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Article

A Taxonomic Study of Candolleomyces Specimens from China Revealed Seven New Species

1
School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand
2
Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand
3
State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China
4
Tibet Plateau Institute of Biology, Lhasa 850000, China
5
Shennongjia National Park Administration, Shennongjia 442421, China
6
Hubei Provincial Key Laboratory on Conservation Biology of the Shennongjia Golden Snub-Nosed Monkey, Shennongjia 442421, China
7
Institue of Ecology, Chinese Research Academy of Environmental Sciences, Beijing 100012, China
8
State Key Laboratory for Managing Biotic and Chemical Treats to the Quality and Safety of Agro-Products, Institute of Plant Protection and Microbiology, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, China
9
Institute of Digital Agriculture, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, China
10
College of Life Sciences, University of Chinese Academy of Sciences, Beijing 100049, China
*
Authors to whom correspondence should be addressed.
J. Fungi 2024, 10(7), 499; https://doi.org/10.3390/jof10070499
Submission received: 31 May 2024 / Revised: 4 July 2024 / Accepted: 16 July 2024 / Published: 19 July 2024
(This article belongs to the Special Issue Diversity, Phylogeny and Ecology of Forest Fungi)

Abstract

:
Based on phylogenetic analysis, Candolleomyces (Psathyrellaceae, Agaricales) was established with Psathyrella candolleana as the type species. The basidiomes range from small to large and are typically terrestrial, lignicolous, and rarely fimicolous. We analysed the Candolleomyces species collected during five years in China, and based on morphological and molecular data (nrITS, nrLSU, and tef-1α), we propose seven new Candolleomyces species viz. C. brevisporus, C. gyirongicus, C. lignicola, C. luridus, C. shennongdingicus, C. shennongjianus, and C. sichuanicus. Full descriptions, colour photographs, illustrations, phylogenetic analyses results, and comparisons with related Candolleomyces species of the new taxa are provided. This study enriches the species diversity of Candolleomyces in China.

1. Introduction

The recently established genus Candolleomyces D. Wächt. & A. Melzer is characterised by small to large basidiomes, being terrestrial, lignicolous, and rarely fimicolous, the veils are often fibrillose, scaly, or granulose, but very fugacious, stipes occasionally with an annulus, basidiospores are medium-sized, it is pale to medium-dark in colour, with a central, usually invisible, germ pore, the presence of cheilocystidia, and absence of pleurocystidia. Most of the species of Candolleomyces historically belonged to Psathyrella (Fr.) Quél. [1,2]. Smith (1972) and Kits van Waveren (1985) made remarkable works on Psathyrella from Europe and North America, and clarified the morphological species concept within this genus [3,4]. The former identified 11 subgenera viz. Candolleana, Conocybella, Cystopsathyra, Homophron, Lacrymaria, Mycophila, Panaeolina, Pannucia, Psathyrella, Psathyroides, and Pseudostropharia [3,4]. Meanwhile, the latter put Psathyrella into two subgenera, Psathyra and Psathyrella [3,4]. Molecular phylogeny based on the nuclear ribosomal internal transcribed spacer region (nrITS) and the nuclear ribosomal large subunit ribosomal RNA gene (nrLSU) did not distinguish the species well within Psathyrella [5]. Therefore, two protein-coding genes viz. the translation elongator factor alpha (tef-1α) and beta-tubulin (β-tub) were also employed for species classification of Psathyrella [6,7,8]. Based on numerous specimen studies, and morphological and phylogenetic analyses, Wächter and Melzer [9] revised the Psathyrellaceae and divided Psathyrella into 18 subclades, with one of them established as the new genus Candolleomyces and C. candolleanus (Fr.) D. Wächt. & A. Melzer as the species type.
In the field, the members of c are similar to those species of Psathyrella, as they share many similar macro-morphological features and ecological niches. However, Candolleomyces species can be distinguished micro-morphologically by the absence of pleurocystidia, and slightly thicker-walled sphaerocysts can be observed in the veil [9]. Therefore, accurate identification requires proper examination of micromorphological characterisation and molecular phylogenetic analyses. Up to now, 43 species of Candolleomyces were reported worldwide [9,10,11,12,13,14,15,16,17,18,19,20]. Of these, 15 species were reported in China [11,12,19,21].
Most species of this genus are terrestrial, lignicolous, and rarely fimicolous, and can grow on rotten wood, plant debris, or litter [9,14]. Recently two species (C. brunneovagabundus and C. albovagabundus) were described to be in marine habitats [19]. Candolleomyces species, recognised as specialised saprophytes [9,14], were reported to be in temperate, tropical, and subtropical regions across Africa, Asia, Europe, North America, and South America [11,12,13,14,17,22]. In addition, some species of Candolleomyces were reported to have edible and medicinal values, while a few species were found to be poisonous fungi [23,24,25]. The type species C. candolleanus was reported to have medicinal value, although it may cause gastroenteritis and neurotoxicity [23,24,25]. Candolleomyces tuberculatus was reported to have edible value [23,24,25]. Based on poisoning incidents, C. yanshanensis, which was previously uncertain of its edibility, was found to be poisonous and can cause psycho-neurological disorders [26].
In the present study, we collected 31 specimens from Beijing City, Guizhou, Hubei, Sichuan, and Yunnan provinces, and the Xizang Autonomous Region in China, and conducted comprehensive phylogenetic analyses using the nrITS, nrLSU, and tef-1α gene regions. Based on the phylogenetic and morphological analyses, seven new species from China are proposed.

2. Materials and Methods

2.1. Morphological Characteristic Examination

Fresh specimens were collected and photographed (Canon EOS 80D, Tokyo, Japan) in the field from Beijing City, Yunnan, Sichuan, Guizhou, and Hubei provinces, and the Xizang Autonomous Region in China from 2019 to 2023. To avoid mixing or crushing, specimens were packed separately in aluminium foils. Macroscopic characteristics were recorded when fresh, including features of the pileus, veil, context, lamellae, stipe, odour, and chemical reactions. The samples were completely dried with a food drier at 50 °C, sealed in plastic bags, and deposited in the Mycological Herbarium, Institute of Microbiology, Chinese Academy of Sciences (HMAS).
Microscopic characteristics, such as basidia, basidiospores, pileipellis, and cheilocystidia, were observed under an Olympus CX31 light microscope (Olympus, Tokyo, Japan), and at least 30 measurements were made for each character. The description of morphological characteristics followed the protocols of Largent [27]. Additionally, 5% KOH and sterilised water were used for microscopic characterisation. Measured values are given as (a)b–c(d), in which a is the lowest value, b–c includes at least 90% of the values, and d is the highest value. The Q value is the ratio of the length and width of a spore [11]. The colour designation refers to the Methuen Handbook of Colour.

2.2. DNA Extraction, PCR and Sequencing

DNA was extracted from dried specimens using a Broad-spectrum Plant Rapid Genomic DNA Kit (Biomed, Shiyan, China). Primers ITS1 and ITS4 were used for the nuclear internal transcribed spacer (nrITS) of the rDNA region [28], LR7/LR0R were used to amplify the large subunit nuclear ribosomal DNA (nrLSU) region [29], and EF983F/EF2218R were used to amplify the translation elongation factor subunit 1 alpha (tef-1α) region [30]. PCR was performed in 25 µL reactions consisting of 2 µL genomic DNA, 1 µL of each forward and reverse primers, 9 µL ddH2O, and 12 µL 2 × Es Taq MasterMix (Beijing Cowin Biotech Co., Ltd., Beijing, China). The PCR programmes follow Zhao et al. [31] and Bau and Yan [11]. The PCR products were detected by electrophoresis and sent to BGI Genomics Co., Ltd., Shenzhen, China, for purification and sequencing.

