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Research Article
Four new species of Erioscyphella (Leotiomycetes, Helotiales) from southwestern China
expand article infoLe Luo‡§|, Kandawatte W. Thilini Chethana§, Qi Zhao‡, Hong-Li Su‡§, Cui-Jin-Yi Li§‡, Vinodhini Thiyagaraja‡, Fatimah Al-Otibi, Kevin D. Hyde‡§
‡ Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, China
§ Mae Fah Luang University, Chiang Rai, Thailand
| Guiyang Institute of Humanities and Technology, Guiyang, Thailand
¶ King Saud University, Riyadh, Saudi Arabia

Corresponding author: Kandawatte W. Thilini Chethana ( kandawatte.thi@mfu.ac.th )

Corresponding author: Kevin D. Hyde ( kdhyde3@gmail.com )

Academic editor: Danushka Sandaruwan Tennakoon
© 2025 Le Luo, Kandawatte W. Thilini Chethana, Qi Zhao, Hong-Li Su, Cui-Jin-Yi Li, Vinodhini Thiyagaraja, Fatimah Al-Otibi, Kevin D. Hyde.
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: Luo L, Thilini Chethana KW, Zhao Q, Su H-L, Li C-J-Y, Thiyagaraja V, Al-Otibi F, Hyde KD (2025) Four new species of Erioscyphella (Leotiomycetes, Helotiales) from southwestern China. MycoKeys 114: 29-48. https://doi.org/10.3897/mycokeys.114.138647
Open Access

Abstract

Erioscyphella is found across various regions and is part of the family Lachnaceae (Helotiales). It is distinguished by its white to orange disc-shaped apothecia, white to brown receptacles, and granulated hairs that contain amorphous or resinous material. These hairs lack swelling apices and crystals. Additionally, this genus is unique for its long ascospores. In the present study, we collected eight specimens from southwestern China. Morphological and phylogenetic analyses based on the combined LSU, ITS, mtSSU and RPB2 dataset showed that our specimens represent four new species of Erioscyphella, including E. ailaoensis, E. baimana, E. gelangheica and E. tengyueica. Here, we provide complete morphological descriptions with illustrations and sequence data essential for future taxonomic and evolutionary research.

Key words:

4 novel species, Lachnaceae, morphology, phylogeny, taxonomy

Introduction

The monophyletic genus Erioscyphella belongs to the family Lachnaceae (Dennis 1954; Spooner 1987; Cantrell and Haines 1997; Guatimosim et al. 2016; Tochihara and Hosoya 2022; Wijayawardene et al. 2020, 2022; Hyde et al. 2024) and includes 22 records (Index Fungorum 2024). The original description of Erioscyphella by Kirschstein (1939), which lacked typification, was inaccurately characterized based on traits that lack taxonomic significance, such as filiform, colored, and pigmented ascospores, as well as lanceolate paraphyses (Korf 1978; Perić and Baral 2014). It was not until Haines (1984) selected E. longispora, under which Peziza abnormis was later synonymized, as the lectotype of Erioscyphella that a more accurate characterization was established. These features are now recognized as insufficient for distinguishing this genus from related taxa. This genus was initially confused with the closely related genus Lachnum, but detailed morphological and molecular phylogenetic studies based on LSU, ITS, mtSSU and RPB2 data have since clarified their distinct characteristics and proposed a new concept based on the examination of Japanese materials (Tochihara and Hosoya 2022). Erioscyphella is characterized by scattered and cupulate apothecia on leaves or bamboo sheaths, straight or irregularly curved, septate and granulated hairs, mostly covered by apical amorphous materials or resinous material, lanceolate or filiform paraphyses, 8-spored asci with an amyloid apical pore and fusiform to long needle-like ascospores (Kirschstein 1939; Perić and Baral 2014; Tochihara and Hosoya 2022). There are no reports of asexual morph in this genus.

