forests from northeastern Ohio, USA (GenBank accession no. FM999645) [93]. By contrast, T. rancida is more common in deciduous litter, especially in Fagus forests, and only occasionally occurs in coniferous forests [88]. Together, these habitat records place the three new species in a more specific regional and ecological context and support continued targeted taxonomic surveys of macrofungi in Shenyang and adjacent areas.

Author Contributions

Investigation and writing draft: Z.-Q.Y. Data measurement and analysis: L.-J.Z. Conceptualization and supervision: H.-S.Y. and H.B.L. All authors have read and agreed to the published version of the manuscript.

Funding

This research was financed by the National Natural Science Foundation of China (Project No. 32570013), and the Fund of CAS Key Laboratory of Forest Ecology and Silviculture, Institute of Applied Ecology, Chinese Academy of Sciences (No. KLFES-2028).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The sequences generated in the present study were submitted to GenBank through the NCBI website (https://www.ncbi.nlm.nih.gov/; accessed on 10 March 2026), and the accession numbers were listed in Table 1. The nomenclatural information for the three new species was registered in Fungal Names (https://nmdc.cn/fungalnames/; accessed on 17 April 2026). The sequence alignments and phylogenetic trees were deposited in TreeBASE (https://www.treebase.org/; accessed on 11 May 2026). All other data supporting the findings of this study are included in the article.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Maximum likelihood tree illustrating the phylogeny of Descolea and related taxa based on the combined ITS and LSU nuclear rDNA sequences dataset. Branches are labeled with maximum likelihood bootstrap values ≥ 50% and Bayesian posterior probabilities ≥ 0.95; lower values are indicated by “-”. Specimen numbers and countries of collection are indicated after species names. New species in bold (black).
Figure 1. Maximum likelihood tree illustrating the phylogeny of Descolea and related taxa based on the combined ITS and LSU nuclear rDNA sequences dataset. Branches are labeled with maximum likelihood bootstrap values ≥ 50% and Bayesian posterior probabilities ≥ 0.95; lower values are indicated by “-”. Specimen numbers and countries of collection are indicated after species names. New species in bold (black).
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Figure 2. Maximum likelihood tree illustrating the phylogeny of Leucocoprinus and related taxa based on the combined ITS and LSU nuclear rDNA sequences dataset. Branches are labeled with maximum likelihood bootstrap values ≥ 50% and Bayesian posterior probabilities ≥ 0.95; lower values are indicated by “-”. Specimen numbers and countries of collection are indicated after species names. New species in bold (black).
Figure 2. Maximum likelihood tree illustrating the phylogeny of Leucocoprinus and related taxa based on the combined ITS and LSU nuclear rDNA sequences dataset. Branches are labeled with maximum likelihood bootstrap values ≥ 50% and Bayesian posterior probabilities ≥ 0.95; lower values are indicated by “-”. Specimen numbers and countries of collection are indicated after species names. New species in bold (black).
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Figure 3. Maximum likelihood tree illustrating the phylogeny of Tephrocybe and related taxa based on the combined ITS and LSU nuclear rDNA sequences dataset. Branches are labeled with maximum likelihood bootstrap values ≥ 50% and Bayesian posterior probabilities ≥ 0.95; lower values are indicated by “-”. Specimen numbers and countries of collection are indicated after species names. New species in bold (black).
Figure 3. Maximum likelihood tree illustrating the phylogeny of Tephrocybe and related taxa based on the combined ITS and LSU nuclear rDNA sequences dataset. Branches are labeled with maximum likelihood bootstrap values ≥ 50% and Bayesian posterior probabilities ≥ 0.95; lower values are indicated by “-”. Specimen numbers and countries of collection are indicated after species names. New species in bold (black).
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Figure 4. Fresh basidiomata of Descolea laevis (holotype, IFP 020246; specimen no. Yuan 21,385). Photos by Hai-Sheng Yuan.