2.3. Phylogenetic Analyses

Considering the results of BLAST searches against GenBank and previous studies, we analysed the nrITS, nrLSU and tef-1α sequences of 94 taxa. The details are presented in Table 1. The sequences were aligned by Muscle version 3.6 separately [32], then manually adjusted in BioEdit version 7.0.4 to remove the ambiguous areas [33], and assembled in PhyloSuite version 1.2.3 [34]. The final alignments were deposited in TreeBASE (study no. 31401). Maximum likelihood (ML) analysis of concatenated sequences was carried out using raxmlGUI 1.3 with a GTRGAMMA model and one thousand rapid bootstrap (BS) replicates [35]. The best partitioning scheme and evolutionary models for three pre-defined partitions were selected using PartitionFinder2 v2.1.1 [36], with greedy algorithm and AICc criterion: GTR+I+G for nrLSU, GTR+G for nrITS, and GTR+I+G for tef-1α. Bayesian Inference (BI) analysis was performed using MrBayes v3.2.7a [37]. Six Markov chains were run for two million generations, and trees were sampled every 100th generation. Burn-ins were determined in Tracer version 1.6 with an ESS value higher than 200, and the remaining trees were used to calculate Bayesian posterior probabilities (PP). The trees were displayed in FigTree version 1.4.0 [38].

3. Results

3.1. Phylogeny

Eighty-nine specimens from 43 Candolleomyces species were included in the phylogenetic analyses with Hausknechtia floriformis (Hauskn.) D. Wächt. & A. Melzer and H. leucosticta (Pat.) Tkalčec, J.Q. Yan, C.F. Nie & C.K. Pradeep as outgroups. In total, 84 new sequences were generated in this study, which were from 28 specimens from China, all with the nrITS, nrLSU, and tef-1α sequences. The combined dataset with 3033 characters including gaps (679 for nrITS, 1322 for nrLSU, and 1032 for tef-1α) was included in the phylogenetic analyses. The phylogenetic tree of ML and MrBayes were almost identical. The ML tree is shown in Figure 1 with bootstrap values and Bayesian posterior probabilities indicated on the branches.