The species of Erioscyphella are primarily distributed in temperate and tropical regions, commonly inhabiting decaying wood and plant debris. Erioscyphella species are distributed mainly in China and a few other countries; E. curvispora is described from Montenegro (Perić and Baral 2014), E. griseibambusicola, E. latispora, E. lunata, E. lushanensis and E. subinsulae are collected from China (Guatimosim et al. 2016; Tello and Baral 2016; Li et al. 2022; Su et al. 2023); E. euterpes collected from Puerto Rico (Guatimosim et al. 2016). In contrast to the above-mentioned species that show a narrower distribution, E. abnormis shows a worldwide distribution, especially endemic in tropical regions (Tello and Baral 2016). They play a crucial role in breaking down organic matter, thereby aiding in nutrient cycling within their ecosystems. The ecological role of Erioscyphella as decomposers underscores their importance in forest ecosystems (Tochihara and Hosoya 2022; Niego et al. 2023a, 2023b). Ongoing research aims to better define species boundaries and elucidate the phylogenetic relationships within the genus through morphological studies and DNA sequencing. This work aids in the precise classification and discovery of new species.

During the investigation of Leotiomycetes in southwest China (Li et al. 2022, 2024a, 2024b; Su et al. 2022, 2023; Luo et al. 2024; Thiyagaraja et al. 2024), eight collections of Erioscyphella were obtained. We used morphological and phylogenetic analyses based on LSU, ITS, mtSSU and RPB2 data to confirm that these eight collections differ from all known species of Erioscyphella. We introduce four species to accommodate these collections. Here, we provide complete morphologies, illustrations, and their phylogenetic relationships for future taxonomic and evolutionary studies.

Material and methods

Specimen collection and morphological examination

We collected eight specimens from southwest China. All samples were collected from highly humid, natural broadleaf forests and protected areas with minimal human access. Altitudes were determined by the GPS device. The fruiting bodies were discovered on the surface of extremely wet, decaying wood litter. The samples were dehydrated in a dehydrator at a temperature range of 25–30 °C. After studying the morphology of the specimens and getting their genomic DNA, they were deposited at the Cryptogamic Herbarium of the Kunming Institute of Botany, Chinese Academy of Sciences (KUN-HKAS). Facesoffungi and Index Fungorum numbers were obtained as in Jayasiri et al. (2015) and Index Fungorum (2024), respectively. All the species identifications followed Chethana et al. (2021). The morphological descriptions were submitted to the Greater Mekong Subregion database (Chaiwan et al. 2021). The dried specimens were examined with a stereomicroscope (C-PSN, Nikon, Japan) and were captured with a digital camera (Canon EOS 70D, Japan) connected to the stereomicroscope. Free-hand sections of the dried specimens were mounted in a drop of water for observing microscopic characteristics, such as apothecia, exciple, paraphyses, asci and ascospores, using a Nikon compound microscope (Nikon, Japan) equipped with a DS-Ri2 camera. In addition, the sections were pretreated with Melzer’s reagent for the Iodine test (MLZ) (Tochihara and Hosoya 2022). Microstructures were measured using the Tarosoft (R) Image Frame Work program v.0.97 (Tarosoft, Thailand). The obtained measurements were presented in the format of (a–) b–c(–d), where ‘a’ represented the minimum value, ‘d’ represented the maximum value, and the range ‘b–c’ reflected the 90% confidence interval. The x̄ indicated the average value of measurements. Ascospore measurements were given as [n/m/p], indicating that the n number of ascospores were measured from m ascomata of the p number of collections. Images used for figures were processed with Adobe Photoshop CS6 Extended version 13.0 × 64 (Adobe Systems, USA).