Figure 4. Fresh basidiomata of Descolea laevis (holotype, IFP 020246; specimen no. Yuan 21,385). Photos by Hai-Sheng Yuan.
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Figure 5. Microscopic features of Descolea laevis (drawn from holotype, IFP 020246; specimen no. Yuan 21,385) (A) Basidiospores (B) Basidia (C) Cheilocystidia (D) Pleurocystidia (E) Pileipellis elements. Scale bars: 10 μm.
Figure 5. Microscopic features of Descolea laevis (drawn from holotype, IFP 020246; specimen no. Yuan 21,385) (A) Basidiospores (B) Basidia (C) Cheilocystidia (D) Pleurocystidia (E) Pileipellis elements. Scale bars: 10 μm.
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Figure 6. Fresh basidiomata of Leucocoprinus shenyangensis (holotype, IFP 020248; specimen no. Yuan 20,026). Photos by Hai-Sheng Yuan.
Figure 6. Fresh basidiomata of Leucocoprinus shenyangensis (holotype, IFP 020248; specimen no. Yuan 20,026). Photos by Hai-Sheng Yuan.
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Figure 7. Microscopic features of Leucocoprinus shenyangensis (drawn from holotype, IFP 020248; specimen no. Yuan 20,026) (A) Pileipellis elements (B) Basidiospores (C) Basidia (D) Cheilocystidia. Scale bars: 10 μm.
Figure 7. Microscopic features of Leucocoprinus shenyangensis (drawn from holotype, IFP 020248; specimen no. Yuan 20,026) (A) Pileipellis elements (B) Basidiospores (C) Basidia (D) Cheilocystidia. Scale bars: 10 μm.
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Figure 8. Fresh basidiomata of Tephrocybe umbonata (holotype, IFP 020250; specimen no. Yuan 21,497). Photos by Hai-Sheng Yuan.
Figure 8. Fresh basidiomata of Tephrocybe umbonata (holotype, IFP 020250; specimen no. Yuan 21,497). Photos by Hai-Sheng Yuan.
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Figure 9. Microscopic features of Tephrocybe umbonata (drawn from holotype, IFP 020250; specimen no. Yuan 21,497) (A) Basidiospores (B) Basidia and Marginal cells (C) Pleurocystidia (D) Pileipellis elements. Scale bars: 10 μm.
Figure 9. Microscopic features of Tephrocybe umbonata (drawn from holotype, IFP 020250; specimen no. Yuan 21,497) (A) Basidiospores (B) Basidia and Marginal cells (C) Pleurocystidia (D) Pileipellis elements. Scale bars: 10 μm.
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Table 1. Species and GenBank numbers used in phylogenetic analysis in this study.
Table 1. Species and GenBank numbers used in phylogenetic analysis in this study.
Species NameITSLSUSpecimen No.CountryReferences
Agaricus bisporus (J.E. Lange) ImbachMH859080MH870797CBS11668The NetherlandsVu et al., 2019 [35]
A. bisporusMH859081MH870798CBS11768The NetherlandsVu et al., 2019 [35]
Descolea alba (Bull.) Kuhar, Nouhra & M.E. Sm.AJ296296/MA-Fungi40133PortugalMartín and Moreno 2001 [36]