3.2. Taxonomy

Candolleomyces brevisporus R.L. Zhao, B. Cao & X.X. Han, sp. nov., Figure 2.
Fungal Names: FN571747.
Holotype: CHINA. Guizhou Province, Doupeng Mountain, 26°37′41″ N, 107°36′54″ E, 1057.8 m asl, 25 September 2021, Yang Liu and Chen-Hao Li, ZRL20211844 (holotype HMAS 258919). GenBank: OR822167 (nrITS), OR822149 (nrLSU), OR819986 (tef-1α).
Etymology: ‘brevisporus’ (Latin) referring to the shorter spores, a distinguishing characteristic of the species.
Diagnosis: Candolleomyces brevisporus, is distinguishable by its pileus, not hygrophanous. Basidiospores (5.0)5.7–6.7(7.3) × (3.3)3.7–4.2(4.5) μm, germ pores are distinct. Pileipellis is a two to three-layered irregular epithelium composed of subglobose cells. Cheilocystidia claviform to utriform, rarely pyriform.
Pileus is 12–33 mm diam, plano-convex to nearly plane, becoming slightly concave at maturity, not hygrophanous, yellowish grey (4B2) to grey (5B1), darker in the centre, occasionally with yellowish grey (5F6) veil elements, becoming white (3A1) as pileus dries. Veil is yellowish grey (5F6), dispersed, fibrillose, falling off easily. Context is thin and very fragile, the same colour as the pileus. Lamellae is moderately close, adnate to slightly adnexed, grey (5B1) to brownish orange (5C3), and edge white as basidiospores mature. Stipes are 19–37 × 1–3 mm, smooth, cylindrical, hollow, equal, and slightly yellowish white (3A2) at the apex. Odour is indistinct.
Basidiospores are (5.0)5.7–6.7(7.3) × (3.3)3.7–4.2(4.5) μm, Q = 1.4–1.7, ellipsoid to oblong-ellipsoid, brown to dark brown in 5% KOH, smooth, and germ pores are distinct. Basidia 13.0–16.4 × 6.7–8.0 μm, clavate, hyaline, and four or two-spored. Pileipellis is a two to three-layered irregular epithelium composed of subglobose cells, oval, (29.2)30.0–42.0(54.0) μm broad, and hyaline. Cheilocystidia is (17.8)20.9–27.7(31.5) × (11.6)12.3–14.9(16.0) μm, claviform to utriform, and rarely pyriform. Trama of gills is irregular. Pleurocystidia is absent.
Habit and habitat: solitary, in pairs, or scattered on the ground with rich humus in broad-leaved forests or broad-leaved shrubs. So far only found in China.
Other specimens examined: CHINA. Guizhou Province, Doupeng Mountain, 26°37′41″ N, 107°36′54″ E, 1057.8 m asl, 25 September 2021, Yang Liu and Chen-Hao Li, ZRL20211843 (HMAS 258920).
Notes: Candolleomyces brevisporus is reminiscent of C. subcacao T. Bau & J.Q. Yan with its dirty white pileus and pale brown veil. Both were originally described in China, but C. subcacao differs from C. brevisporus by having larger basidiospores (6.8–8.0 × 3.9–4.9 μm), and longer basidia (17–22 × 6.1–7.3 μm) [11]. In the multigene tree (Figure 1), Candolleomyces brevisporus formed a monophyletic sister clade to C. lignicola with high support, but the former pileus is not hygrophanous, pale grey to greyish brown, and the two have different nrITS and tef-1α sequences (Figure 3).
Candolleomyces gyirongicus R.L. Zhao, B. Cao & X.X. Han, sp. nov., Figure 4.
Fungal Names: FN 571921.
Holotype: CHINA. Xizang Autonomous Region, Shigatse Municipality, Gyirong County, Gyironggou, 28°24′ N, 85°18′ E, 2935 m asl, 1 August 2022, Mao-Qiang He, Bin Cao, ZRL20220470 (holotype HMAS 287612). GenBank: PP734613 (nrITS), PP734624 (nrLSU), PP729326 (tef-1α).
Etymology: refers to Gyirong County, the locality of the type specimen.
Diagnosis: Candolleomyces gyirongicus, is distinguishable by its pileus, slightly hygrophanous. Basidiospores (5.5)6.1–6.9(8.0) × (3.2)3.8–4.3(4.7) μm, often with germ pores. Pileipellis is a one to two-layered irregular epithelium composed of subglobose cells. Cheilocystidia utriform, sometimes claviform.
Pileus is 15–56 mm diam, paraboloid to hemispherical when young, broadly conical, convex to broadly convex, becoming plano-convex to nearly plane when mature, slightly hygrophanous, with grey (5B1) veil elements at a young stage, white (5A1), orange-white (5A2) to golden brown (5D7), paler at the margin, and usually white (5A1) to orange-white (5A2). Veil is grey (5B1), fibrillose, and evanescent. Context is 0.2–0.5 mm broad at the centre, same color as pileus, and fragile. Lamellae is very close to moderately close, adnate to adnexed, orange-white (5A2) to pale orange (5A3) when immature, becoming greyish orange (5B5), and nougat (5D3) to greyish brown (6D3) when mature. Stipes are 30–100 × 3–6 mm, smooth, hollow, with white (5A1) fibrils at the base, and white (5A1) to yellowish white (3A2). Odour is not distinctive. Taste is indistinct.
Basidiospores are (5.5)6.1–6.9(8.0) × (3.2)3.8–4.3(4.7) μm, Q = 1.5–1.7, ellipsoid to oblong-ellipsoid, pale brown to dark brown in 5% KOH, smooth, often with germ pore. Basidia is 15.5–19.7 × 7.2–8.5 μm, clavate, hyaline, and four-spored. Pileipellis is a one to two-layered irregular epithelium composed of subglobose cells, (19.9)24.8–31.5(39.5) μm broad, and hyaline. Cheilocystidia is (25.7)37.7–53.1(61.1) × (7.1)9.1–12.7(15.1) μm, utriform, and sometimes claviform. Trama of gills is irregular. Pleurocystidia is absent.
Habit and habitat: solitary, scattered on soil, in bush, and broad-leaved or deciduous coniferous forest. So far only found in China.
Other specimens examined: CHINA. Yunnan Province, Jingdong County, Ailao Mountain, 24°52′ N, 101°03′ E, 2443 m asl, 4 July 2021, Rui-Lin Zhao, Mao-Qiang He, Xin-Yu Zhu, Ming-Zhe Zhang, ZRL20210352 (HMAS 287607); Xizang Autonomous Region, Nyingchi Municipality, Zayü County, 28°36′ N, 98°05′ E, 4110 m asl, 21 July 2021, Rui-Lin Zhao, Ming-Yu Zhu, Bin Cao, ZRL20210621 (HMAS 287608); Xizang Autonomous Region, Nyingchi Municipality, Mêdog County, Xironggou, 29°42′ N, 95°35′ E, 2800 m asl, 25 July 2021, Bin Cao, Xin-Yu Zhu, Ming-Zhe Zhang, ZRL20210861 (HMAS 287609); Xizang Autonomous Region, Nyingchi Municipality, Mêdog County, Xironggou, 29°42′N, 95°35′ E, 2800 m asl, 25 July 2021, Zhi-Lin Ling, Mao-Qiang He, ZRL20210966 (HMAS 287610); Xizang Autonomous Region, Shigatse Municipality, Dinggyê County, Chentang Town, Jiuyan hot spring, 27°55′ N, 87°21′ E, 3060 m asl, 29 July 2021, Rui-Lin Zhao, Xin-Yu Zhu, ZRL20220325 (HMAS 287611); Xizang Autonomous Region, Shigatse Municipality, Gyirong County, Gyironggou, 28°26′ N, 85°15′ E, 3024 m asl, 1 August 2022, Dorji Phurbu, Jia-Xin Li, ZRL20220628 (HMAS 287613); and Xizang Autonomous Region, Shigatse Municipality, Gyirong County, Gyironggou, 28°26′ N, 85°15′ E, 3024 m asl, 1 August 2022, Dorji Phurbu, Jia-Xin Li, ZRL20220631 (HMAS 287614).
Notes: In the field, Candolleomyces gyirongicus is morphologically similar to C. candolleanus. However, C. gyirongicus can be distinguished from C. candolleanus by its smaller basidiospores, which measure (5.5)6.1–6.9(8.0) × (3.2)3.8–4.3(4.7) μm, and larger basidia (15.5–19.7 × 7.2–8.5 μm) [48,49]. In the multigene tree (Figure 1), Candolleomyces gyirongicus formed a monophyletic sister clade to C. shennongjianus with high support. However, C. gyirongicus has a narrower context, thinner stipe, as well as smaller basidiospores, slightly bigger basidia, and longer but narrower cheilocystidia. Additionally, there are some differences in the sequence of tef-1α (Figure 3). Candolleomyces gyirongicus is introduced as a new species based on morphology and phylogenetic analyses.
Candolleomyces lignicola R.L. Zhao, B. Cao & X.X. Han, sp. nov., Figure 5.
Fungal Names: FN571749.
Holotype: CHINA. Yunnan Province, Chuxiong, Zixi Mountain, 25°01′06″ N, 101°23′19″ E, 2235 m asl, 18 July 2021, Rui-Lin Zhao, Bin Cao and Xin-Yu Zhu, ZRL20210496 (holotype HMAS 258921). GenBank: OR822169 (nrITS), OR822151 (nrLSU), OR819988 (tef-1α).
Etymology: ‘lignicola’ (Latin) refers to the habitat, this species grows mainly on rotting wood.
Diagnosis: Candolleomyces lignicola differs by its pileus, hygrophanous. Basidiospores (4.5)5.5–6.9(7.6) × (3.5)3.7–4.3(4.8) μm, a germ pore is absent or indistinct. Pileipellis is a two to three-layered irregular epithelium composed of irregular subglobose cells, irregular oval. Cheilocystidia claviform to somewhat broadly claviform or subsphaeropenduculate. Habitat on rotten wood.
Pileus has a 28–53 mm diam, flabellate, flattening with age, with or without obtuse umbo, hygrophanous, dark blonde (5D4) to yellowish brown (5D8) at the centre and golden blonde (5C4) to pale orange (5A3) toward the margin, becoming orange-white (5A2) as pileus dries, with split margins when mature. Veil is white (5A1), fibrillose, and evanescent. Context is thin and very fragile, the same colour as the pileus. Lamellae is close to moderately close, adnate to adnexed, nougat (5D3) to elay (5D5), and edge white (5A1) as basidiospores mature. Stipes are 23–51 × 3–9 mm, cylindrical, hollow, equal, white (5A1) to orange-white (5A2), and has a surface covered with yellowish white (3A2) fibrillose. Odour is not distinctive.
