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Research Article
Taxonomy and phylogeny of Auriculariales (Agaricomycetes, Basidiomycota) with descriptions of four new species from south-western China
expand article infoJunhong Dong‡, Yonggao Zhu‡, Chengbin Qian‡, Changlin Zhao‡
‡ Southwest Forestry University, Kunming, China

Corresponding author: Changlin Zhao ( fungichanglinz@163.com )

Academic editor: Ajay Kumar Gautam
© 2024 Junhong Dong, Yonggao Zhu, Chengbin Qian, Changlin Zhao.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Dong J, Zhu Y, Qian C, Zhao C (2024) Taxonomy and phylogeny of Auriculariales (Agaricomycetes, Basidiomycota) with descriptions of four new species from south-western China. MycoKeys 108: 115-146. https://doi.org/10.3897/mycokeys.108.128659
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Abstract

The wood-inhabiting fungi play an integral role in wood degradation and the cycle of matter in the ecological system. They are considered as the “key player” in wood decomposition, because of their ability to produce lignocellulosic enzymes that break down woody lignin, cellulose and hemicellulose. In the present study, four new wood-inhabiting fungal species, Adustochaete albomarginata, Ad. punctata, Alloexidiopsis grandinea and Al. xantha collected from southern China, are proposed, based on a combination of morphological features and molecular evidence. Adustochaete albomarginata is characterised by resupinate basidiomata with cream to buff, a smooth, cracked, hymenial surface, a monomitic hyphal system with clamped generative hyphae and subcylindrical to allantoid basidiospores (12–17.5 × 6.5–9 µm). Adustochaete punctata is characterised by resupinate basidiomata with cream, a smooth, punctate hymenial surface, a monomitic hyphal system with clamped generative hyphae and subcylindrical to allantoid basidiospores (13.5–18 × 6–8.2 µm). Alloexidiopsis grandinea is characterised by resupinate basidiomata with buff to slightly yellowish, a grandinioid hymenial surface, a monomitic hyphal system with clamped generative hyphae and allantoid basidiospores (10–12.3 × 5–5.8 µm). Additionally, Alloexidiopsis xantha is characterised by resupinate basidiomata with cream to slightly buff, a smooth hymenial surface, a monomitic hyphal system with clamped generative hyphae and subcylindrical to allantoid basidiospores measuring 20–24 × 5–6.2 µm. Sequences of the internal transcribed spacers (ITS) and the large subunit (nrLSU) of the nuclear ribosomal DNA (rDNA) markers of the studied samples were generated. Phylogenetic analyses were performed with the Maximum Likelihood, Maximum Parsimony and Bayesian Inference methods. The phylogram, based on the ITS+nLSU rDNA gene regions, revealed that four new species were assigned to the genera Adustochaete and Alloexidiopsis within the order Auriculariales, individually. The phylogenetic tree inferred from the ITS sequences highlighted that Ad. albomarginata was retrieved as a sister to Ad. yunnanensis and the species Ad. punctata was sister to Ad. rava. The topology, based on the ITS sequences, showed that Al. grandinea was retrieved as a sister to Al. schistacea and the taxon Al. xantha formed a monophyletic lineage. Furthermore, two identification keys to Adustochaete and Alloexidiopsis worldwide are provided.

Key words

Biodiversity, molecular systematics, taxonomy, wood-inhabiting fungi, Yunnan Province

Introduction

In forest ecosystems, fungi play an essential ecological role to drive carbon cycling in forest soils, mediate mineral nutrition of plants and alleviate carbon limitations (Tedersoo et al. 2014). The fungal order Auriculariales is a group mainly composed of wood-inhabiting fungi in Agaricomycetes Doweld (Basidiomycota) (Hibbett et al. 2007). The type genus of this order is Auricularia Bull., in which several other gelatinous genera Exidia Fr., Guepinia Fr. and Pseudohydnum P. Karst., comprise important edible and medicinal fungi (Wu et al. 2019; Liu et al. 2022). Therefore, interest in species diversity in gelatinous genera has increased significantly in recent years (Chen et al. 2020; Shen and Fan 2020; Ye et al. 2020; Wang and Thorn 2021; Wu et al. 2021; Tohtirjap et al. 2023).

Contrary to the gelatinous genera, most species in the order Auriculariales are tough, include saprophytic species with resupinate, effused-reflexed, hydnoid, cerebriform, coralloid or pileate basidiomata (Wells and Bandoni 2001; Miettinen et al. 2012; Hibbett et al. 2014; Malysheva and Spirin 2017; Alvarenga et al. 2019; Spirin et al. 2019a, 2019b; Liu et al. 2022; Tohtirjap et al. 2023). Species with the stereoid basidiocarps are widely distributed in many orders of the Agaricomycetes, although they are certainly a minority in the order Auriculariales (Malysheva and Spirin 2017).

The genus Adustochaete Alvarenga & K.H. Larss. was erected by Alvarenga and Larsson and typed by the taxon Ad. rava Alvarenga & K.H. Larss. It is characterised by the resupinate basidiomata, spiny or tuberculate hymenophore, a monomitic hyphal structure with clamp connections on generative hyphae, present cystidia and hyphidia, ellipsoid-ovoid to obconical basidia, cylindrical to broadly cylindrical, straight or curved basidiospores (Alvarenga et al. 2019). The genus Alloexidiopsis L.W. Zhou & S.L. Liu is typified by Al. schistacea L.W. Zhou & S.L. Liu, which is c characterised by annual, resupinate basidiomata, smooth or with sterile spines hymenophore, a monomitic hyphal structure with clamp connections on generative hyphae, present cystidia and hyphidia, ellipsoid to ovoid, septate basidia, and cylindrical to broadly cylindrical, slightly curved (allantoid) basidiospores (Liu et al. 2022). Based on the MycoBank database (http://www.mycobank.org, accessed on 25 July 2024) and the Index Fungorum (http://www.indexfungorum.org, accessed on 25 July 2024), the genera Adustochaete and Alloexidiopsis have registered four and six species, respectively (Alvarenga et al. 2019; Guan et al. 2020; Hyde et al. 2020; Li et al. 2022a, 2022b; Li and Zhao 2022; Liu et al. 2022; Dong et al. 2024).

Classification of the kingdom of fungi has been updated continuously, based on the frequent inclusion of data from DNA sequences in many phylogenetic studies (Wijayawardene et al. 2020, 2022). Based on the early embrace of molecular systematics by mycologists, both the discovery and classification of fungi, amongst the more basal branches of the tree, are now coming to light from genomic analyses and environmental DNA surveys that have been conducted (James et al. 2020). Based on both the morphological and phylogenetic evidence, the generic concepts of Eichleriella Bres., Hirneolina (Pat.) Bres. and Tremellochaete Raitv. were revised, in which Malysheva and Spirin (2017) proposed that the genus Heteroradulum Lloyd ex Spirin and Malysheva was validated. The genus Eichleriella was accepted to be a monophyletic genus, while both genera Exidiopsis (Bref.) Möller and Heterochaete Pat. seemed to be synonymous, with priority given to the latter genus (Malysheva and Spirin 2017; Alvarenga et al. 2019; Alvarenga and Gibertoni 2021). However, certain species of Exidiopsis, even sequenced ones such as E. calcea (Pers.) K. Wells and E. grisea (Bres.) Bourdot & Maire, still have no appropriate placement at the generic level (Malysheva and Spirin 2017; Li et al. 2022a; Liu et al. 2022).