D. albella (Massee & Rodway) Kuhar, Nouhra & M.E. Sm.PV650409/MEL2523797AAustraliaNCBI Database
D. angustispora (A.A. Francis & Bougher) Kuhar, Nouhra & M.E. Sm.DQ328058/H7216AustraliaFrancis and Bougher 2004 [37]
D. antarctica SingerAF325647/HorakNZ5182New ZealandPeintner et al., 2001 [38]
D. archeureta (Halling) Kuhar, Nouhra & M.E. Sm.KY523096/MES-1786ChileKuhar et al., 2017 [2]
D. archeuretaKY523092/MES-1584ChileKuhar et al., 2017 [2]
D. australiensis (G.W. Beaton, Pegler & T.W.K. Young) Kuhar, Nouhra & M.E. Sm.AF325627/Claridge2679AustraliaPeintner et al., 2001 [38]
D. australiensisAF325628/Claridge2621AustraliaPeintner et al., 2001 [38]
D. brunnea (E. Horak) Kuhar, Nouhra & M.E. Sm.OP339609/FLAS:F-70573-MES-4047ChileNCBI Database
D. brunneaOP339580/FLAS:F-70526-MES-4000ChileNCBI Database
D. ferruginea (J.W. Cribb) Kuhar, Nouhra & M.E. Sm.DQ328083/H5803AustraliaKuhar et al., 2017 [2]
D. flavoannulata (Lj.N. Vassiljeva) E. HorakPP345433/KV-20-15IndiaNCBI Database
D. gunnii (Berk. ex Massee) E. HorakAF325652/NZ752New ZealandPeintner et al., 2001 [38]
D. gunniiAF325653/NZ2042New ZealandPeintner et al., 2001 [38]
D. indoquercina S. Choudhary, P. Uniyal & Y.P. SharmaOR979479/SC/PU/12IndiaNCBI Database
D. inferna Kuhar, Nouhra & M.E. Sm.NR154032/CORD:MES1315 (T)ArgentinaKuhar et al., 2017 [2]
D. infernaOP339680/FLAS:F70672MES-4146ChileNCBI Database
D. laevisZ.Q. You & H.S. YuanPZ235489PZ235510Yuan21385 (T)ChinaPresent study
D. laevisPZ235490PZ235511Yuan21391ChinaPresent study
D. maculata BougherDQ192181DQ457664E8078(PERTH)AustraliaMatheny et al., 2006 [39]
D. maculataAF325651/E4986AustraliaPeintner et al., 2001 [38]
D. phlebophora E. HorakAF325657/E4912AustraliaPeintner et al., 2001 [38]
D. phlebophoraJX178627/OTA:60177New ZealandTeasdale et al., 2013 [40]
D. pretiosa E. HorakMN267170/ZT9000IndonesiaNCBI Database
D. quercina J. Khan & NaseerMF966638/LAH35219PakistanKhan et al., 2017 [3]
D. quercinaMF966637MF966635LAH35218PakistanKhan et al., 2017 [3]
D. recedens (Sacc.) SingerAF325648/E4591AustraliaPeintner et al., 2001 [38]
D. recedensAF325649/E4459AustraliaPeintner et al., 2001 [38]
D. tenuipes (Setch.) Neville & PoumaratAF325624/Trappe24776AustraliaPeintner et al., 2001 [38]
D. tenuipesAF325623/Trappe24524AustraliaPeintner et al., 2001 [38]
Hebeloma plesiocistum Beker, U. Eberh. & VilaNR119686/LIP:JVG1021214 (T)SpainEberhardt et al., 2009 [41]
H. theobrominum Quadr.NR120177/ROHB:925LQ (T)ItalyEberhardt et al., 2013 [42]
Leucocoprinus albosquamosus (Y.R. Ma, Z.W. Ge & T.Z. Liu) M. Asif, Saba & VellingaOM976879OM976865CFSZ20662ChinaMa et al., 2022 [8]
L. antillarum Justo, Bizzi & AngeliniMN482989/CA9Dominican RepublicJusto et al., 2021 [43]
L. asiaticus (Qasim, Nawaz & Khalid) M. Asif, Saba & VellingaKP164972/LAH5872011PakistanGe et al., 2015 [44]