Basidiospores are (4.5)5.5–6.9(7.6) × (3.5)3.7–4.3(4.8) μm, Q = 1.4–1.7, ellipsoid to oblong-ellipsoid, pale brown to brown in 5% KOH, smooth, a germ pore is absent or indistinct. Basidia is 12.1–15.8 × 6.1–7.9 μm, clavate, hyaline, and 4 or 2-spored. Pileipellis is a two to three-layered irregular epithelium composed of irregular subglobose cells, irregular oval, (19.0)24.0–35.8(43.6) μm broad, and hyaline. Cheilocystidia is (18.4)22.1–32.1(40.6) × (7.5)10.7–15.3(18.9) μm, claviform to somewhat broadly claviform or subsphaeropenduculate, and rarely with deposits. Trama of gills is irregular. Pleurocystidia is absent.
Habit and habitat: solitary, in pairs, or scattered on rotten wood in broad-leaved forests or broad-leaved shrubs.
Other specimens examined: CHINA. Yunnan Province, Nangunhe Nature Reserve, 23°22′06″ N, 99°21′22″ E, 1633 m asl, 3 July 2021, Rui-Lin Zhao, Mao-Qiang He, and Ming-Zhe Zhang, ZRL20210404 (HMAS 258922).
Notes: Candolleomyces lignicola can easily be mistaken for C. yanshanensis in the field due to their similar macroscopic characteristics. However, C. yanshanensis differs from C. lignicola due to its slightly larger basidiospores (5.8–8.2 × 3.3–5.4 μm) and longer basidia (17–31 × 5.8–7.5 μm) [12]. Candolleomyces lignicola distinguishes itself from the sister species C. brevisporus by its broader pileus, wider lamellae, and longer and thicker stipes. Additionally, there are differences in their nrITS and tef-1α sequences (Figure 3). Notably, Candolleomyces lignicola was collected on wood rather than soil.
Candolleomyces luridus R.L. Zhao, B. Cao & X.X. Han, sp. nov., Figure 6.
Fungal Names: FN571751.
Holotype: CHINA. Xizang Autonomous Region, Shigatse Municipality, Gyirong County, Gyironggou, 28°14′24″ N, 85°10′48″ E, 2935 m asl, 1 August 2022, Dorji Phurbu and Jia-Xin Li, ZRL20220606 (holotype HMAS 258913). GenBank: OR822161 (nrITS), OR822143 (nrLSU), OR819980 (tef-1α).
Etymology: ‘luridus’ (Latin) refers to the yellowish brown colours of the pileus.
Diagnosis: Candolleomyces luridus is distinguished by its pileus, hygrophanous. Basidiospores are (5.3)6.1–7.1(8.3) × (3.4)3.9–4.6(5.2) μm, germ pores are distinct. Pileipellis a one to two-layered irregular epithelium composed of irregular subglobose cells, that is an irregular oval. Cheilocystidia is narrowly utriform to utriform, sometimes subclaviform.
Pileus is 21–50 mm diam, broadly conical when young and convex when mature, with or without obtuse umbo, hygrophanous, golden yellow (5B7) to yellowish brown (5D8), with orange-white (5A2) veil elements at a young stage, striate up to halfway from the margin or indistinct, sometimes cleft or lobed. Veil is white (5A1), fibrillose, and falls off easily. Context is thin and very fragile, the same colour as the pileus. Lamellae is adnate to adnexed, pale orange (5A3) to elay (5D5), and edge is orange-white (5A2) to white (5A1) as basidiospores mature. Stipes are 39–72 × 3–8 mm, cylindrical, hollow, equal, orange-white (5A2) to pale orange (5A3), with a surface covered with slight white fibrils, and is evanescent. Odour is not distinctive.
Basidiospores are (5.3)6.1–7.1(8.3) × (3.4)3.9–4.6(5.2) μm, Q = 1.4–1.7, ellipsoid to oblong-ellipsoid, pale brown to brown in 5% KOH, smooth, and germ pores are distinct. Basidia is 16.7–19.5 × 7.3–8.6 μm, clavate, hyaline, and four or two-spored. Pileipellis is a one to two-layered irregular epithelium composed of irregular subglobose cells, is an irregular oval, (14.3)22.0–36.0(42.9) μm broad, and hyaline. Cheilocystidia is (20.6)26.0–41.4(53.8) × (7.6)9.0–11.8(14.4) μm, narrowly utriform to utriform, sometimes subclaviform, and rarely with deposits. Trama of gills is irregular. Pleurocystidia is absent.
Habit and habitat: in pairs, scattered or clustered on humus-rich ground or decaying wood in broad-leaved or deciduous coniferous forests.
Other specimens examined: CHINA. Xizang Autonomous Region, Shigatse Municipality, Gyirong County, Gyironggou, 28°14′24″ N, 85°10′48″ E, 2935 m asl, 1 August 2022, Dorji Phurbu and Jia-Xin Li, ZRL20220625 (HMAS 258914) and ZRL20220627 (HMAS 258915); Sichuan Province, Liangshan Yi Autonomous Prefecture, Yanyuan County, Xiamosuogou, 27°39′31″ N, 101°16′6″ E, 1953 m asl, 8 August 2019, Rui-Lin Zhao, Bin Cao, and Zhi-Lin Ling, ZRL20190449 (HMAS 258912); Sichuan Province, Ganzi Tibetan Autonomous Prefecture, Xiangcheng County, Fozhuxia Nature Reserve, 29°3′53″ N, 99°56′16″ E, 3090 m asl, 21 August 2020, Rui-Lin Zhao and Xi-Xi Han, ZRL20201771 (HMAS 258911); Beijing City, Miyun District, Taishitun Town, Bailongtan, 40°29’32″ N, 117°4’0″ E, 302 m asl, 28 August 2023, Bin Cao, Ming-Yu Zhu and Bei Han, ZRL20230723 (HMAS 287929); and Beijing City, Pinggu District, Laoquankou Village, 40°29’32″ N, 117°4’0″ E, 229 m asl, 15 August 2023, Jia-Xin Li, Wen-Qiang Yang and Ze-Zhi Wang, ZRL20233312 (HMAS 287930).
Notes: Candolleomyces luridus is easily confused with C. candolleanus and C. gyirongicus in the field due to its similar macroscopic characteristics, but C. candolleanus differs from C. luridus in having larger basidiospores (7–8 × 4.5–5.5 μm) and smaller basidia (14–17 × 6–7 μm) [48,49], while C. gyirongicus has a longer cheilocystidia. The nrITS and tef-1α sequences of C. luridus are distinct from other members of Candolleomyces (Figure 3). Therefore, we introduce C. luridus as a new species based on morphology and phylogenetic analyses (Figure 1, Figure 2 and Figure 6).
Candolleomyces shennongdingicus R.L. Zhao, B. Cao & X.X. Han, sp. nov., Figure 7.
Fungal Names: FN571750.
Holotype: CHINA. Hubei Province, Shennongjia National Park, Shennongding, Jinhou Ridge, 31°16′48″ N, 110°10′47.9″ E, 2498 m asl, 25 August 2022, Rui-Lin Zhao and Mao-Qiang He, ZRL20220855 (holotype HMAS 258918). GenBank: OR822166 (nrITS), OR822148 (nrLSU), OR819985 (tef-1α).
Etymology: ‘shennongdingicus’ refers to the location Shennongding where the holotype was collected.
Diagnosis: Candolleomyces shennongdingicus is recognised by the pileus, hygrophanous. Basidiospores are (5.3)6.2–7.4(8.5) × (3.4)3.8–4.3(4.7) μm, and germ pores are distinct but small. Pileipellis is a one to two-layered irregular epithelium composed of subglobose cells. Cheilocystidia is narrowly utriform, seldom cylindrical to claviform.
Pileus has a 25–53 mm diam, parabolic when young and convex when mature, with or without obtuse umbo, hygrophanous, is darker in the centre, golden yellow (5B7) to golden brown (5D7) at the centre and golden blonde (5C4) to nougat (5D3) toward the margin, with striate up to halfway from the margin or indistinct. Veil is white (5A1), fibrillose, and gradually disappearing in later stages. Context is grey (5C1), thin, and very fragile. Lamellae is adnate, moderately close, orange-grey (5B2) to grey (5D1), and the edge becomes white as basidiospores mature. Stipes are 34–53 × 3–5 mm, cylindrical, hollow, equal, and white (5A1) to orange-white(5A2), with the same color flocculent fibres. Odour is not distinctive.
Basidiospores are (5.3)6.2–7.4(8.5) × (3.4)3.8–4.3(4.7) μm, Q = 1.5–1.8, ellipsoid to oblong, brown to dark brown in 5% KOH, smooth, the germ pore is distinct but small. Basidia 15.8–21.2 × 6.4–7.9 μm, short clavate, hyaline, 4 or 2-spored. Pileipellis is a one to two-layered irregular epithelium composed of subglobose cells, (14.9)19.8–29.5(37.4) μm broad, hyaline. Cheilocystidia (23.8)29.8–40.5(49.1) × (8.2)9.5–13.3(15.3) μm, narrowly utriform, seldom cylindrical to claviform, thin-walled, rarely with deposits. Trama of gills is irregular. Pleurocystidia is absent.
Habit and habitat: solitary, scattered or clustered on the ground with rich humus in broad-leaved or deciduous coniferous forests.
Other specimens examined: CHINA. Xizang Autonomous Region, Shigatse Municipality, Dinggyê County, Chentang Town, Xiaerba Village, 27°31′12″ N, 87°15′0″ E, 2600 m asl, 30 July 2022, Dorji Phurbu, ZRL20220339 (HMAS 258917); Xizang Autonomous Region, Shigatse Municipality, Dinggyê County, Chentang Town, Jiuyan hot spring, 27°33′0″ N, 87°12′36″ E, 3060 m asl, 29 July 2022, Mao-Qiang He, Bin Cao, Jia-Xin Li, ZRL20220411 (HMAS 258916).
Notes: In the field, Candolleomyces shennongdingicus can be easily confused with C. shennongjianus at first glance, as both species have parabolic pileus when young and convex when mature, and yellowish brown pileus. Additionally, both species exhibit white stipes with pale yellowish brown bases. However, C. shennongdingicus can be distinguished by its slightly smaller basidiospores, longer basidia, as well as fusiform, seldom cylindrical to clavate, and smaller cheilocystidia. Moreover, phylogenetic analysis reveals that C. shennongdingicus is distinct from C. shennongjianus (Figure 3). Based on morphology and phylogenetic analyses, Candolleomyces shennongdingicus is introduced as a new species (Figure 1 and Figure 7).