DNA extraction, PCR amplifications and sequencing

Genomic DNA was extracted from the dried apothecia (around 50–100 mg) using a TSP101 DNA extraction kit (TSINGKE, China). Following the latest studies (Tochihara and Hosoya 2022; Su et al. 2023; Luo et al. 2024), LSU, ITS, mtSSU and RPB2 were used for PCR amplification, using the primers LR0R/LR5 (Vilgalys and Hester 1990), ITS1F/ITS4 (White et al. 1990; Gardes and Bruns 1993), mrSSU1/mrSSU3R (Zoller et al. 1999) and fRPB2-5F/fRPB2-7cR (Liu et al. 1999), respectively. For LSU, ITS, mtSSU and RPB2, the total volume of PCR amplifications was 25 μL, comprising 12.5 μL 2 × PCR G013 Taq MasterMix with Dye (Applied Biological Materials, Canada), 1 μL of each primer (10 μM), 2 μL genomic DNA, and 8.5 μL of sterilized, distilled water. Amplifications of LSU, ITS and RPB2 were conducted under the following conditions: pre-denaturation at 95 °C for 5 min, followed by 35 cycles of denaturation at 95 °C for 20 sec, annealing at 56 °C (LSU)/53 °C (ITS and RPB2) for 10 sec, elongation at 72 °C for 20 sec, and final elongation at 72 °C for 7 min. For mtSSU, the total volume of PCR amplifications was 25 μL, which comprised 21 μL 1 × PCR TSE101 Mix (TSINGKE, China), 1 μL of each primer (10 μM), and 2 μL genomic DNA. Amplifications of mtSSU were conducted under the following conditions: initial denaturation at 98 °C for 3 min, followed by 40 cycles of denaturation at 98 °C for 1 min, annealing at 52 °C for 1 min, elongation at 72 °C for 1 min, and final elongation at 72 °C for 10 min. Gel electrophoresis with 1% TAE and TSJ003 GoldView nucleic acid dye (TSINGKE, China) was used to confirm the obtained PCR products. Finally, the PCR products were sequenced at the Tsingke Biotechnology Co., Ltd., Kunming, China. Newly produced sequences were deposited in the GenBank and the accession numbers were given in Table 1.

Table 1.
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Taxa included in the phylogenetic analyses and the GenBank accession numbers of LSU, ITS, mtSSU and RPB2 sequences.