L. atroviridis (Y.R. Ma, Z.W. Ge & T.Z. Liu) M. Asif, Saba & VellingaOM976852OM976868SYAUFUNGI073ChinaMa et al., 2022 [8]
L. aurantiopileus Maula, Asif, A.K. Rani & AfshanPP383877PP583799LAH38131 (T)PakistanMaula et al., 2026 [45]
L. aurantioruber (Y.R. Ma, Z.W. Ge & T.Z. Liu) M. Asif, Saba & VellingaOM976875OM976863CFSZ19756ChinaMa et al., 2022 [8]
L. badius (S. Hussain, Pfister, Afshan & Khalid) M. Asif, Saba & VellingaKU647734/LAHSH210PakistanHussain et al., 2018 [46]
L. beninensis SarawiPX634061/SeSa186BeninSarawi et al., 2026 [47]
L. birnbaumii (Corda) SingerPQ321881PQ319807HTBM1220ChinaYang et al., 2024 [10]
L. brebissonii (Godey) Locq.AF482859AY176446ecv1784LFranceVellinga et al., 2003 [48]
L. brunneocanus (Fei Yu, Jun F. Liang & Z.W. Ge) M. Asif, Saba & VellingaKP096237/ZWGe97ChinaGe et al., 2015 [44]
L. brunneodiscus (A.K. Dutta & K. Acharya) Kun L. Yang, Jia Y. Lin & Zhu L. YangNR198110NG244261CUHAM708 (T)IndiaDutta et al., 2021 [49]
L. brunneosporus B.E. Lechner & J.M. SuárezMT796198MT796197LB3ArgentinaSuarez et al., 2021 [50]
L. brunneus (Zia Ullah, Jabeen & Khalid) M. Asif, Saba & VellingaMH990662/LAH35862PakistanUllah et al., 2019 [51]
L. bulbiger (Justo, Bizzi & Angelini) M. Asif, Saba & VellingaMN483028/ANGE197Dominican RepublicJusto et al., 2021 [43]
L. caeruleovertens (Justo, Bizzi & Angelini) M. Asif, Saba & VellingaMN483032/ANGE734Dominican RepublicJusto et al., 2021 [43]
L. candidus (Y.R. Ma, Z.W. Ge & T.Z. Liu) M. Asif & SabaOM976877OM976864CFSZ20964ChinaMa et al., 2022 [8]
L. centricastaneus (Y.R. Ma, Z.W. Ge & T.Z. Liu) M. Asif, Saba & VellingaOM976855OM976871SYAUFUNGI076ChinaMa et al., 2022 [8]
L. cepistipes (Sowerby) Pat.PV470803PV476031xml2014128ChinaLi et al., 2025 [52]
L. cinerascens (Quél.) Locq.PX527069PX527070FJAU77966ChinaNCBI Database
L. croceovelutinus Bon & BoiffardEU166351/ecv3131UCUSAVellinga and Sundberg 2008 [53]
L. crystallifer (Vellinga) Migl. & DonatoAF482863AY1764123IX1998GermanyVellinga et al., 2003 [48]
L. cygneus (J.E. Lange) BonKR673661/KA130934Republic of KoreaKim et al., 2015 [54]
L. dacrytus (Vellinga) Kun L. Yang, Jia Y. Lin & Zhu L. YangMT196954/TENN:074972USASwenie and Matheny 2023 [55]
L. dahranwalanus Asif, Saba & RazaOQ947827OQ947833AsifSP71AS67 (T)PakistanAsif et al., 2024 [9]
L. domingensis Justo, Bizzi, Angelini &VizziniMN483016/ANGE418 (T)Dominican RepublicJusto et al., 2021 [43]
L. flammeotinctus (Kauffman) RedheadGU136165/ecv3315UCUSAVellinga 2010 [56]
L. flavescens (Morgan) H.V. Sm.MW567852/SDRussellMycoMa-p1216USANCBI Database
L. flavirobustus R.L. Zhao & J.X. LiPV470825PV475985ZRL20210081 (T)ChinaLi et al., 2025 [52]
L. flavovirens (Jun F. Liang, Zhu L. Yang & J. Xu) Kun L. Yang, Jia Y. Lin & Zhu L. YangEU416293EU416294HKAS29580ChinaLiang et al., 2010 [57]