Candolleomyces shennongjianus R.L. Zhao, B. Cao & X.X. Han, sp. nov., Figure 8.
Fungal Names: FN571748.
Holotype: CHINA. Hubei Province, Shennongjia National Park, Shennongding, Jinhou Ridge, 31°16′48″ N, 110°10′47.9″ E, 2498 m asl, 25 August 2022, Rui-Lin Zhao, Mao-Qiang He, ZRL20220858 (holotype HMAS 258909). GenBank: OR822157 (nrITS), OR822139 (nrLSU), OR819976 (tef-1α).
Etymology: shennongjianus refers to the location Shennongjia National Park, where the type specimen was collected.
Diagnosis: Candolleomyces shennongjianus is distinguishable by its pileus, hygrophanous. Basidiospores are (5.9)6.7–8.4(9.4) × (3.9)4.2–4.9(5.3) μm, the germ pore is distinct but small. Pileipellis is a two to three-layered irregular epithelium composed of irregular subglobose cells, and is a irregular oval. Cheilocystidia is utriform, subclaviform, and sometimes pyriform.
Pileus is 23–63 mm diam, paraboloid when young, obtusely conical, convex, or plano-convex when mature, with or without obtuse umbo, and sometimes cleft or lobed; surface is glabrous, dull, hygrophanous, oak brown (5D6) to bronze (5E5), darker in the centre, and striate up to halfway from the margin or indistinct. Veil is white (5A1), dispersed, fibrillose, and falls off easily. Context is 0.5–1.0 mm broad at the centre, the same colour as pileus. Lamellae is moderately close, adnate to slightly adnexed, grey (5C1), brownish orange (5C3) to hair brown (5E4), and the edge becomes white as basidiospores mature. Stipes are 45–70 × 4–9 mm, sometimes with occasional white flocculation, hollow, white (5A1) to pale orange (5A3), and sometimes dark blond (5D4) at the base. Odour is indistinct.
Basidiospores are (5.9)6.7–8.4(9.4) × (3.9)4.2–4.9(5.3) μm, Q = 1.5–1.8, ellipsoid to oblong, brown (#b06500) to dark brown (#4f484c) in 5% KOH, abundant, smooth, germ pores are distinct but small. Basidia is 13.3–18.3 × 6.6–8.7 μm, clavate, hyaline, and four-spored. Pileipellis is a two to three-layered irregular epithelium composed of irregular subglobose cells, is an irregular oval, (15.7)19.2–28.8(38.0) μm broad, and hyaline. Cheilocystidia is (27.5)35.0–45.3(51.3) × (8.2)11.2–14.7(16.9) μm, utriform, subclaviform, sometimes pyriform, rarely with deposits, and thin-walled. Trama of gills is irregular. Pleurocystidia is absent.
Habit and habitat: Solitary, in pairs, or scattered on the ground with rich humus in broad-leaved or deciduous coniferous forests.
Other specimens examined: CHINA. Hubei Province, Shennongjia National Park, Shennongding, Jinhou Ridge, 31°16′48″ N, 110°10′47.9″ E, 2498 m asl, 25 August 2022, Rui-Lin Zhao and Mao-Qiang He, ZRL20220857 (HMAS 258910); and Hubei Province, Shennongjia National Park, Shennongding, Guanyin Cave, 31°17′24″ N, 110°10′12″ E, 2283 m asl, 2 September 2022, Rui-Lin Zhao, Bin Cao, Xi-Xi Han and Xin-Yu Zhu, ZRL20221427 (HMAS 258907) and ZRL20221467 (HMAS 258908).
Notes: Candolleomyces shennongjianus is morphologically similar to C. asiaticus. However, C. asiaticus can be distinguished by its broader basidiospores (7.2–7.6 × 4.5–6 vs. 6.7–8.4 × 4.2–4.9 μm), larger basidia (19.3–22.5 × 9.4–10.5 vs. 13.3–18.3 × 6.6–8.7 μm), and shorter cheilocystidia (21–38 × 9.6–16 vs. 35.0–45.3 × 11.2–14.7 μm) [14]. In addition, there are differences in their nrITS and tef-1α sequences (Figure 3). Based on morphological characteristics and phylogenetic analyses, C. shennongjianus is introduced as a new species (Figure 1, Figure 3 and Figure 8).
Candolleomyces sichuanicus R.L. Zhao, B. Cao & X.X. Han, sp. nov., Figure 9.
Fungal Names: FN 571922.
Holotype: CHINA. Sichuan Province, Ganzi Tibetan Autonomous Prefecture, Derong County, Xiayong Nature Reserve, 28°22′ N, 99°21′ E, 3399 m asl, 22 August 2020, Bin Cao, Jia-Xin Li, ZRL20201861 (holotype HMAS 287616). GenBank: PP734617 (nrITS), PP734628 (nrLSU), PP729330 (tef-1α).
Etymology: refers to Sichuan Province, the locality of the type specimen.
Diagnosis: Candolleomyces sichuanicus differs from other species by its pileus, not hygrophanous. Basidiospores is (5.6)6.4–7.9(8.6) × (3.4)4.0–4.8(5.1) μm, sometimes germ pores are absent. Pileipellis a two to three-layered irregular epithelium composed of irregular subglobose cells, and is an irregular oval. Cheilocystidia is utriform, rarely subclaviform.
Pileus is 8–42 mm diam, paraboloid to hemispherical when young and convex to plano-convex when mature, sometimes cleft or lobed, moist, smooth, not hygrophanous, not striate to rimos, and golden blonde (5C4) to yellowish brown (5D8). Veil is white (5A1), fibrillose, and evanescent. Context is thin and very fragile, and the same colour as the pileus. Lamellae is adnexed, grey (5B1) to nougat (5D3), and the edge becomes white (5A1) as spores mature. Stipes are 21–68(74) × 3–7 mm, hollow, and white (5A1) to grey (5B1). Odour is not distinctive. Taste is indistinct.
Basidiospores are (5.6)6.4–7.9(8.6) × (3.4)4.0–4.8(5.1) μm, Q = 1.5–1.8, ellipsoid to oblong, pale brown to brown in water, abundant, smooth, and sometimes germ pores are absent. Basidia is 17.4–22.4 × 7.9–8.9 μm, clavate, hyaline, and four-spored. Pileipellis is a two to three-layered irregular epithelium composed of irregular subglobose cells, is an irregular oval, (12.2)17.9–26.3(33.2) μm broad, and hyaline. Cheilocystidia is (27.7)35.0–44.6(55.1) × (9.9)11.7–15.1(17.8) μm, utriform, rarely subclaviform. Trama of gills is irregular. Pleurocystidia is absent.
Habit and habitat: Scattered or clustered on the ground with rich humus in broad-leaved or deciduous coniferous forests. So far only found in China in July/August.
Other specimens examined: CHINA. Sichuan Province, Ganzi Tibetan Autonomous Prefecture, Yajiang County, Gexigou National Nature Reserve, 30°3′ N, 100°56′ E, 2953 m asl, 15 August 2020, Rui-Lin Zhao, Ming-Zhe Zhang, Mei-Qi Wang, ZRL20200271 (HMAS 287615).
Notes: Candolleomyces sichuanicus is morphologically similar to C. cladii-marisci and C. gyirongicus. Candolleomyces cladii-marisci differs from C. sichuanicus by having larger basidiospores (7–9.5 × 4–5.5 vs. 6.4–7.9 × 4.0–4.8 μm), smaller basidia (8.5–20.5 × 6–9 vs. 17.4–22.4 × 7.9–8.9 μm), and narrower cheilocystidia (21.5–54 × 6–11 vs. 35.0–44.6 × 11.7–15.1 μm) [13]. In contrast, Candolleomyces gyirongicus can be distinguished by its longer stipe, smaller basidiospores, shorter basidia, and longer but narrower cheilocystidia. Phylogenetic analysis and morphological characteristics supported the proposal of this new species (Figure 1, Figure 3 and Figure 9).
Key to Candolleomyces species distributed in Chinese
1a Spores very pale, nearly hyaline in 5% KOH2
1b Spores pale yellowish brown, greyish brown or darker8
2a Spores mostly larger than 8.0 μm3
2b Spores less than 8.0 μm4
3a Spores larger than 8.5 μm and mostly wider than 5.0 μmC. luteopallidus
3b Not as aboveC. sulcatotuberculosus
4a Spores less than 7.0 μm5
4b Not as above7
5a Basidiomata slender, spores nearly hyaline in waterC. subminutisporus
5b Basidiomata stout, spores orange-white to pale orange in water6
6a Pileus 5–20 mm, brown to golden brownC. subcandolleanus
6b Pileus 5–25 mm, incanus to nudeC. incanus
7a Basidiomata stout, spores up to 5.5 μm wideC. singeri
7b Basidiomata slender, spores up to 4.5 μm wideC. subsinger
8a Spores larger than 10.0 μmC. typhae
8b Not as above9
9a Spores without a germ pore, sequestrate basidoma and marine habits10
9b Spores with a germ pore11
10a Brownish basidomaC. brunneovagabundus
10b Whitish basidomaC. albovagabundus
11a Sometimes germ pore absent12
11b Germ pore distinct15
12a Pileipellis isa one to two-layered irregular epitheliumC. gyirongicus
12b Pileipellis is a two to three-layered irregular epithelium13
13a Spores up to 5.0 μm wideC. yanshanensis
13b Not as above14
14a Cheilocystidia claviform to somewhat broadly claviform or subsphaeropenduculateC. lignicola
14b Cheilocystidia utriform, rarely subclaviformC. sichuanicus
15a Spores up to 5.0 μm wide, larger than 8.0 μmC. leucotephrus
15b Not as above16
16a Spores less than 4.0 μm wide, pileus 6–22 mmC. albipes
16b Spores 3.7–4.9 μm wide, pileus 10–100 mm17
17a Basidiomata slender, pileus yellowish grey to grey, brown, becoming white as dries18
17b Basidiomata stout, orange-white, golden yellow to yellowish brown19
18a Spores less than 6.8 μmC. brevisporus
18b Not as aboveC. subcacao
19a Spores less than 7.5 μm20
19b Spores up to 7.5 μm, germ pore distinct but small21
20a Spores 6.2–7.4 × 3.8–4.3 μm, germ pore distinct but smallC. shennongdingicus
20b Spores 6.1–7.1 × 3.9–4.6 μm, germ pore distinctC. luridus
21a Pileus 23–63 mm, spores 6.7–8.4 × 4.2–4.9 μmC. shennongjianus
21b Pileus 10–100 mm, spores 6.1–9.0 × 3.7–4.5 μmC. candolleanus