Species Strain Gene accession No. References
ITS LSU mtSSU RPB2
Capitotricha bicolor TNS-F-65670 LC424834 LC424942 LC533244 LC425011 Tochihara and Hosoya (2022)
Capitotricha rubi TNS-F-65752 LC438560 LC438573 LC533243 LC440395 Tochihara and Hosoya (2022)
Erioscyphella abnormis TNS-F-16609 AB705234 LC533175 LC533256 LC533184 Tochihara and Hosoya (2022)
Erioscyphella abnormis TNSF38452 LC669457 LC533171 LC533262 LC533210 Tochihara and Hosoya (2022)
Erioscyphella abnormis TNS-F-80478 LC424837 LC424949 LC533283 - Tochihara and Hosoya (2019)
Erioscyphella abnormis TNS-F-46841 LC669474 LC533170 LC533279 LC533209 Tochihara and Hosoya (2022)
Erioscyphella ailaoensis HKAS135686 (T) PQ349783 PQ349775 PQ358800 PQ424108 This study
Erioscyphella ailaoensis HKAS135687 PQ349784 PQ349776 PQ358801 PQ424109 This study
Erioscyphella alba MFLU16-0614(T) MK584965 MK591990 - - Ekanayaka et al. (2019)
Erioscyphella aseptata MFLU16-0590(T) MK584957 MK591986 - MK388223 Ekanayaka et al. (2019)
Erioscyphella baimana HKAS135697 (T) PQ349785 PQ349777 PQ358802 PQ424110 This study
Erioscyphella baimana HKAS135696 PQ349786 PQ349778 PQ358803 PQ424111 This study
Erioscyphella boninensis TNS-F-26520(T) NR185389 LC533151 LC533254 LC533196 Tochihara and Hosoya (2022)
Erioscyphella brasiliensis MFLU16-0577b MK584967 MK591993 - - Ekanayaka et al. (2019)
Erioscyphella brasiliensis TNS-F-46419 LC669456 LC533133 LC533278 LC549672 Tochihara and Hosoya (2022)
Erioscyphella curvispora KL 381(T) MH190414 MH190415 - - Perić and Baral (2014)
Erioscyphella euterpes PR 147 U58640 - - - Cantrell and Haines (1997)
Erioscyphella fusiforme MFLU15-0230(T) MK584948 MK591975 - MK614728 Ekanayaka et al. (2019)
Erioscyphella gelangheica HKAS135689 (T) PQ349787 PQ349779 PQ358804 - This study
Erioscyphella gelangheica HKAS135695 PQ349788 PQ349780 PQ358805 - This study
Erioscyphella griseibambusicola HKAS124657 OP451797 OP451791 OP451844 OP432252 Su et al. (2023)
Erioscyphella griseibambusicola HKAS124656(T) OP451796 OP451790 OP451843 OP432251 Su et al. (2023)
Erioscyphella hainanensis TNS-F-35056 LC669465 LC533169 LC533275 LC533206 Tochihara and Hosoya (2022)
Erioscyphella hainanensis TNS-F-35049 LC669452 LC533168 LC533274 LC533205 Tochihara and Hosoya (2022)
Erioscyphella insulae TNS-F-26500 LC669448 LC533149 LC533252 LC533194 Tochihara and Hosoya (2022)
Erioscyphella insulae TNS-F-39720(T) LC669451 LC533177 LC533261 LC533207 Tochihara and Hosoya (2022)
Erioscyphella latispora HKAS124391 OP113849 OP113850 - OP715727 Li et al. (2022)
Erioscyphella latispora HKAS124389(T) OP310823 OP113844 - OP715728 Li et al. (2022)
Erioscyphella lunata JA-CUSSTA 8292 KX501132 KX501133 - - Tello and Baral (2016)
Erioscyphella lushanensis HMAS81575 JF937582 - - - Zhao and Zhuang (2011)
Erioscyphella otanii TNS-F-81472(T) NR185393 LC533179 LC533286 LC533226 Tochihara and Hosoya (2022)
Erioscyphella papillaris TNS-F-81272(T) NR185391 LC533161 LC533285 LC533204 Tochihara and Hosoya (2022)
Erioscyphella paralushanensis TNS-F-61920(T) NR185390 LC533141 LC533267 LC533220 Tochihara and Hosoya (2022)
Erioscyphella sasibrevispora TNS-F-80399 LC669470 LC533173 LC533268 LC533216 Tochihara and Hosoya (2022)
Erioscyphella sasibrevispora TNS-F-81401(T) LC669472 LC533174 LC533269 LC533217 Tochihara and Hosoya (2022)
Erioscyphella sclerotii TNS-F-26492 LC669438 LC533152 LC533255 LC533197 Tochihara and Hosoya (2022)
Erioscyphella sclerotii TNS-F-38480 LC669458 LC533134 LC533263 LC549673 Tochihara and Hosoya (2022)
Erioscyphella sclerotii MFLU 16-0569 MK584951 MK591980 - - Ekanayaka et al. (2019)
Erioscyphella sclerotii MFLU 18-0688 MK584969 MK591995 - - Ekanayaka et al. (2019)
Erioscyphella sinensis TNS-F-32161 LC669449 LC533167 LC533273 LC533219 Tochihara and Hosoya (2022)
Erioscyphella sinensis TNS-F-16838 AB481280 LC533164 LC533235 AB481364 Tochihara and Hosoya (2022)
Erioscyphella subinsulae HKAS 124659 OP451799 OP451793 OP451846 OP432254 Su et al. (2023)
Erioscyphella subinsulae HKAS 124660 OP451800 OP451794 OP451847 OP432255 Su et al. (2023)
Erioscyphella subinsulae HKAS 124661 OP451801 OP451795 OP451848 OP432256 Su et al. (2023)
Erioscyphella subinsulae HKAS 124658(T) OP451798 OP451792 OP451845 OP432253 Su et al. (2023)
Erioscyphella tengyueica HKAS135688 (T) PQ349789 PQ349781 PQ358806 - This study
Erioscyphella tengyueica HKAS135693 PQ349790 PQ349782 PQ358807 - This study
Lachnellula calyciformis TNS-F-81248 LC438561 LC438574 LC533247 LC438590 Tochihara and Hosoya (2022)
Lachnellula suecica TNS-F-16529 AB481248 LC424944 LC533231 AB481341 Tochihara and Hosoya (2022)
Neodasyscypha cerina TNS-F-65625 LC424836 LC424948 LC533242 LC425013 Tochihara and Hosoya (2022)