L. fragilissimus (Ravenel ex Berk. & M.A. Curtis) Pat.LT716029KY418844ZRL20151466ChinaZhao et al., 2017 [58]
L. fuligineopunctatus Justo, Bizzi & AngeliniNR173872/JBSD130948 (T)Dominican RepublicJusto et al., 2021 [43]
L. glaber R.L. Zhao & J.X. LiPV470824PV475895ZRL20210059 (T)ChinaLi et al., 2025 [52]
L. glareicolor (S. Ashraf, Naseer & Khalid) M. Asif, Saba & VellingaOP605604OP782028LAH37455PakistanAshraf et al., 2023 [59]
L. griseodiscus (Bon) Migl. & DonatoGQ329059/MCVE13719ItalyOsmundson et al., 2013 [60]
L. griseofloccosus Lagardère & Eyssart.MH257568/GE17001FranceLagardère and Eyssartier 2016 [61]
L. griseosquamosus (Sysouph. & Thongkl.) Kun L. Yang, Jia Y. Lin & Zhu L. YangPQ871415/LAH38519PakistanNCBI Database
L. guatopoensis (Dennis) M. Asif, Saba & VellingaMN483031/ANGE419Dominican RepublicJusto et al., 2021 [43]
L. gujratensis (A. Rehman, Usman, Afshan & Khalid) M. Asif, Saba & VellingaOP526420OP885328LAH37457PakistanRehman et al., 2023 [62]
L. heinemannii Migl.MN483010/AJ487USAJusto et al., 2021 [43]
L. houaynhangensis (Sysouph.) Kun L. Yang, Jia Y. Lin & Zhu L. YangKX640915/HNL502947LaosSysouphan-thong et al., 2018 [63]
L. ianthinus (Sacc.) P. MohrPQ796880/NFSG2024092201-ACUnited KingdomNCBI Database
L. inflatus Raithelh.MK685764/TRS12100701BrazilSolomon et al., 2019 [64]
L. karjaticus (P.B. Patil, N.P. Patil, S. Chahar & S. Maurya) R.L. Zhao & J.X. LiNR198656NG243935AMH10515 (T)IndiaPatil et al., 2024 [65]
L. lahorensiformis (S. Hussain, H. Ahmad, Afshan & Khalid) M. Asif, Saba & VellingaKU647730KU900516FHSHL2PakistanHussain et al., 2018 [46]
L. lahorensis (Qasim, T. Amir & Nawaz) M. Asif, Saba & VellingaKJ701796/LAH2481336PakistanQasim et al., 2015 [66]
L. lateritiopurpureus (Lj.N. Vassiljeva) M. Asif, Saba & VellingaJX133174/VLAM4729RussiaMalysheva et al., 2013 [67]
L. leucothites (Vittad.) RedheadPV470789PV476017ZRL20235205ChinaLi et al., 2025 [52]
L. lidensis (Migl. & P. Alvarado) Migl. & DonatoMT416130/MVLeu011ItalyMigliozzi and Alvarado 2021 [68]
L. littoralis (Ménier) M. Asif, Saba & VellingaGQ329041/MCVE702ItalyOsmundson et al., 2013 [60]
L. longistriatus (Peck) H.V. Sm. & N.S. WeberMH211956/FLASF61532USANCBI Database
L. lugdunensis S. Basso & N. SchwabPX527021PX527063FJAU77962ChinaNCBI Database
L. margaritifer (Justo, Bizzi & Angelini) M. Asif, Saba & VellingaMN482997/ANGE509Dominican RepublicJusto et al., 2021 [43]
L. marriageae (D.A. Reid) Migl. & DonatoPV026219/MV040040ItalyMigliozzi and Donato 2025 [69]
L. medioflavoides (Bon) Migl. & DonatoGQ329055/MCVE2324ItalyOsmundson et al., 2013 [60]
L. medioflavus (Boud.) BonPQ570805/MRG20231002FranceNCBI Database
L. melanotrichus (Malençon & Bertault) Migl. & DonatoAY176417/ecv2262NetherlandsVellinga 2004 [5]