In recent years, the species diversity of the resupinate Auriculariales have been described or better defined using morphological and molecular analyses and the results showed the hidden diversity of this group and several corticioid genera, for example, Adustochaete, Alloexidiopsis, Amphistereum Spirin & Malysheva, Crystallodon Alvarenga, Heteroradulum, Metulochaete Alvarenga, Proterochaete Spirin & Malysheva and Sclerotrema Spirin & Malysheva, which have been established and described, based on the morphological and phylogenetic studies (Malysheva and Spirin 2017; Alvarenga et al. 2019; Spirin et al. 2019a, 2019b; Alvarenga and Gibertoni 2021; Liu et al. 2022).

During investigations on wood-inhabiting fungi in the Yunnan-Guizhou Plateau, China, many specimens were collected. To clarify the placement and relationships of these specimens, we carried out a phylogenetic and taxonomic study, based on the ITS+nLSU and ITS sequences. These specimens were assigned to the genera Adustochaete and Alloexidiopsis within the order Auriculariales. Therefore, four new species Ad. albomarginata, Ad. punctata, Al. grandinea and Al. xantha are proposed with description and illustrations, based on the morphological characteristics and phylogenetic analyses.

Materials and methods

Sample collection and herbarium specimen preparation

The fresh fruiting bodies were collected on the fallen angiosperm branches from Dali, Dehong, Diqing, Lincang and Zhaotong of Yunnan Province, China. The samples were photographed in situ and fresh macroscopic details were recorded. Photographs were recorded by a Nikon D7100 camera. All the photos were focus-stacked using Helicon Focus software. Macroscopic details were recorded and transported to a field station where the fruit body was dried on an electronic food dryer at 45 °C. Once dried, the specimens were sealed in an envelope and zip-lock plastic bags and labelled (Zhang et al. 2024). The dried specimens were deposited in the Herbarium of the Southwest Forestry University (SWFC), Kunming, Yunnan Province, China.

Morphology

The macromorphological descriptions were based on field notes and photos captured in the field and lab. The colour terminology follows Petersen (1996). The micromorphological data were obtained from the dried specimens after observation under a light microscope with a magnification of 10 × 100 oil (Zhao et al. 2023). Sections mounted in 5% potassium hydroxide (KOH) and 2% phloxine B dye (C20H2Br4Cl4Na2O5) and we also used other reagents, including Cotton Blue and Melzer’s reagent to observe micromorphology following Wu et al. (2022b). To show the variation in spore sizes, 5% of measurements were excluded from each end of the range and shown in parentheses. At least thirty basidiospores from each specimen were measured. Stalks were excluded from basidia measurements and the hilar appendage was excluded from basidiospores measurements. The following abbreviations are used: KOH = 5% potassium hydroxide water solution, CB– = acyanophilous, IKI– = both inamyloid and non-dextrinoid, L = mean spore length (arithmetic average for all spores), W = mean spore width (arithmetic average for all spores), Q = variation in the L/W ratios between the specimens studied, Qm represented the average Q of basidiospores measured ± standard deviation and n = a/b (number of spores (a) measured from given number (b) of specimens).

Molecular phylogeny

The CTAB rapid plant genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd., Beijing, China) was used to obtain genomic DNA from the dried specimens according to the manufacturer’s instructions. The ITS region was amplified with ITS5 and ITS4 primers (White et al. 1990). The nLSU region was amplified with the LR0R and LR7 (Vilgalys and Hester 1990; Rehner and Samuels 1994). The PCR procedure for ITS was as follows: initial denaturation at 95 °C for 3 min, followed by 35 cycles at 94 °C for 40 s, 58 °C for 45 s and 72 °C for 1 min and a final extension of 72 °C for 10 min. The PCR procedure for nLSU was as follows: initial denaturation at 94 °C for 1 min, followed by 35 cycles at 94 °C for 30 s, 48 °C for 1 min and 72 °C for 1.5 min and a final extension of 72 °C for 10 min. The PCR products were purified and sequenced at Kunming Tsingke Biological Technology Limited Company (Yunnan Province, P.R. China). The newly-generated sequences were deposited in NCBI GenBank (Table 1).

Table 1.
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List of species, specimens, and GenBank accession number of sequences used in this study.