*Names in indicate the specimens from the current study. Names with (T) indicate type specimens and - denotes unavailable data in the GenBank.

Phylogenetic analyses

New DNA sequences generated from forward and reverse primers were assembled using BioEdit v.7.2.5 (Hall 1999) to obtain consensus sequences. The concatenated sequences were used to search for the closer relatives in the NCBI (Johnson et al. 2008). According to the close relatives and recent studies, the newly generated sequences and some published sequences were used for the phylogenetic analyses (Table 1). Capitotricha bicolor (TNS-F-65670), C. rubi (TNS-F-65752), Lachnellula calyciformis (TNS-F-81248), L. suecica (TNS-F-16529) and Neodasyscypha cerina (TNS-F-65625) were used as the outgroup taxa. The phylogenetic analysis was conducted based on the datasets including reference DNA sequences and newly generated DNA sequences using OFPT (Zeng et al. 2023) with the following protocol. Datasets of each gene region were first independently aligned with ‘auto’ strategy (based on data size) by MAFFT (Katoh and Standley 2013) and trimmed with ‘gappyout’ method (based on gaps’ distribution) by TrimAl (Capella-Gutiérrez et al. 2009). The best-fit nucleotide substitution model for each dataset was then selected based on the Bayesian information criterion (BIC) from twenty-two common DNA substitution models with rate heterogeneity by ModelFinder (Kalyaanamoorthy et al. 2017). Afterwards, all datasets were concatenated with partition information for the subsequent phylogenetic analyses. Maximum likelihood with 1000 replicates was performed using ultrafast bootstrap approximation (Hoang et al. 2018) with SH-like approximate likelihood ratio test (SH-aLRT) (Guindon et al. 2010) in IQ-TREE (Nguyen et al. 2015). The consensus tree was summarized based on the extended majority rule. Bayesian inference (BI) analyses were run in the CIPRES Science Gateway v.3.3 (Miller et al. 2010). The best-fit nucleotide substitution models were determined using jModelTest2 on XSEDE (2.1.6). The BI was performed in MrBayes on XSEDE v. 3.2.7a (Ronquist et al. 2012), with four simultaneous Markov chain Monte Carlo (MCMC) chains and four runs for 3,000,000 generations, with trees sampled at each 300th generation. The first 25% of trees were discarded as burn-in, and BI posterior probabilities (PP) were conducted from the remaining trees. The consensus phylograms were visualized on FigTree v. 1.4.4, and edited with Adobe Illustrator CC 2019, Adobe Systems (USA). Decisions as to whether species are new followed the polyphasic approach as recommended by Chethana et al. (2021) and Maharachchikumbura et al. (2021).