L. menieri (Sacc.) Migl. & DonatoKP300879/HAHuijserNetherlandsGe et al., 2015 [44]
L. microlepis Justo, Bizzi & AngeliniNR173874/JBSD139951 (T)Dominican RepublicJusto et al., 2021 [43]
L. nigrosquamosus R.L. Zhao & J.X. LiPV470838PV476015ZRL20231501ChinaLi et al., 2025 [52]
L. nivalis (W.F. Chiu) M. Asif, Saba & VellingaMK106151/CM229PakistanJabeen et al., 2020 [70]
L. nympharum (Kalchbr.) M. Asif, Saba & VellingaOM974312OM967233WGS1146ChinaMa et al., 2022 [8]
L. orientiflavus (Z.W. Ge) M. Asif, Saba & VellingaGU084262JN940290HKAS54260ChinaGe 2010 [71]
L. pabbiensis (Usman & Khalid) M. Asif, Saba & VellingaMG973423/LAH35302 (T)PakistanUsman and Khalid 2018 [72]
L. pakistaniensis (Jabeen & Khalid) Asif & SabaKU647727KU900515SJF13PakistanHussain et al., 2018 [46]
L. parvipileus Justo, Bizzi, Angelini & VizziniNR200351/JBSD130965 (T)Dominican RepublicJusto et al., 2021 [43]
L. pegleri (Justo, Bizzi & Angelini) M. Asif, Saba & VellingaNR173878/JBSD130930 (T)Dominican RepublicJusto et al., 2021 [43]
L. phantasmaticus Kun L. Yang, Jia Y. Lin & Zhu L. YangPV620895PV616973HTBM1745ChinaYang et al., 2025 [73]
L. proximus (E.F. Malysheva, Svetash. & E.M. Bulakh) M. Asif, Saba & VellingaJX133171/LE262855RussiaMalysheva et al., 2013 [67]
L. purpureolilacinus (Huijsman) M. Asif, Saba & VellingaGQ329053/MCVE2261ItalyOsmundson et al., 2013 [60]
L. rhodelephantinus (Boisselet & Eyssart.) Migl. & DonatoMT984270/GE19050 (T)FranceBoisselet and Eyssartier 2020 [74]
L. roseovertens (Justo, Bizzi & Angelini) M. Asif, Saba & VellingaNR173879/JBSD130932 (T)Dominican RepublicJusto et al., 2021 [43]
L. rubrobrunneus (E.F. Malysheva, Svetash. & E.M. Bulakh) M. Asif, Saba & VellingaJX133168/LE262863RussiaMalysheva et al., 2013 [67]
L. rubroconfusus (Migl. & Coccia) RedheadKP300875/ZT13003ZTSwitzerlandGe et al., 2015 [44]
L. rubrotinctus (Peck) RedheadJN944081JN940295KUNHKAS54240ChinaNCBI Database
L. sabinae (Angelini, Justo & Vizzini) Kun L. Yang, Jia Y. Lin & Zhu L. YangKM983667KM983669ANGE306Dominican RepublicJusto et al., 2015 [75]
L. sardous (Zecchin & Migl.) Migl. & DonatoKU041693/MCVE20105ItalyMuñoz et al., 2015 [7]
L. scissus Justo, Bizzi & AngeliniNR173873/JBSD139953 (T)Dominican RepublicJusto et al., 2021 [43]
L. serenus (Fr.) M. Asif, Saba & VellingaAF482871AY176421ecv1930LBelgiumVellinga et al., 2003 [48]
L. shenyangensis Z.Q. You & H.S. YuanPV460238PZ235509Yuan20026 (T)ChinaPresent study
L. shenyangensisPV460239PZ235508Yuan19598ChinaPresent study
L. silvestris (Justo, Bizzi & Angelini) M. Asif, Saba & VellingaNR173876/JBSD130934 (T)Dominican RepublicJusto et al., 2021 [43]
L. stillatus (Justo, Bizzi & Angelini) M. Asif, Saba & VellingaNR173877/JBSD130937 (T)Dominican RepublicJusto et al., 2021 [43]