4. Discussion

At present, 15 species in Candolleomyces were reported from China viz. C. albipes [50], C. albovagabundus [19], C. brunneovagabundus [19], C. candolleanus [21], C. incanus [12], C. leucotephrus [21], C. luteopallidus [21], C. singer [21], C. subcacao [11], C. subcandolleanus [12], C. subminutisporus [11], C. subsingeri [21], C. sulcatotuberculosus [11], C. typhae [21], and C. yanshanensis [12]. Yan (2018) reported the distribution of C. leucotephrus, C. singer, and C. subsingeri in China based on morphological characteristics and ITS sequences [21]. Subsequently, in 2021, Bau and Yan supplemented these specimens with LSU, tef-1α, and β-Tub sequences [11]. Additionally, they identified the sample with the voucher of HFJAU1515 as C. sulcatotuberculosus and provided its ITS, tef-1α, and β-Tub sequences but did not describe its morphology [11]. Furthermore, Psathyrella typhae var. bispora was reported as a new variety in China in 2018 [21], and it became synonymised with Candolleomyces typhae [21]. However, only its SSU sequence is available in NCBI, lacking the complete sequence data needed to verify its presence in China through phylogenetic analysis. On the other hand, C. albipes has only morphological descriptions in China, with no associated molecular data, leaving its actual presence in the region uncertain and requiring further investigation [50]. Furthermore, although C. singeri was previously reported in Hubei, this paper marks the first record of the species in the Shennongjia National Park. In the recorded 43 Candolleomyces species, 36 species have nrITS sequences, 27 species have nrLSU sequences, and 16 species have tef-1α sequences. However, only 14 species have nrITS, nrLSU, and tef-1α sequences. All five new species revealed in this study provided sequences of nrITS, nrLSU, and tef-1α.
Significant progress for Candolleomyces was made in the study, but there are still some challenges. Existing classifications primarily rely on a combination of morphological characters and molecular data [18,20]. However, due to the variability of morphological traits and the limited availability of gene sequence data, the identification and classification of certain species remain problematic [9]. In recent years, the development of molecular biology techniques, such as tef-1α and β-tub gene sequence analyses, greatly facilitated systematic taxonomic studies of the genus [11,12,19]. The ongoing discovery of well-defined boundaries in new taxa, as demonstrated by this study, enhances our understanding of species within this genus. In addition, the edible and medicinal values, as well as the toxicity, of only a few species, such as C. candolleanus, C. tuberculatus, and C. yanshanensis, were clarified, while those of most other species remain unknown. Meanwhile, the ecological functions and distribution ranges of the majority of species are still ambiguous. The diversity of the genus Candolleomyces continues to increase with the discovery of new species, necessitating more comprehensive field investigations, as well as morphological and molecular studies to refine the taxonomic system [12]. Future research should focus on integrating morphological, molecular, and ecological methods to further elucidate the phylogenetic relationships and species diversity within the genus.

5. Supplementary Note

In the paper “Zhi-Lin Yuan, Fu-Cheng Lin, Chu-Long Zhang, Christian P. Kubicek, A new species of Harpophora (Magnaporthaceae) recovered from healthy wild rice (Oryza granulata) roots, representing a novel member of a beneficial dark septate endophyte [51], the authors propose corrections for the invalid publication name, which does not conform to Nom. inval., Art. 40.7 (Melbourne). Two strain numbers are provided, where the holotypes lyophilised culture no. R5-6-1 was deposited. These were corrected as follows: China General Microbiological Culture Collection Center (CGMCC 2737) was designated as the holotype, and Centraalbureau voor Schimmelcultures (CBS 125863) was designated as the paratype.
Harpophora oryzae Z.L. Yuan, C.L. Zhang & F.C. Lin
Holotype: CGMCC 2737
Paratype: CBS 125863

Author Contributions

Conceptualisation, X.-X.H.; field sampling, X.-X.H., D.P., G.-F.M., Y.-Z.L., Y.-J.M. and D.-M.L.; molecular experiments and data analysis, X.-X.H. and F.-C.L.; morphological observation, X.-X.H. and F.-C.L.; writing—original draft preparation, X.-X.H.; writing—review and editing, N.T., R.-L.Z. and B.C.; supervision, N.T. and B.C.; project administration, B.C.; funding acquisition, B.C. and R.-L.Z. All authors have read and agreed to the published version of the manuscript.

Funding

This work was supported by the Project of Background Resources Survey in Shennongjia National Park (SNJNP2022004), the Open Project Fund of Hubei Provincial Key Laboratory for Conservation Biology of Shennongjia Snub-nosed Monkeys (SNJGKL2022004), the Project of Science and Technology Programs of Tibet (XZ202202YD0031C), the Survey of Wildlife Resources in Key Areas of Tibet (ZL202203601), the Biodiversity Survey and Assessment Project of the Ministry of Ecology and Environment, China (2019HJ2096001006), and the National Natural Science Foundation of China (31961143010, 31970010, 31470152).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

All sequence data are available in NCBI GenBank following the accession numbers in the manuscript.

Acknowledgments

We thank the Mycological Herbarium, Institute of Microbiology, Chinese Academy of Sciences (HMAS), and all the individuals in our laboratory. We thank Luis A. Parra and Shaun Pennycook (Manaaki Whenua Landcare Research, New Zealand) for his valuable suggestions on the fungal nomenclature. We thank Qianlan Ma (International Department, Beijing BAYI School) for her help in this study.