Results

Phylogenetic analysis

The phylogenetic analyses were based on 45 Erioscyphella taxa, including C. rubi (TNS-F-65752), C. bicolor (TNS-F-65670), L. calyciformis (TNS-F-81248), L. suecica (TNS-F-16529) and N. cerina (TNS-F- 65625) as the outgroup taxa. The alignment comprised 4 partitions and 3944 total sites (ITS: 758 bp; LSU: 1105 bp; mtSSU: 947 bp; RPB2: 1134 bp), with 14.359% gaps and completely undetermined characters. The ML tree has the same topology as the BI tree. The best ML tree with a final optimization likelihood of -25316.078809 is displayed in Fig. 1. In the BI analyses, the final average standard deviation of split frequencies was 0.007666, which revealed convergence. In the multi-gene phylogenetic tree based on the combined ITS, LSU, mtSSU and RPB2 dataset, all taxa of Erioscyphella clustered together. Erioscyphella ailaoensis was sister to the clade comprising E. sclerotii and E. abnormis, with 73% maximum likelihood bootstrap (MLBP) and 0.89 Bayesian posterior probabilities (BPP) support. Erioscyphella gelangheica was sister to the clade comprising E. sclerotii and E. abnormis, E. ailaoensis, E. brasiliensis and E. aseptata with 78% MLBP and 0.88 BPP support. Erioscyphella baimana was sister to E. latispora with 99% MLBP and 1.00 BPP support. Besides, E. tengyueica was sister to E. papillaris with 91% MLBP and 0.98 BPP. The phylogenetic result showed that Erioscyphella species clustered together, similar to previously published studies (Su et al. 2023).

Figure 1. 

The Maximum Likelihood tree based on the combined LSU, ITS, mtSSU and RPB2 sequence data for Erioscyphella. Capitotricha bicolor (TNS-F-65670), C. rubi (TNS-F-65752), Lachnellula calyciformis (TNS-F-81248), L. suecica (TNS-F-16529) and Neodasyscypha cerina (TNS-F-65625) are used as the outgroup taxa. The MLBP ≥ 70% and BPP ≥ 0.90 are shown at the nodes as MLBP/BPP. MLBS < 70% and BPP < 0.90 are expressed as a hyphen (“-”). Names with (T) indicate type specimens. Names in red indicate new species.

Taxonomy

Erioscyphella ailaoensis L. Luo, K.D. Hyde & H.L. Su, sp. nov.

Index Fungorum: IF902529
Facesoffungi Number: FoF16389
Fig. 2

Etymology.

The epithet “ailaoensis” refers to the collection site, Ailao Mountain, where the holotype specimen was collected.

Holotype.

HKAS135686.

Description.

Saprobic on the dead bark. Sexual morph: Apothecia scattered to partly gregarious, superficial, 1–2.4 mm in diameter, 0.4–1.4 mm high when dry, discoid to cupulate, shortly stipitate, externally covered with short, white to brown hairs. Disc concave, surface slightly rough, yellow to brown. Margin flat to slightly involute, pale yellow, covered with yellow to pale brown hairs. Receptacle discoid to cupulate, yellow to pale brown, clothed entirely with short, yellow to pale brown hairs. Stipe 0.2–1 mm in diameter, 0.2–0.6 mm long when dry, cylindrical, solitary, yellow to pale brown, clothed with yellow to pale brown hairs. Hairs 22–92 × 3.0–4.3 µm (x̄ = 51 × 3.7 µm, n = 30), clavate to cylindrical, straight to slightly curved, septate, hyaline, thick-walled, covered with hyaline granules, obtuse apex, apical amorphous or resinous material. Hymenium 120–230 µm (x̄ = 153 µm, n = 12), concave, surface slightly rough, light yellowish brown in dry. Medullary excipulum 40–90 µm (x̄ = 58 µm, n = 20), thin, hyaline to light yellow, thin-walled, smooth cells of textura oblita, 2.1–5.9 µm (x̄ = 3.7 µm, n = 50) in diameter. Ectal excipulum 55–80 µm (x̄ = 65 µm, n = 20), thin-walled, smooth, light yellowish cells of textura prismatica to globulosa, 2–4.9 µm (x̄ = 3.0 µm, n = 60) in diameter. Paraphyses 100–138 × 1.6–3.9 µm (x̄ = 117 × 2.8 µm, n = 25), longer than asci, filiform, straight to slightly curved, aseptate, hyaline, thin-walled, rough, with slightly acute apex. Asci 85–143 × (4.5–) 5.5–9.0(–9.5) µm (x̄ = 100 × 7.3 µm, n = 34), 8-spored, unitunicate, overlapping fascicles, clavate, straight to slightly curved, inoperculate, hyaline, apically thickened wall, laterally relatively thin, slightly smooth, with an apical, non-amyloid pore and tapered ends, J- in MLZ. Ascospores (50/14/2) (43.0–)45.5–97(–101.0) × 1.4–2.4(–2.6) µm, (x̄ = 68 × 1.9 µm), fascicled, filiform, multi-septate, thin-walled, hyaline, rough with taper, obtuse ends, without oil guttules, hyaline, slightly smooth. Asexual morph: Not observed.