L. straminellus (Bagl.) Narducci & CarotiPX527025PX527066FJAU78063ChinaNCBI Database
L. subcrystallifer (Z.W. Ge & Zhu L. Yang) M. Asif, Saba & VellingaKP205399/ZWGe796ChinaGe et al., 2015 [44]
L. subhymenoderma (Bon & A. Caball.) Kun L. Yang, Jia Y. Lin & Zhu L. YangKT992146/AC2638 (T)SpainNCBI Database
L. sublittoralis (Kühner ex Hora) Migl. & DonatoAY176442AY176443Vellinga2235LNetherlandsVellinga 2004 [5]
L. subpurpureolilacinus (Z.W. Ge & Zhu L. Yang) M. Asif, Saba & VellingaKP096233/ZWGe406ChinaGe et al., 2015 [44]
L. subvolvatus (Malençon & Bertault) M. Asif, Saba & VellingaKP300878/AMBrandsnHunga-ryLHungaryGe et al., 2015 [44]
L. sultanii (S. Hussain, H. Ahmad & Khalid) M. Asif, Saba & VellingaKU647732KU900519FHSH115PakistanHussain et al., 2018 [46]
L. tangerinus (Y. Yuan & Jun F. Liang) Kun L. Yang, Jia Y. Lin & Zhu L. YangPQ321878PQ319804HTBM0679ChinaYang et al., 2024 [10]
L. taniae (C. Heisecke & M.A. Neves) Kun L. Yang, Jia Y. Lin & Zhu L. YangMT952879/MAN1206 (T)BrazilHeisecke et al., 2022 [76]
L. tener (P.D. Orton) M. Asif, Saba & VellingaGQ329043/MCVE751ItalyOsmundson et al., 2013 [60]
L. tephrolepis Justo, Bizzi, Angelini & VizziniNR200350/JBSD130966 (T)Dominican RepublicJusto et al., 2021 [43]
L. thallensis Z. Khan, Izhar & KhalidOP972578OQ568217LAH37619PakistanKhan et al., 2023 [77]
L. truncatus (Z.W. Ge & Zhu L. Yang) M. Asif, Saba & VellingaNR155319/HKAS49288ChinaGe et al., 2015 [44]
L. turgipes (Justo, Bizzi & Angelini) M. Asif, Saba & VellingaNR173880/JBSD130940 (T)Dominican RepublicJusto et al., 2021 [43]
L. umbonatus (S. Hussain, H. Ahmad & Afshan) M. Asif, Saba & VellingaKU647737KU900521LAHSHL1PakistanHussain et al., 2018 [46]
L. vassiljevae (E.F. Malysheva, Svetash. & E.M. Bulakh) M. Asif, Saba & VellingaJX133170/LE289338RussiaMalysheva et al., 2013 [67]
L. viridariorum (G. Muñoz, A. Caball., Salom & Vizzini) Kun L. Yang, Jia Y. Lin & Zhu L. YangKU041692/JCS296LSpainMuñoz et al., 2015 [7]
L. viridiflavus (Petch) E. Ludw.GU574745/CALI10XI2004ChinaLiang et al., 2010 [57]
L. viscidulus (Heinem.) M. Asif, Saba & VellingaPP756657/ANKAcar1305TurkeyNCBI Database
L. volvatus (Bon & A. Caball.) Migl. & DonatoPV026213/MV161494ItalyMigliozzi and Donato 2025 [78]
L. wichanskyi (Pilát) Migl. & DonatoAF482874/IXX1987The NetherlandsVellinga et al., 2003 [48]
Rhizocybe alba Y.X. Ding & E.J. TianOP626999OP646425KUN-HKAS123143ChinaHe et al., 2023 [79]
R. vermicularis (Fr.) Vizzini, P. Alvarado, G. Moreno &ConsiglioKJ681034KJ681040AH44080SpainAlvarado et al., 2015 [80]
Tephrocybe ambusta (Fr.) DonkAF357058/CBS450.87FranceHofstetter et al., 2002 [81]
T. ambustaAF357057/CBS452.87SwitzerlandHofstetter et al., 2002 [81]
T. anthracophila (Lasch) P.D. OrtonOM905952/BR5020155848666Ukrainevan de Peppel et al., 2022 [20]