Species Name Sample No. GenBank Accession No. Country References
ITS nLSU
Adustochaete albomarginata CLZhao 22774 * PP852049 PP849033 China Present study
Adustochaete interrupta LR 23435 MK391518 MK391527 Brazil Alvarenga et al. (2019)
Adustochaete nivea RLMA 531 MN165954 MN165989 USA Liu et al. (2022)
Adustochaete punctata CLZhao 29669 PP852050 — China Present study
Adustochaete punctata CLZhao 29671 PP852051 PP849034 China Present study
Adustochaete punctata CLZhao 29675 * PP852052 PP849035 China Present study
Adustochaete punctata CLZhao 29685 PP852053 PP849036 China Present study
Adustochaete punctata CLZhao 29686 PP852054 PP849037 China Present study
Adustochaete punctata CLZhao 29706 PP852055 — China Present study
Adustochaete punctata CLZhao 29710 PP852056 PP849038 China Present study
Adustochaete punctata CLZhao 29711 PP852057 PP849039 China Present study
Adustochaete rava RC 841 MK391516 — Brazil Alvarenga et al. (2019)
Adustochaete rava KHL 15526 MK391517 MK391526 Brazil Alvarenga et al. (2019)
Adustochaete yunnanensis CLZhao 8212 MZ911964 MZ950629 China Li and Zhao (2022)
Adustochaete yunnanensis CLZhao 4671 MZ911965 — China Li and Zhao (2022)
Adustochaete yunnanensis CLZhao 4401 MZ911966 MZ950630 China Li and Zhao (2022)
Alloexidiopsis australiensis LWZ 20180514-18 OM801934 OM801919 China Liu et al. (2022)
Alloexidiopsis australiensis LWZ 20180513-22 OM801933 OM801918 China Liu et al. (2022)
Alloexidiopsis calcea LWZ 20180904-14 OM801935 OM801920 China Liu et al. (2022)
Alloexidiopsis calcea MW 331 AF291280 AF291326 Germany Weiß and Oberwinkler (2001)
Alloexidiopsis grandinea CLZhao 33798 * PP852058 — China Present study
Alloexidiopsis grandinea CLZhao 34279 PP852059 — China Present study
Alloexidiopsis nivea CLZhao 11204 MZ352947 MZ352938 China Li et al. (2022a)
Alloexidiopsis nivea CLZhao 11210 MZ352948 MZ352939 China Li et al. (2022a)
Alloexidiopsis schistacea LWZ 20200819-21a OM801939 OM801932 China Liu et al. (2022)
Alloexidiopsis xantha CLZhao 25093 * PP852060 PP849040 China Present study
Alloexidiopsis yunnanensis CLZhao 8106 MT215569 MT215565 China Guan et al. (2020)
Alloexidiopsis yunnanensis CLZhao 4023 MT215568 MT215564 China Guan et al. (2020)
Amphistereum leveilleanum FP-106715 KX262119 KX262168 USA Malysheva and Spirin (2017)
Amphistereum schrenkii HHB 8476 KX262130 KX262178 USA Malysheva and Spirin (2017)
Aporpium caryae Miettinen 14774 JX044145 — Finland Miettinen et al. (2012)
Aporpium caryae WD 2207 AB871751 AB871730 Japan Sotome et al. (2014)
Auricularia auricula-judae JT 04 KT152099 KT152115 UK Tohtirjap et al. (2023)
Auricularia cornea Dai 13621 MZ618936 MZ669905 China Tohtirjap et al. (2023)
Auricularia polytricha TUFC 12920 AB871752 AB871733 Japan Sotome et al. (2014)
Auricularia tibetica Dai 13336 MZ618943 MZ669915 China Tohtirjap et al. (2023)
Bourdotia galzinii Otto MiettinenX3067 MG757511 MG757511 Spain Malysheva et al. (2018)
Crystallodon subgelatinosum RC 1609-URM93444 MN475884 MN475888 Brazil Alvarenga and Gibertoni (2021)
Crystallodon subgelatinosum TBG BF-18001-URM93445 MN475885 MN475889 Brazil Alvarenga and Gibertoni (2021)
Ductifera sucina KW3886 AY509551 AY509551 Canada Liu et al. (2022)
Eichleriella bactriana TAAM 55071 KX262121 KX262170 Russia Malysheva and Spirin (2017)
Eichleriella crocata TAAM 101077 KX262100 KX262147 Russia Malysheva and Spirin (2017)
Eichleriella leucophaea Barsukova LE 303261 KX262111 KX262161 Russia Malysheva and Spirin (2017)
Eichleriella tenuicula ValCB 1 MK391515 MK391525 Brazil Alvarenga et al. (2019)
Elmerina cladophora Miettinen 14314 MG757509 MG757509 Indonesia Malysheva et al. (2018)
Elmerina sclerodontia Miettinen 16431 MG757512 MG757512 Malaysia Malysheva et al. (2018)
Exidia glandulosa YC Dai 21232 MT663362 MT664781 China Wu et al. (2020)
Exidia glandulosa YC Dai 21233 MT663363 MT664782 China Wu et al. (2020)
Exidia pithya MW 313 AF291275 AF291321 Germany Weiß and Oberwinkler (2001)
Grammatus labyrinthinus Yuan 1600 KM379139 KM379140 China Alvarenga et al. (2019)
Grammatus semis OM10618 KX262146 KX262194 China Malysheva and Spirin (2017)
Heteroradulum adnatum LR 23453 KX262116 KX262165 Mexico Tohtirjap et al. (2023)
Heteroradulum kmetii VS 6466 KX262104 KX262152 Russia Malysheva and Spirin (2017)
Hyalodon piceicola Spirin 2689 MG735414 MG735422 Russia Spirin et al. (2019a)
Hyalodon piceicola Spirin 11063 MG735415 MG735423 Russia Spirin et al. (2019a)
Mycostilla vermiformis Spirin 11330 MG735417 MG735425 Russia Spirin et al. (2019a)
Mycostilla vermiformis OF 188059 MG735418 — Russia Spirin et al. (2019a)
Myxarium cinnamomescens OF160494 KY801882 KY801909 Russia Spirin et al. (2018)
Myxarium grilletii VS9016 MK098896 MK098944 Russia Spirin et al. (2019b)
Myxarium hyalinum TL2012 443455 KY801880 KY801907 Russia Spirin et al. (2018)
Myxarium legonii VS 8986 MK098899 MK098947 Russia Spirin et al. (2019b)
Protodaedalea foliacea Miettinen 13 054 MG757507 MG757507 Finland Malysheva et al. (2018)
Protodaedalea hispida Spirin 5139 MG757510 MG757510 Finland Malysheva et al. (2018)
Protodontia africana AS 171126 1104 MK098978 MK098973 Russia Spirin et al. (2019b)
Protohydnum cartilagineum SP 467240 MG735419 MG735426 Russia Malysheva et al. (2018)
Protomerulius dubius VS 3019 MK484041 MK480553 Russia Spirin et al. (2019a)
Protomerulius minor KHL 15937 MK484060 MK480569 Russia Spirin et al. (2019a)
Protomerulius substuppeus O 19171 JX134482 JQ764649 China Spirin et al. (2019a)
Pseudohydnum gelatinosum F14063 AF384861 AF384861 Canada Weiß and Oberwinkler (2001)
Pseudohydnum gelatinosum AFTOL ID1875 DQ520094 DQ520094 Germany Lutzoni et al. (2004)
Stypellopsis farlowii Larsson 12337 MG857095 MG857099 Russia Spirin et al. (2018)
Stypellopsis hyperborea J Norden 9751 MG857097 MG857101 Russia Spirin et al. (2018)
Tremellochaete atlantica URM90199 MG594381 MG594383 Brazil Alvarenga et al. (2019)
Tremellochaete japonica TAA 42689 AF291274 AF291320 Russia Weiß and Oberwinkler (2001)
Tremiscus helvelloides AFTOL ID1680 DQ520100 DQ520100 Germany Lutzoni et al. (2004)
Sistotrema brinkmannii isolate 236 JX535169 JX535170 Netherlands Alvarenga and Gibertoni (2021)

New species is shown in bold; * is shown type material, holotype.

The sequences were aligned in MAFFT v. 7 (Katoh et al. 2019) using the G-INS-i strategy. The alignment was adjusted manually using AliView v. 1.27 (Larsson 2014). The dataset was aligned first and then the sequences of ITS+nLSU were combined with Mesquite v. 3.51. The combined ITS+nLSU sequences and ITS datasets were used to infer the position of the new species and related species. The sequence of Sistotrema brinkmannii (Bres.) J. Erikss. obtained from GenBank was used as an outgroup to root trees in the ITS+nLSU analysis (Fig. 1) in the order Auriculariales (Tohtirjap et al. 2023). The sequence of Amphistereum leveilleanum (Berk. & M.A. Curtis) Spirin & Malysheva obtained from GenBank was used as an outgroup to root trees in the ITS analysis in the genus Adustochaete (Fig. 2). The sequence of Heteroradulum kmetii (Bres.) Spirin & Malysheva obtained from GenBank was used as an outgroup to root trees in the ITS analysis in the genus Alloexidiopsis (Fig. 3).