Figure 2. 

Erioscyphella ailaoensis (HKAS135686, holotype) a–c dried ascomata on the bark d vertical section of an ascoma e excipulum f hairs g paraphyses h–k asci (j, k asci in MLZ) l–o ascospores (l–n ascospores in MLZ). Scale bars: 100 µm (d); 50 µm (e–k); 30 µm (l–o).

Material examined.

China • Yunnan Province, Puer City, Jingdong County, Ailao Mountain, altitude 2478 m, on the decayed unidentified bark, 8 June 2022, Hongli Su, SU872 (HKAS135686, holotype); China • Xizang Province, Shigatse City, altitude 1774 m, on the decayed unidentified twig, 6 July 2022, Hongli Su, SU1423 (HKAS135687, paratype).

Notes.

Our specimens, HKAS135686 and HKAS135687, were grouped as a distinct clade, separated from the clade comprising E. abnormis and E. sclerotii by 73% MLBS and 0.89 BIPP (Fig. 1). The new species exhibited morphological differences from E. abnormis and E. sclerotii by having J- apical pores, whereas the latter species have apical pores that are J- in MLZ. In contrast to the septate paraphyses of E. abnormis, E. ailaoensis has aseptate paraphyses (Han et al. 2021). Furthermore, asci and ascospores of E. ailaoensis are longer than those of E. sclerotii (Nagao 1996; Perić and Baral 2014). Therefore, E. ailaoensis is introduced here as a new species.

Erioscyphella baimana L. Luo, K.D. Hyde & Q. Zhao, sp. nov.

Index Fungorum: IF902530
Facesoffungi Number: FoF16391
Fig. 3

Etymology.

The epithet “baimana” refers to the collection site, Baima Mountain, where the holotype specimen was collected.

Holotype.

HKAS 135697.

Description.