T. anthracophilaOM905953/K(M):164958Englandvan de Peppel et al., 2022 [20]
T. atrata (Fr.) DonkKP192645/BBF:GC98102503FranceBellanger et al., 2015 [82]
T. confusa (P.D. Orton) P.D. OrtonKP192548/PAM00100616FranceBellanger et al., 2015 [82]
T. confusaKP192611/AB92-11-424FranceBellanger et al., 2015 [82]
T. coracina (Fr.) M.M. MoserOM905954/K(M):195497Englandvan de Peppel et al., 2022 [20]
T. coracinaKP192632/BBF:GC08101102FranceBellanger et al., 2015 [82]
T. erosa (Fr.) BonKP192541/PAM01101601FranceBellanger et al., 2015 [82]
T. erosaKP192634/GC97062013FranceBellanger et al., 2015 [82]
T. fibrosipes Métrod ex BonKP192576/AB02-10-175FranceBellanger et al., 2015 [82]
T. fuscipes P.D. OrtonOM905955/K(M):90837Englandvan de Peppel et al., 2022 [20]
T. impexa (P. Karst.) M.M. MoserOM905956/K(M):191686Englandvan de Peppel et al., 2022 [20]
T. mephitica (Fr.) M.M. MoserOM905958/K(M):196494Englandvan de Peppel et al., 2022 [20]
T. ochraceobrunnea (Métrod ex Consiglio & Contu) P.-A. Moreau & Courtec.KP192549/LIP:PAM02090608FranceBellanger et al., 2015 [82]
T. platypus (Kühner) M.M. MoserOM905961/BR5020153733858Belgiumvan de Peppel et al., 2022 [20]
T. rancida (Fr.) DonkOM905966OM906004CORT012400USAvan de Peppel et al., 2022 [20]
T. rancidaOM905967OM906005TR2017The Netherlandsvan de Peppel et al., 2022 [20]
T. striaepilea (Fr.) DonkKP192647/BBF:GC98101106FranceBellanger et al., 2015 [82]
T. substriipilea Contu & VizziniPX632623/GE22008FranceEyssartier 2025 [83]
T. umbonata Z.Q. You & H.S. YuanPZ235487PZ235512Yuan21497 (T)ChinaPresent study
T. umbonataPZ235488PZ235513Yuan21507aChinaPresent study
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MDPI and ACS Style

You, Z.-Q.; Zhou, L.-J.; Yuan, H.-S.; Lee, H.B. Morphological and Molecular Identification of Three New Macrofungal Species from Shenyang and Adjacent Areas, Northeast China. J. Fungi 2026, 12, 491. https://doi.org/10.3390/jof12070491

AMA Style

You Z-Q, Zhou L-J, Yuan H-S, Lee HB. Morphological and Molecular Identification of Three New Macrofungal Species from Shenyang and Adjacent Areas, Northeast China. Journal of Fungi. 2026; 12(7):491. https://doi.org/10.3390/jof12070491

Chicago/Turabian Style

You, Zi-Qi, Lin-Jiang Zhou, Hai-Sheng Yuan, and Hyang Burm Lee. 2026. "Morphological and Molecular Identification of Three New Macrofungal Species from Shenyang and Adjacent Areas, Northeast China" Journal of Fungi 12, no. 7: 491. https://doi.org/10.3390/jof12070491

APA Style

You, Z.-Q., Zhou, L.-J., Yuan, H.-S., & Lee, H. B. (2026). Morphological and Molecular Identification of Three New Macrofungal Species from Shenyang and Adjacent Areas, Northeast China. Journal of Fungi, 12(7), 491. https://doi.org/10.3390/jof12070491

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