Figure 1. 

Maximum parsimony strict consensus tree illustrating the phylogeny of Adustochaete and Alloexidiopsis and related genera in the order Auriculariales, based on ITS+nLSU sequences. Branches are labelled with Maximum Likelihood bootstrap value ≥ 70%, parsimony bootstrap value ≥ 50% and Bayesian posterior probabilities ≥ 0.95.

Figure 2. 

Maximum parsimony strict consensus tree illustrating the phylogeny of the two new species and related genera in the genus Adustochaete, based on ITS sequences. Branches are labelled with Maximum Likelihood bootstrap value ≥ 70%, parsimony bootstrap value ≥ 50% and Bayesian posterior probabilities ≥ 0.95.

Figure 3. 

Maximum parsimony strict consensus tree illustrating the phylogeny of the two new species and related genera in the genus Alloexidiopsis, based on ITS sequences. Branches are labelled with Maximum Likelihood bootstrap value ≥ 70%, parsimony bootstrap value ≥ 50% and Bayesian posterior probabilities ≥ 0.95.

Maximum Parsimony (MP), Maximum Likelihood (ML) and Bayesian Inference (BI) analyses were applied to the combined three datasets following a previous study (Zhao and Wu 2017) and the tree construction procedure was performed in PAUP* v. 4.0b10 (Swofford 2002). All of the characters were equally weighted and gaps were treated as missing data. Using the heuristic search option with TBR branch swapping and 1000 random sequence additions, trees were inferred. Maxtrees were set to 5000, branches of zero length were collapsed and all parsimonious trees were saved. Clade robustness was assessed using bootstrap (BT) analysis with 1000 replicates (Felsenstein 1985). Descriptive tree statistics, tree length (TL), the consistency index (CI), the retention index (RI), the rescaled consistency index (RC) and the homoplasy index (HI) were calculated for each maximum parsimonious tree generated. The multiple sequence alignment was also analysed using Maximum Likelihood (ML) in RAxML-HPC2 on XSEDE v. 8.2.8 with default parameters (Miller et al. 2012). Branch support (BS) for ML analysis was determined by 1000 bootstrap replicates.

jModelTest v. 2 (Darriba et al. 2012) was used to determine the best-fit evolution model for each dataset for the purposes of Bayesian Inference (BI), which was performed using MrBayes 3.2.7a with a GTR+I+G model of DNA substitution and a gamma distribution rate variation across sites (Ronquist et al. 2012). The first one-quarter of all the generations were discarded as burn-in. The majority-rule consensus tree of all the remaining trees was calculated. Branches were considered significantly supported if they received a Maximum Likelihood bootstrap value (BS) of > 70%, a Maximum Parsimony bootstrap value (BT) of > 70% or Bayesian Posterior Probabilities (BPP) of > 0.95.

Results

Sequence similarity search

The results of BLAST queries in NCBI, based on ITS and nLSU separately, showed the sequences producing significant alignment descriptions:

Adustochaete albomarginata: in ITS BLAST results, Ad. rava, Exidia saccharina Fr., Ea. qinghaiensis S.R. Wang & Thorn, Ad. nivea Alvarenga and Exidiopsis mucedinea (Pat.) K. Wells were found as the top ten taxa (maximum record descriptions: Max score 830; Total score 830; Query cover 96%; E value 0.0; Ident 92.93%). In nLSU BLAST results, Alloexidiopsis yunnanensis (C.L. Zhao) L.W. Zhou & S.L. Liu, Auricularia asiatica Bandara & K.D. Hyde, Au. brasiliana Y.C. Dai & F. Wu and Steccherinum nandinae (F. Wu, P. Du & X.M. Tian) Z.B. Liu, Y.C. Dai & Jing Si were found as the top ten taxa (maximum record descriptions: Max score 2398; Total score 2398; Query cover 98%; E value 0.0; Ident 98.60%).

Adustochaete punctata: in ITS BLAST results, Ad. rava, Ad. nivea, Exidiopsis mucedinea and Exidia candida Lloyd were found as the top ten taxa (maximum record descriptions: Max score 959; Total score 959; Query cover 96%; E value 0.0; Ident 96.74%). In nLSU BLAST results, Ad. rava, Ad. yunnanensis Y.F. Li & C.L. Zhao., Auricularia thailandica Bandara & K.D. Hyde, Au. scissa Looney, Birkebak & Matheny, Au. nigricans (Sw.) Birkebak, Looney & Sánchez-García and Alloexidiopsis yunnanensis were found as the top ten taxa (maximum record descriptions: Max score 2464; Total score 2464; Query cover 98%; E value 0.0; Ident 99.34%).

Alloexidiopsis grandinea: in ITS BLAST results, Ad. nivea and Al. schistacea were found as the top ten taxa (maximum record descriptions: Max score 861; Total score 861; Query cover 91%; E value 0.0; Ident 94.94%).

Alloexidiopsis xantha: in ITS BLAST results, Al. sinensis J.H. Dong & C.L. Zhao was found as the top ten taxa (maximum record descriptions: Max score 832; Total score 832; Query cover 98%; E value 0.0; Ident 92.42%). In nLSU BLAST results, Al. sinensis and Al. yunnanensis were found as the top ten taxa (maximum record descriptions: Max score 2457; Total score 2457; Query cover 99%; E value 0.0; Ident 99.05%).

The aligned dataset comprised 70 specimens representing 53 species. Four Markov chains were run for two runs from random starting trees, each for two million generations for the combine ITS+nLSU (Fig. 1) dataset with trees and parameters sampled every 1000 generations. The dataset had an aligned length of 2333 characters, of which 1301 characters are constant, 368 are variable and parsimony uninformative and 664 are parsimony informative. Maximum parsimony analysis yielded 120 equally parsimonious trees (TL = 4342, CI = 0.4000, HI = 0.6000, RI = 0.5288 and RC = 0.2115). The best model for the ITS+nLSU dataset, estimated and applied in the Bayesian analysis, was SYM+I+G. Both Bayesian analysis and ML analysis resulted in a similar topology to MP analysis with an average standard deviation of split frequencies = 0.008542 (BI) and the effective sample size (ESS) for Bayesian analysis across the two runs is double of the average ESS (avg. ESS) = 395.5.