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. Molecular phylogenetic analyses of Candolleomyces species by the maximum likelihood (ML) method based on combined nrITS, nrLSU, and tef-1α sequences. Maximum likelihood bootstrap support values (ML) ≥ 60% and Bayesian posterior probabilities (PP) ≥ 0.95 are shown at the nodes as ML/PP. Candolleomyces species produced in this study are indicated in bold.
Figure 1. Molecular phylogenetic analyses of Candolleomyces species by the maximum likelihood (ML) method based on combined nrITS, nrLSU, and tef-1α sequences. Maximum likelihood bootstrap support values (ML) ≥ 60% and Bayesian posterior probabilities (PP) ≥ 0.95 are shown at the nodes as ML/PP. Candolleomyces species produced in this study are indicated in bold.
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Figure 2. Basidiomata and microscopic features of Candolleomyces brevisporus. (AC) Basidiomata: (A,B) HMAS 258919 (holotype); (C) HMAS 258920, (D) Basidiospores, (E) Pileipellis, (F) Cheilocystidia, and (G) Basidia. Scale bars: 10 mm (AC); 5 μm (D); 20 μm (E); and 10 μm (F,G).
Figure 2. Basidiomata and microscopic features of Candolleomyces brevisporus. (AC) Basidiomata: (A,B) HMAS 258919 (holotype); (C) HMAS 258920, (D) Basidiospores, (E) Pileipellis, (F) Cheilocystidia, and (G) Basidia. Scale bars: 10 mm (AC); 5 μm (D); 20 μm (E); and 10 μm (F,G).
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Figure 3. Nucleotide differences of Candolleomyces brevisporus, C. shennongdingicus, C. gyirongicus, C. lignicola, and C. shennongjianus across ITS, LSU, and tef-1α. The numbers at the top indicate the positions of the polymorphic sites in each fragment. The dashes indicate the lack of data for the respective positions.
Figure 3. Nucleotide differences of Candolleomyces brevisporus, C. shennongdingicus, C. gyirongicus, C. lignicola, and C. shennongjianus across ITS, LSU, and tef-1α. The numbers at the top indicate the positions of the polymorphic sites in each fragment. The dashes indicate the lack of data for the respective positions.
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Figure 4. Basidiomata and microscopic features of Candolleomyces gyirongicus. (AF) Basidiomata: (A) HMAS 287607; (B) HMAS 287610; (C,D) HMAS 287611; (E) HMAS 287612 (holotype); (F) HMAS 287614, (G) Basidiospores, (H) Pileipellis, (I) Cheilocystidia, and (J) Basidia. Scale bars: 10 mm (AF); 5 μm (G); 20 μm (H); and 10 μm (I,J).
Figure 4. Basidiomata and microscopic features of Candolleomyces gyirongicus. (AF) Basidiomata: (A) HMAS 287607; (B) HMAS 287610; (C,D) HMAS 287611; (E) HMAS 287612 (holotype); (F) HMAS 287614, (G) Basidiospores, (H) Pileipellis, (I) Cheilocystidia, and (J) Basidia. Scale bars: 10 mm (AF); 5 μm (G); 20 μm (H); and 10 μm (I,J).
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Figure 5. Basidiomata and microscopic features of Candolleomyces lignicola. (A,B) Basidiomata: (A) HMAS 258922; (B) HMAS 258921 (holotype), (C) Basidiospores, (D) Pileipellis, (E) Cheilocystidia, (F) Basidia. Scale bars: 10 mm (A,B); 5 μm (C); 20 μm (D); and 10 μm (E,F).
Figure 5. Basidiomata and microscopic features of Candolleomyces lignicola. (A,B) Basidiomata: (A) HMAS 258922; (B) HMAS 258921 (holotype), (C) Basidiospores, (D) Pileipellis, (E) Cheilocystidia, (F) Basidia. Scale bars: 10 mm (A,B); 5 μm (C); 20 μm (D); and 10 μm (E,F).
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Figure 6. Basidiomata and microscopic features of Candolleomyces luridus. (AD) Basidiomata: (A) HMAS 258913, (B) HMAS 258913 (holotype), (C) HMAS 258914, (D) HMAS 258915, (E) Basidiospores, (F) Pileipellis, (G) Cheilocystidia, and (H) Basidia. Scale bars: 10 mm (AD); 5 μm (E); 20 μm (F); and 10 μm (G,H).
Figure 6. Basidiomata and microscopic features of Candolleomyces luridus. (AD) Basidiomata: (A) HMAS 258913, (B) HMAS 258913 (holotype), (C) HMAS 258914, (D) HMAS 258915, (E) Basidiospores, (F) Pileipellis, (G) Cheilocystidia, and (H) Basidia. Scale bars: 10 mm (AD); 5 μm (E); 20 μm (F); and 10 μm (G,H).
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Figure 7. Basidiomata and microscopic features of Candolleomyces shennongdingicus. (A,B) Basidiomata: (A) HMAS 258918 (holotype); (B) HMAS 258916, (C) Basidiospores, (D) Pileipellis, (E) Cheilocystidia, (F) Basidia. Scale bars: 10 mm (A,B); 5 μm (C); 20 μm (D); 10 μm (E,F).
Figure 7. Basidiomata and microscopic features of Candolleomyces shennongdingicus. (A,B) Basidiomata: (A) HMAS 258918 (holotype); (B) HMAS 258916, (C) Basidiospores, (D) Pileipellis, (E) Cheilocystidia, (F) Basidia. Scale bars: 10 mm (A,B); 5 μm (C); 20 μm (D); 10 μm (E,F).
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Figure 8. Basidiomata and microscopic features of Candolleomyces shennongjianus. (AC) Basidiomata: (A) HMAS 258909 (holotype); (B) HMAS 258907; (C) HMAS 258910, (D) Basidiospores, (E) Pileipellis, (F) Cheilocystidia, and (G) Basidia. Scale bars: 10 mm (AC); 5 μm (D); 20 μm (E); and 10 μm (F,G).
Figure 8. Basidiomata and microscopic features of Candolleomyces shennongjianus. (AC) Basidiomata: (A) HMAS 258909 (holotype); (B) HMAS 258907; (C) HMAS 258910, (D) Basidiospores, (E) Pileipellis, (F) Cheilocystidia, and (G) Basidia. Scale bars: 10 mm (AC); 5 μm (D); 20 μm (E); and 10 μm (F,G).
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Figure 9. Basidiomata and microscopic features of Candolleomyces sichuanicus. (AC) Basidiomata: (A) HMAS 287615; (B,C) HMAS 287616 (holotype), (D) Basidiospores, (E) Pileipellis, (F) Cheilocystidia, and (G) Basidia. Scale bars: 10 mm (AC); 5 μm (D); 20 μm (E); and 10 μm (F,G).
Figure 9. Basidiomata and microscopic features of Candolleomyces sichuanicus. (AC) Basidiomata: (A) HMAS 287615; (B,C) HMAS 287616 (holotype), (D) Basidiospores, (E) Pileipellis, (F) Cheilocystidia, and (G) Basidia. Scale bars: 10 mm (AC); 5 μm (D); 20 μm (E); and 10 μm (F,G).
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Table 1. Sequences used in the phylogenetic analysis in this study. Missing sequences are indicated by “–”.
Table 1. Sequences used in the phylogenetic analysis in this study. Missing sequences are indicated by “–”.
TaxonVoucherCountrynrITSnrLSUtef-1αReference
Candolleomyces aberdarensisGLMF116094KenyaMH880928[39]
C. aberdarensisIHIB618KenyaMK421517MK421517[10]
C. albipesDED8340Sao TomeKX017209[40]
C. albosquamosusTBGT18600 TypeIndiaOQ676550[15]
C. albosquamosusTBGT18895IndiaOQ676549[15]
C. albovagabundusHTBM1139 ParatypeChinaOR711038OR711054[19]