The aligned dataset comprised 17 specimens representing seven species. Four Markov chains were run for two runs from random starting trees, each for 0.5 million generations for the ITS (Fig. 2) dataset with trees and parameters sampled every 1000 generations. The dataset had an aligned length of 522 characters, of which 413 characters are constant, 47 are variable and parsimony uninformative and 62 are parsimony informative. Maximum parsimony analysis yielded four equally parsimonious trees (TL = 161, CI = 0.8075, HI = 0.1925, RI = 0.8306 and RC = 0.6707). The best model for the ITS dataset, estimated and applied in the Bayesian analysis, was SYM+G. Both Bayesian analysis and ML analysis resulted in a similar topology to MP analysis with an average standard deviation of split frequencies = 0.006786 (BI) and the effective sample size (ESS) for Bayesian analysis across the two runs is double the average ESS (avg. ESS) = 617.

The aligned dataset comprised 13 specimens representing eight species. Four Markov chains were run for two runs from random starting trees, each for 0.3 million generations for the ITS (Fig. 3) dataset with trees and parameters sampled every 1000 generations. The dataset had an aligned length of 562 characters, of which 417 characters are constant, 64 are variable and parsimony uninformative and 81 are parsimony informative. Maximum parsimony analysis yielded two equally parsimonious trees (TL = 218, CI = 0.784, HI = 0.2156, RI = 0.7814 and RC = 0.6129). The best model for the ITS dataset, estimated and applied in the Bayesian analysis, was SYM+G. Both Bayesian analysis and ML analysis resulted in a similar topology to MP analysis with an average standard deviation of split frequencies = 0.007707 (BI) and the effective sample size (ESS) for Bayesian analysis across the two runs is double of the average ESS (avg. ESS) = 639.5.

The phylogram, based on the combined ITS+nLSU sequences (Fig. 1) analysis, showed that four new species Ad. albomarginata, Ad. punctata, Al. grandinea and Al. xantha were assigned to the genera Adustochaete and Alloexidiopsis within the order Auriculariales, individually. The phylogenetic tree, based on ITS sequences (Fig. 2), revealed that Ad. albomarginata was retrieved as a sister to Ad. yunnanensis. The taxon Ad. punctata was sister to Ad. rava. The topology, based on the ITS sequences (Fig. 3), revealed that Al. grandinea was retrieved as a sister to Al. schistacea and the species Al. xantha formed a monophyletic lineage.

Taxonomy

Adustochaete albomarginata J.H Dong & C.L. Zhao, sp. nov.

MycoBank No: 854168
Figs 4, 5, 6

Diagnosis

Differs from other Adustochaete species by its soft membranaceous basidiomata with cream to buff, smooth, cracked hymenial surface, a monomitic hyphal system with clamped generative hyphae and subcylindrical to allantoid basidiospores measuring 12–17.5 × 6.5–9 µm.

Holotype

China • Yunnan Province, Dali, Weishan County, Leqiu Town, Zhongyao Village, 25°01′N, 100°19′E, altitude 1910 m, on the fallen branch of angiosperm, leg. C.L. Zhao, 19 July 2022, CLZhao 22774 (SWFC).

Figure 4. 

Basidiomata of Adustochaete albomarginata in general and detailed views (CLZhao 22774, holotype). Scale bars: 1 cm (A); 1 mm (B).

Etymology

albomarginata (Latin or Greek origin): referring to the white margin of the basidiomata.

Basidiomata

Annual, resupinate, closely adnate, soft membranaceous, very hard to separate from substrate, without odour or taste when fresh, becoming coriaceous upon drying, up to 5 cm long, 1.5 cm wide, 50–100 µm thick. Hymenial surface smooth, white to cream when fresh, turning to cream to buff upon drying, cracked. Sterile margin white, thinning out, up to 0.5 mm wide.

Figure 5. 

Sections of hymenium of Adustochaete albomarginata (holotype, CLZhao 22774) A basidiospores B basidia C cystidia D hyphidia. Scale bars: 20 µm (A–D); 10 × 100 Oil.

Hyphal system

Monomitic, generative hyphae with clamp connections, colourless, thin-walled, unbranched, interwoven, 2.5–3.5 µm in diameter; IKI–, CB–, tissues unchanged in KOH. Hymenium. Cystidia numerous, thin-walled, subclavate to fusiform with an acute or obtuse apex, occasionally sinuous in the basal, 23.5–48.5 × 10–13.5 µm, with a clamp connection at base; cystidioles absent. Hyphidia arising from generative hyphae, nodulose, branched, colourless, thin-walled, 2.5–5 µm in diameter. Basidia ellipsoid to ovoid, longitudinally septate, two to four-celled, 17–24.5 × 11–16.5 µm; basidioles dominant, similar to basidia in shape, but slightly smaller. Basidiospores. Subcylindrical to allantoid, slightly curved, colourless, smooth, thin-walled, with 1–2 oil drops, IKI–, CB–, (11.5–)12–17.5(–18) × 6.5–9(–9.5) µm, L = 14.66 µm, W = 7.80 µm, Q = 1.72–1.99, Qm = 1.88 ± 0.08 (n = 30/1).

Figure 6. 

Microscopic structures of Adustochaete albomarginata (holotype, CLZhao 22774) A basidiospores B basidia C basidioles D cystidia E hyphidia F part of the vertical section of hymenium. Scale bars: 10 µm (A–F).

Adustochaete punctata J.H Dong & C.L. Zhao, sp. nov.

MycoBank No: 854170
Figs 7, 8, 9

Diagnosis

Differs from other Adustochaete species by its membranaceous basidiomata with cream, smooth, punctate hymenial surface, a monomitic hyphal system with clamped generative hyphae and subcylindrical to allantoid basidiospores measuring 13.5–18 × 6–8.2 µm.

Holotype

China • Yunnan Province, Dehong, Yingjiang County, Tongbiguan Provincial Nature Reserve, 23°48′N, 97°38′E, altitude 1500 m, on the fallen branch of angiosperm, leg. C.L. Zhao, 17 July 2023, CLZhao 29675 (SWFC).

Etymology

punctata (Latin or Greek origin): referring to the punctate hymenial surface of the specimen.

Basidiomata

Annual, resupinate, closely adnate, membranaceous, very hard to separate from substrate, without odour or taste when fresh, becoming coriaceous upon drying, up to 10 cm long, 1.5 cm wide, 100–250 µm thick. Hymenial surface smooth, punctate, white to cream when fresh, turning to cream upon drying. Sterile margin cream, thinning out, up to 1 mm wide.

Figure 7. 

Basidiomata of Adustochaete punctata in general and detailed views (CLZhao 29675, holotype). Scale bars: 1 cm (A); 1 mm (B).

Hyphal system

Monomitic, generative hyphae with clamp connections, colourless, thin-walled, unbranched, interwoven, 1.5–3.5 µm in diameter; IKI–, CB–, tissues unchanged in KOH. Hymenium. Cystidia numerous, thin-walled, subcylindrical to clavate with an obtuse apex, occasionally sinuous in the basal, 15.5–23.5 × 5.5–7.5 µm, with a clamp connection at base; cystidioles absent. Hyphidia arising from generative hyphae, nodulose, branched, colourless, thin-walled, 1.5–5 μm in diameter. Basidia ellipsoid to ovoid, longitudinally septate, two to four-celled, 17–25 × 16.5–21 µm; basidioles dominant, similar to basidia in shape, but slightly smaller. Basidiospores. Subcylindrical to allantoid, slightly curved, colourless, smooth, thin-walled, with several oil drops, IKI–, CB–, (13–)13.5–18(–18.5) × (5.5–)6–8.2(–8.5) µm, L = 15.78 µm, W = 6.79 µm, Q = 2.15–2.40 Qm = 2.32 ± 0.08 (n = 90/3).

Figure 8. 

Sections of hymenium of Adustochaete punctata (holotype, CLZhao 29675) A basidiospores B basidia C cystidia D hyphidia. Scale bars: 20 µm (A–D); 10 × 100 Oil.

Additional specimens examined

China • Yunnan Province, Dehong, Yingjiang County, Tongbiguan Provincial Nature Reserve, 23°48′N, 97°38′E, altitude 1500 m, on the fallen branch of angiosperm, leg. C.L. Zhao, 17 July 2023, CLZhao 29669; CLZhao 29671; CLZhao 29685; CLZhao 29686; CLZhao 29706; CLZhao 29710; CLZhao 29711 (SWFC).

Figure 9. 

Microscopic structures of Adustochaete punctata (holotype, CLZhao 29675) A basidiospores B basidia C basidioles D cystidia E hyphidia F part of the vertical section of hymenium. Scale bars: 10 µm (A–F).

Alloexidiopsis grandinea J.H Dong & C.L. Zhao, sp. nov.

MycoBank No: 854171
Figs 10, 11, 12

Diagnosis

Differs from other Alloexidiopsis species by its membranaceous basidiomata with buff to slightly yellowish, grandinioid hymenial surface, a monomitic hyphal system with clamped generative hyphae and cylindrical to allantoid basidiospores measuring 10–12.3 × 5–5.8 µm.

Holotype

China • Yunnan Province, Zhaotong, Wumengshan National Nature Reserve, 28°03′N, 104°20′E, altitude 1500 m, on the fallen branch of angiosperm, leg. C.L. Zhao, 21 September 2023, CLZhao 33798 (SWFC).

Etymology

grandinea (Latin or Greek origin): referring to the grandinioid hymenial surface.

Basidiomata

Annual, resupinate, closely adnate, membranaceous, very hard to separate from substrate, without odour or taste when fresh, becoming coriaceous upon drying, up to 20 cm long, 3 cm wide, 50–100 µm thick. Hymenial surface grandinioid, white to buff when fresh, turning to buff to slightly yellowish upon drying. Sterile margin cream to buff, thinning out, up to 1 mm wide.

Figure 10. 

Basidiomata of Alloexidiopsis grandinea in general and detailed views (CLZhao 33798, holotype). Scale bars: 1 cm (A); 1 mm (B).

Hyphal system

Monomitic, generative hyphae with clamp connections, colourless, thin-walled, rarely branched, interwoven, 2–4 µm in diameter; IKI–, CB–, tissues unchanged in KOH. Hymenium. Cystidia numerous, thin-walled, fusiform with an acute apex, occasionally sinuous in the basal, 20–42.5 × 5.5–9.5 µm, with a clamp connection at base; cystidioles absent. Hyphidia arising from generative hyphae, nodulose, frequently branched, colourless, thin-walled, 2–5 µm in diameter. Basidia ellipsoid to ovoid, longitudinally septate, two to four-celled, 12.5–14.5 × 9–11.5 µm; basidioles dominant, similar to basidia in shape, but slightly smaller. Basidiospores. Cylindrical to allantoid, slightly curved, colourless, smooth, thin-walled, with 1–2 oil drops, IKI–, CB–, (9.5–)10–12.3(–12.5) × (4.8–)5–5.8(–6) µm, L = 11.08 µm, W = 5.38 µm, Q = 1.95–2.20, Qm = 2.06 ± 0.04 (n = 60/2).

Figure 11. 

Sections of hymenium of Alloexidiopsis grandinea (holotype, CLZhao 33798) A basidiospores B basidia C cystidia D hyphidia. Scale bars: 10 µm (A); 20 µm (B–D); 10 × 100 Oil.

Additional specimen examined

China • Yunnan Province, Diqing, Weixi County, Weiden Town, Fuchuan Village, 27°06′N, 99°10′E, altitude 2900 m, on the fallen branch of angiosperm, leg. C.L. Zhao, 12 October 2023, CLZhao 34279 (SWFC).

Figure 12. 

Microscopic structures of Alloexidiopsis grandinea (holotype, CLZhao 33798) A basidiospores B basidia C basidioles D cystidia E hyphidia F part of the vertical section of hymenium. Scale bars: 10 µm (A–F).

Alloexidiopsis xantha J.H. Dong & C.L. Zhao, sp. nov.

MycoBank No: 854172
Figs 13, 14, 15

Diagnosis

Differs from other Alloexidiopsis species by its coriaceous basidiomata with cream to buff to yellow, smooth, slightly cracked hymenial surface, a monomitic hyphal system with clamped generative hyphae and allantoid to sickle-shaped basidiospores measuring 20–24 × 5–6.2 µm.

Holotype

China • Yunnan Province, Lincang, Yun County, Dumu Village, 24°32′N, 100°23′E, altitude 2100 m, on the fallen branch of angiosperm, leg. C.L. Zhao, 20 October 2022, CLZhao 25093 (SWFC).

Etymology

xantha (Latin or Greek origin): referring to the buff to yellow hymenial surface of the type specimen.

Basidiomata

Annual, resupinate, closely adnate, coriaceous, very hard to separate from substrate, without odour or taste when fresh, becoming leathery upon drying, up to 10 cm long, 2 cm wide, 200–300 µm thick. Hymenial surface smooth, slightly cracked, cream when fresh, turning to cream to buff to yellow upon drying. Sterile margin cream, thinning out, up to 1 mm wide.

Figure 13. 

Basidiomata of Alloexidiopsis xantha in general and detailed views (CLZhao 25093, holotype). Scale bars: 1 cm (A); 1 mm (B).

Hyphal system

Monomitic, generative hyphae with clamp connections, colourless, thin- to thick walled, branched, interwoven, 2.5–3.5 µm in diameter; IKI–, CB–, tissues unchanged in KOH. Hymenium. Cystidia numerous, thin-walled, subcylindrical to subconiform with an obtuse apex, 12.5–17.5 × 3.5–6 µm, with a clamp connection at base; cystidioles absent. Hyphidia arising from generative hyphae, nodulose, frequently branched, colourless, thin-walled, 2.5–4 µm in diameter. Basidia ellipsoid to ovoid, obconical, longitudinally septate, two to four-celled, 18–20.5 × 12–15.5 µm; basidioles dominant, similar to basidia in shape, but slightly smaller. Basidiospores. Allantoid, curved, sickle-shaped, colourless, smooth, thin-walled, IKI–, CB–, (18.5–)20–24(–24.5) × 5–6.2(–6.5) µm, L = 21.66 µm, W = 5.63 µm, Q = 3.60–4.05, Qm = 3.85 ± 0.10 (n = 30/1).