C. albovagabundusHKAS129660 TypeChinaOR711041OR711057OR727285[19]
C. asiaticusLAH36975PakistanOK392606OQ802843[14]
C. asiaticusLAH36809 TypePakistanNR182405NG229114[14]
C. badhyzensisTAA79478 TypeTurkmenistanKC992883KC992883[30]
C. badiophyllusSZMC-NL-2347-FN430699FM876268FM897252[41]
C. bivelatus110114MFBPZHO777ChinaMW554021Non-referenced
C. bivelatusMCVE29117ItalyMF325962MF521811[42]
C. brevisporusHMAS 258919 TypeChinaOR822167OR822149OR819986This study
C. brevisporusHMAS 258920ChinaOR822168OR822150OR819987This study
C. brunneopileatusTBGT18553IndiaOQ878483OR244398[20]
C. brunneopileatusTBGT18698 TypeIndiaOQ878348OR244401[20]
C. brunneovagabundusHKAS129659 TypeChinaOR711031OR711047OR791600[19]
C. brunneovagabundusHTBM1136 ParatypeChinaOR711036OR711052[19]
C. cacaoFP1R4USAKU847452Non-referenced
C. cacaoDED 8339 TypeSao TomeNR148106[40]
C. campanulatusLAH35719 TypePakistanOQ308881OQ802837[18]
C. campanulatusLAH37657PakistanOQ308882OQ802838[18]
C. candolleanusLAS73030 NeotypeSwedenKM030175KM030175[30]
C. candolleanusLÖ38-00SwedenDQ389720DQ389720[5]
C. cladii-marisciCLUF302 TypeItalyMK080112[43]
C. cladii-marisciSDBR-CMUNK0507ThailandMZ145228MZ145244[13]
C. efflorescensPegler2133(K)Sri LankaKC992941[30]
C. eurysporusGLMF126263 TypeViet NamMT651560MT651560[10]
C. gyirongicusHMAS 287607ChinaPP734608PP734619PP729321This study
C. gyirongicusHMAS 287608ChinaPP734609PP734620PP729322This study
C. gyirongicusHMAS 287609ChinaPP734610PP734621PP729323This study
C. gyirongicusHMAS 287610ChinaPP734611PP734622PP729324This study
C. gyirongicusHMAS 287611ChinaPP734612PP734623PP729325This study
C. gyirongicusHMAS 287612 TypeChinaPP734613PP734624PP729326This study
C. gyirongicusHMAS 287613ChinaPP734614PP734625PP729327This study
C. gyirongicusHMAS 287614ChinaPP734615PP734626PP729328This study
C. halophilusMICH AH-14321SpainMG825900[44]
C. incanusBJTC S173ChinaON042760ON042767ON098509[12]
C. incanusBJTC Z777 TypeChinaON042759ON042766ON098508[12]
C. leucotephrusLÖ138-01 (UPS)SwedenKC992885KC992885KJ732775[30]
C. leucotephrusMCVE28705SpainMF325979MF521791[42]
C. lignicolaHMAS 258921 TypeChinaOR822169OR822151OR819988This study
C. lignicolaHMAS 258922ChinaOR822170OR822152OR819989This study
C. luridusHMAS 258911ChinaOR822159OR822141OR819978This study
C. luridusHMAS 258912ChinaOR822160OR822142OR819979This study
C. luridusHMAS 258913 TypeChinaOR822161OR822143OR819980This study
C. luridusHMAS 258914ChinaOR822162OR822144OR819981This study
C. luridusHMAS 258915ChinaOR822163OR822145OR819982This study
C. luteopallidusHMJAU5148ChinaMG734736MW301084MW314073[45]
C. luteopallidusSharp20863 TypeUSAKC992884KC992884[30]
C. niveofloccosusTBGT18412 TypeIndiaOQ878345OR244387[20]
C. niveofloccosusTBGT18513IndiaOQ878251OR244394[20]
C. ruhunensisHKAS123158 TypeSri LankaON685315[17]
C. secotioidesAH31746 TypeMexicoKR003281KR003282KR003283[7]
C. shennongdingicusHMAS 258916ChinaOR822164OR822146OR819983This study
C. shennongdingicusHMAS 258917ChinaOR822165OR822147OR819984This study
C. shennongdingicusHMAS 258918 TypeChinaOR822166OR822148OR819985This study
C. shennongjianusHMAS 258907ChinaOR822155OR822137OR819974This study
C. shennongjianusHMAS 258908ChinaOR822156OR822138OR819975This study
C. shennongjianusHMAS 258909 TypeChinaOR822157OR822139OR819976This study
C. shennongjianusHMAS 258910ChinaOR822158OR822140OR819977This study
C. sichuanicusHMAS 287615ChinaPP734616PP734627PP729329This study
C. sichuanicusHMAS 287616 TypeChinaPP734617PP734628PP729330This study
C. sichuanicusHMAS 287617ChinaPP734618PP734629PP729331This study
C. sindhudeltaeLAH37632 TypePakistanOQ247908OQ247912[16]
C. sindhudeltaeLAH37633PakistanOQ247909OQ247913[16]
C. singeriHMJUA37867ChinaMG734718MW301088MW314077[45]
C. singeriHMJAU37877ChinaMW301073MW301091MW314080[11]
C. singeriHMAS 258923ChinaOR822171OR822153OR819990This study
C. singeriHMAS 258924ChinaOR822172OR822154OR819991This study
C. subcacaoHMJAU37807 TypeChinaMW301064MW301092MW314081[11]
C. subcacaoHMJAU37808ChinaMW301065MW301093MW314082[11]
C. subcandolleanusBJTC Z239 TypeChinaON042755ON042762ON098505[12]
C. subcandolleanusBJTC Z232ChinaON042756ON042763[12]
C. subminutisporusHMJAU37801 TypeChinaMW301066MW301094MW314083[11]
C. subminutisporusHMJAU37916ChinaMW301067MW301095MW314084[11]
C. subsingeriHMIAU37913ChinaMG734725MW301098MW314086[45]
C. subsingeriHMJAU37811 TypeChinaMG734715MW301097MW314085[45]
C. sulcatotuberculosusLÖ55-12GermanyKJ138422KJ138422[46]
C. sulcatotuberculosusHFJAU1515ChinaMW375696MW382965[11]
C. sultaniiLAH35714 TypePakistanOQ308835OQ801565[18]
C. sultaniiLAH37658PakistanOQ308836OQ801566[18]
C. thailandensisSDBR-CMUNK0442ThailandMZ145232[13]
C. thailandensisSDBR-CMUNK0443 TypeThailandMZ146874[13]
C. trinitatensisTL9035EcuadorKC992882KC992882[30]
C. tuberculatusADK4162SwedenKC992886KC992886[30]
C. typhaeLÖ21-04SwedenDQ389721DQ389721KJ732776[5]
C. yanshanensisBJTC Z783ChinaON042757ON042764ON098506[12]
C. yanshanensisBJTC Z110 TypeChinaON042758ON042765ON098507[12]
Hausknechtia floriformisWU22832 TypeVanuatuON745613ON745616ON746007[47]
H. floriformisWU22833VanuatuON745619ON745615ON746009[47]
H. leucostictaHFJAU1486 EpitypeChinaOL435561OL435565OL439896[47]
H. leucostictaHFJAU1526ChinaOL435563OL435566OL439897[47]
The sequences generated in this study are marked in bold.
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Han, X.-X.; Phurbu, D.; Ma, G.-F.; Li, Y.-Z.; Mei, Y.-J.; Liu, D.-M.; Lin, F.-C.; Zhao, R.-L.; Thongklang, N.; Cao, B. A Taxonomic Study of Candolleomyces Specimens from China Revealed Seven New Species. J. Fungi 2024, 10, 499. https://doi.org/10.3390/jof10070499

AMA Style

Han X-X, Phurbu D, Ma G-F, Li Y-Z, Mei Y-J, Liu D-M, Lin F-C, Zhao R-L, Thongklang N, Cao B. A Taxonomic Study of Candolleomyces Specimens from China Revealed Seven New Species. Journal of Fungi. 2024; 10(7):499. https://doi.org/10.3390/jof10070499

Chicago/Turabian Style

Han, Xi-Xi, Dorji Phurbu, Guo-Fei Ma, You-Zhi Li, Yu-Jiao Mei, Dong-Mei Liu, Fu-Cheng Lin, Rui-Lin Zhao, Naritsada Thongklang, and Bin Cao. 2024. "A Taxonomic Study of Candolleomyces Specimens from China Revealed Seven New Species" Journal of Fungi 10, no. 7: 499. https://doi.org/10.3390/jof10070499

APA Style

Han, X. -X., Phurbu, D., Ma, G. -F., Li, Y. -Z., Mei, Y. -J., Liu, D. -M., Lin, F. -C., Zhao, R. -L., Thongklang, N., & Cao, B. (2024). A Taxonomic Study of Candolleomyces Specimens from China Revealed Seven New Species. Journal of Fungi, 10(7), 499. https://doi.org/10.3390/jof10070499

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