Figure 14. 

Sections of hymenium of Alloexidiopsis xantha (holotype, CLZhao 25093) A basidiospores B basidia C hyphidia D cystidia. Scale bars: 20 µm (A–D); 10 × 100 Oil.

Discussion

In the present study, four new species Ad. albomarginata, Ad. punctata, Al. grandinea and Al. xantha are described, based on the phylogenetic analyses and morphological characteristics.

Figure 15. 

Microscopic structures of Alloexidiopsis xantha (holotype, CLZhao 25093) A basidiospores B basidioles C cystidia D basidia E hyphidia F part of the vertical section of hymenium. Scale bars: 10 µm (A–F).

The corticioid species of the order Auriculariales are traditionally placed in Eichleriella, Exidiopsis and Heterochaete according to the morphological characteristics (Liu et al. 2022). On the basis of the erection of six new genera as Adustochaete, Alloexidiopsis, Amphistereum, Crystallodon, Proterochaete and Sclerotrema, they were placed in the corticioid species and three previously known genera were reinstated, for example, Hirneolina, Heteroradulum and Tremellochaete (Malysheva and Spirin 2017; Alvarenga et al. 2019; Alvarenga and Gibertoni 2021; Liu et al. 2022). A multilocus-based phylogeny with a wider sampling of various morphological groups in Auriculariales is urgently needed to achieve a more natural classification of this order, as in other orders within Agaricomycetes (Wang et al. 2021).

Phylogenetically, based on the combined ITS+nLSU sequence data (Fig. 1), it demonstrated that the four new species were nested in the genera Adustochaete and Alloexidiopsis within the order Auriculariales. Based on ITS topology tree (Fig. 2), Ad. albomarginata was retrieved as a sister to Ad. yunnanensis and the species Ad. punctata was sister to Ad. rava. However, Ad. yunnanensis differs from Ad. albomarginata by its grandinioid hymenial surface, longer basidia (25–47.5 × 8.5–14 µm) and smaller cystidia (17.5–24.5 × 3.5–5.8 µm; Li and Zhao (2022)). Ad. rava can be distinguished from Ad. punctata by its spined, sharp-tipped hymenial surface, smaller basidia (14.9–16.2 × 9.7–10.1 μm) and basidiospores (10.2–13.6 × 4.6–5.9 µm; Hyde et al. (2020)). Based on ITS topology tree (Fig. 3), Al. grandinea was retrieved as a sister to Al. schistacea and Al. xantha formed a monophyletic lineage. However, Al. schistacea differs from Al. grandinea by its smooth hymenial surface and longer basidia (15–20 × 7–10 µm; Liu et al. (2022)).

Morphologically, two new species Adustochaete albomarginata and Ad. punctata resemble four similar species in the genus Adustochaete, Ad. interrupta Spirin & Malysheva, Ad. nivea, Ad. rava and Ad. yunnanensis. A morphological comparison between two new Adustochaete species and four similar species are presented in Table 2. Two new species Al. grandinea and Al. xantha are similar to five species in the genus Alloexidiopsis, Al. australiensis S.L. Liu, Z.Q. Shen & L.W. Zhou, Al. calcea (Pers.) L.W. Zhou & S.L. Liu, Al. nivea (J.J. Li & C.L. Zhao) L.W. Zhou & S.L. Liu, Al. schistacea and Al. yunnanensis. A morphological comparison between two new Alloexidiopsis species and six similar species are presented in Table 3.

Table 2.
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A morphological comparison between two new Adustochaete species and four similar species in the genus Adustochaete.

Species name Hymenial surface Hyphae Cystidia Basidia Basidiospores References
Adustochaete albomarginata Smooth/ Cream to buff Thin-walled, unbranched Subclavate to fusiform; 23.5–48.5 × 10–13.5 µm Ellipsoid to ovoid, two to four-celled; 17–24.5 × 11–16.5 µm Subcylindrical to allantoid; 12–17.5 × 6.5–9 µm Present study
Adustochaete interrupta Smooth/ Light ochraceous-grey to brownish Thin-walled Clavate to fusiform; 45–96 × 6–13.5 µm Narrowly ovoid to obconical, four-celled; 15.1–24 × 9.1–11.8 µm Broadly cylindrical; 11.3–14.3 × 5.7–6.2 µm Alvarenga et al. (2019)
Adustochaete nivea Sharp-tipped spines/ White Thin-walled — Narrowly ovoid to obconical, four-celled; 14.9–16.2 × 9.7–10.1 μm Cylindrical; 10.2–13.6 × 4.6–5.9 µm Hyde et al. (2020)
Adustochaete punctata Smooth/ Punctate, white to cream Thin-walled, unbranched Subcylindrical to clavate; 15.5–23.5 × 5.5–7.5 µm Ellipsoid to ovoid, two to four-celled; 17–25 × 16.5–21 µm Subcylindrical to allantoid; 13.5–18 × 6–8.2 µm Present study
Adustochaete rava Sharp-tipped spines/ Pale to dark grey Thin-walled Clavate to fusiform; 27–52 × 4–8 µm Narrowly ovoid to obconical, four-celled; 10.8–15.2 × 7.3–10 µm Cylindrical; 10.2–13.7 × 3.8–4.7 µm Alvarenga et al. (2019)
Adustochaete yunnanensis Grandinioid/ Dark greyish to brownish Thin-walled, branched Clavate to fusiform; 17.5–24.5 × 3.5–5.8 µm Narrowly ovoid to obconical, four-celled; 25–47.5 × 8.5–14 µm Narrow cylindrical to allantoid; 12–20 × 5–7 µm Li and Zhao (2022)
Table 3.
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A morphological comparison between two new Alloexidiopsis species and six similar species in the genus Alloexidiopsis.

Species name Hymenial surface Hyphae Cystidia Basidia Basidiospores References
Alloexidiopsis australiensis Smooth, covered by sterile spines/ Cream to pale orange Thin-walled, branched Cylindrical, ventricose; 21.5–24.5 × 9.5–12 µm Ellipsoid to ovoid, four-celled; 18–21 × 13–18 µm Cylindrical to broadly cylindrical; 13–25 × 7–11 µm Li et al. (2022b)
Alloexidiopsis calcea Granulose to pruinose/ Greyish-white to light ochraceous Thin-walled, branched — Obovate to clavate, two to four-celled; 14–25 × 9.5–15 µm Allantoid to cylindrical, sometimes helicoid; 12–18 × 5–7 µm Wells (1961)
Alloexidiopsis grandinea Grandinioid/ Buff to slightly yellowish Thin-walled, branched Fusiform; 20–42.5 × 5.5–9.5 µm Ellipsoid to ovoid, two to four-celled; 12.5–14.5 × 9–11.5 µm Cylindrical to allantoid; 10–12.3 × 5–5.8 µm Present study
Alloexidiopsis nivea Smooth/ White to slightly cream Thin-walled, unbranched Tubular; 15–34 × 2.5–7 µm Narrowly ovoid to obconical, two to four-celled; 9–19 × 8–15 µm Allantoid; 6.5–13.5 × 2.7–5.5 µm Li et al. (2022a)
Alloexidiopsis schistacea Smooth/ Greyish Thin-walled, branched Cylindrical; 25–50 × 4–6 µm Ellipsoid to ovoid, four-celled; 15–20 × 7–10 µm Cylindrical to broadly cylindrical; 9.5–11 × 4.5–5.5 µm Liu et al. (2022)
Alloexidiopsis sinensis Grandinoid/ Yellowish-brown to rose to slightly purple Thin- to thick-walled, branched Cylindrical; 11.5–15.5 × 3–5.5 µm Ellipsoid to ovoid, two to four-celled; 16–22 × 7.5–10 µm Allantoid; 14.5–23 × 4.5–6.5 µm Dong et al. (2024)
Alloexidiopsis xantha Smooth/ Cream to slightly buff Thin- to thick walled, branched Subcylindrical to subconiform; 12.5–17.5 × 3.5–6 µm Ellipsoid to ovoid, obconical; 18–20.5 × 12–15.5 µm Allantoid, sickle-shaped; 20–24 × 5–6.2 µm Present study
Alloexidiopsis yunnanensis Odontoid/ White to smoke grey Thin-walled, unbranched Clavate to fusiform; 13–35 × 2–6 µm Narrowly ovoid to obconical, two to three-celled; 28–41 × 9–14 µm Cylindrical; 17–24 × 5–8 µm Guan et al. (2020)

In the ecological distribution, both genera species are not an extensively studied group, distributed worldwide and mainly found on hardwood (Alvarenga et al. 2019; Liu et al. 2022). The species of Adustochaete interrupta Spirin & Malysheva was found in Mexico, Ad. nivea was described in Brazil, Ad. rava was found in Brazil and Ad. yunnanensis was found in China. The species of Alloexidiopsis australiensis was found in Australia, Al. calcea was found in Germany and Al. nivea, Al. schistacea, Al. sinensis and Al. yunnanensis were found in China.

Fungi are one of the most diverse groups of organisms on Earth and play a crucial role in ecosystem processes and functions (Hyde 2022). New DNA sequencing techniques have revolutionised the studies of fungal taxonomy and diversity, in which about 150 k species of fungi have been described (Hyde 2022). In recent years, the wood-inhabiting fungi are an extensively studied group of Basidiomycota, which includes a number of poroid, smooth, grandinoid, odontioid and hydnoid basidiomata in China (Wu et al. 2022a, 2022b; Dong et al. 2023a, 2023b; Guan et al. 2023; Liu et al. 2023; Mao et al. 2023; Yang et al. 2023, 2024; Deng et al. 2024; Li et al. 2024; Luo et al. 2024; Zhang et al. 2024; Zhao et al. 2024; Zhou et al. 2024). In the past several years, many corticioid species have been reported and described in the order Auriculariales (Malysheva and Spirin 2017; Alvarenga et al. 2019; Spirin et al. 2019a, 2019a; Alvarenga and Gibertoni 2021; Li et al. 2022a, 2022b; Li and Zhao 2022; Liu et al. 2022), but many new taxa have not yet been discovered. Thus, the corticioid species diversity of the order Auriculariales is still not well known in China, especially in the subtropical and tropical areas. In the present study, four new species, Ad. albomarginata, Ad. punctata, Al. grandinea and Al. xantha were found and reported. This paper enriches our knowledge of fungal diversity in the order Auriculariales. We anticipate that more undescribed corticioid taxa will be discovered throughout China after extensive collection combined with morphological and molecular analyses.

Key to the known species of Adustochaete worldwide

1 Hymenial surface smooth 2
– Hymenial surface grandinioid 4
2 Basidia > 16.5 µm wide Adustochaete punctata
– Basidia < 16.5 µm wide 3
3 Basidiospores > 6.5 µm wide Adustochaete albomarginata
– Basidiospores < 6.5 µm wide Adustochaete interrupta
4 Cystidia absent Adustochaete nivea
– Cystidia present 5
5 Basidiospores > 5 µm wide, basidia > 16 µm long Adustochaete yunnanensis
– Basidiospores < 5 µm wide, basidia < 16 µm long Adustochaete rava

Key to the known species of Alloexidiopsis worldwide

1 Basidiospores allantoid 2
– Basidiospores cylindrical 6
2 Hymenial surface smooth 3
– Hymenial surface grandinoid, granulose to pruinose 4
3 Basidiospores > 13.5 µm long, cystidia subcylindrical to subconiform Alloexidiopsis xantha
– Basidiospores < 13.5 µm long, cystidia tubular Alloexidiopsis nivea
4 Cystidia absent Alloexidiopsis calcea
– Cystidia present 5
5 Cystidia > 5.5 µm wide Alloexidiopsis grandinea
– Cystidia < 5.5 µm wide Alloexidiopsis sinensis
6 Basidia > 28 µm long, cystidia clavate to fusiform Alloexidiopsis yunnanensis
– Basidia < 28 µm long, cystidia cylindrical 7
7 Basidiospores > 11 µm long, cystidia < 25 µm long Alloexidiopsis australiensis
– Basidiospores < 11 µm long, cystidia > 25 µm long Alloexidiopsis schistacea

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

The research was supported by the National Natural Science Foundation of China (Project Nos. 32170004, U2102220), Forestry Innovation Programs of Southwest Forestry University (Grant No: LXXK-2023Z07), the High-level Talents Program of Yunnan Province (YNQR-QNRC-2018-111) and the Research Project of Key Laboratory of Forest Disaster Warning and Control in Universities of Yunnan Province (ZKJS-S-202208).

Author contributions

Conceptualisation, CZ and JD; methodology, CZ and JD; software, CZ, JD and YZ; validation, CZ and JD; formal analysis, CZ and J JD; investigation, CZ and JD; resources CZ; writing – original draft preparation, CZ, JD, YZ and CQ; writing – review and editing, CZ and JD; visualisation, CZ and JD; supervision, CZ and JD; project administration, CZ; funding acquisition, CZ. All authors have read and agreed to the published version of the manuscript.

Author ORCIDs

Junhong Dong https://orcid.org/0000-0001-8740-0805

Yonggao Zhu https://orcid.org/0009-0008-5341-3796

Chengbin Qian https://orcid.org/0009-0003-5329-1016

Changlin Zhao https://orcid.org/0000-0002-8668-1075

Data availability

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

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