Saprobic on dead twigs. Sexual morph: Apothecia superficial, gregarious, 0.3–1.1 mm in diameter, 0.3–1.4 mm high when dry, discoid to cupulate, long stipitate, externally covered with short, white hairs. Disc concave, surface slightly smooth, yellow. Margin flat to slightly involute, white, covered with white hairs. Receptacle cupulate to discoid, white, covered entirely with short, white hairs. Stipe 0.2–0.6 mm in diameter, 0.3–1.1 mm long when dry, cylindrical, solitary, white, clothed with white hairs. Hairs 30–120 × 2.8–4.7 µm (x̄ = 74 × 3.7 µm, n = 30), clavate to cylindrical, straight to slightly curved, aseptate, hyaline, thin-walled, covered with fine granules, obtuse apex, lacks apical amorphous. Hymenium 165–230 µm (x̄ = 195 µm, n = 12), concave, surface slightly smooth, yellow in dry. Medullary excipulum 35–120 µm (x̄ = 70 µm, n = 18), thin, hyaline, thin-walled cells of textura porrecta, 1.3–3.8 µm (x̄ = 2.5 µm, n = 50) in diameter. Ectal excipulum 40–120 µm (x̄ = 68 µm, n = 18) thin, thin-walled, smooth, light yellowish cells of textura porrecta to oblita, 1.9–6.3 µm (x̄ = 4.1 µm, n = 60) in diameter. Paraphyses 95–170 × 1.2–2.5 µm (x̄ = 140 × 1.6 µm, n = 25), longer than asci, filiform, straight to slightly curved, aseptate, hyaline, light smooth, with slightly obtuse apex. Asci 100–152 × 3.6–9.5 µm (x̄ = 123 × 7.2 µm, n = 34), 8-spored, unitunicate, clavate, straight to slightly curved, inoperculate, hyaline, slightly smooth, with an apical, amyloid pore and rounded ends, J+ in MLZ, tapered long stipitate base. Ascospores (85/7/2) (29.5–)30.5–37.5(–40.0) × 2.2–5.0 (–5.5) µm, (x̄ = 32.9 × 4.1 µm, n = 85), biseriate, fusoid-clavate with blunt ends, fusiform, 1–3-septate, thin-walled, hyaline, slightly smooth, tapering towards the obtuse ends, with longitudinal striations, without oil guttules. Asexual morph: Not observed.

Figure 3. 

Erioscyphella baimana (HKAS 135697, holotype) a–c dried ascomata on the twig d a vertical section of part of an ascoma e excipulum f, g hairs h paraphyses i–l asci (k, l asci in MLZ) m–r ascospores. Scale bars: 100 µm (d–f); 50 µm (g–l); 30 µm (m–r).

Material examined.

China • Yunnan Province, Diqing City, Deqin County, Baima Mountain, altitude 3485 m, on the decayed unidentified twig, 21 July 2022, Le Luo, Ly7 (HKAS 135697, holotype); • ibid., Le Luo, Ly26 (HKAS 135696, isotype).

Notes.

Our specimens, HKAS 135697 and HKAS 135696, were grouped in a distinct clade, separated from E. latispora by 99% MLBS and 1.00 BIPP (Fig. 1). Our species, E. baimana, morphologically differs from E. latispora by having aseptate hairs, ascospores without oil guttules whereas the latter species possess septate hairs, asci with rounded to subconical apex, and ascospores with four or more large guttules. Therefore, E. baimana is introduced here as a new species.

Erioscyphella gelangheica L. Luo, K.D. Hyde, H.L. Su & C.J.Y. Li, sp. nov.

Index Fungorum: IF902529
Facesoffungi Number: FoF16390
Fig. 4

Etymology.

The epithet “gelangheica” refers to the collection site Gelanghe township where the holotype specimen was collected.

Holotype.

HKAS135689.

Description.

Saprobic on dead bark. Sexual morph: Apothecia scattered to partly gregarious, superficial, 0.24–0.5 mm in diameter, 0.35–0.5 mm high when dry, discoid to cupulate, long stipitate, externally covered with short, white to yellowish hairs. Discs concave, surface slightly rough, white to yellow. Margin flat to slightly involute, white to pale yellow, covered with white to pale yellow hairs. Receptacle discoid to cupulate, white to pale brown, clothed entirely with short, white to slightly yellow hairs. Stipe 0.06–0.18 mm in diameter, 0.14–0.3 mm long when dry, cylindrical, solitary, white to pale yellow, clothed with white to pale yellow hairs. Hairs 28–117 × 1.6–3.9 µm (x̄ = 60 × 2.9 µm, n = 30), clavate to cylindrical, straight to slightly curved, septate, hyaline, thin-walled, covered with hyaline granules, obtuse apex. Hymenium 65–120 µm (x̄ = 85 µm, n = 12), concave, surface slightly rough, light yellow in dry. Medullary excipulum 23.5–65 µm (x̄ = 37 µm, n = 18), thin, hyaline to light yellow, thin-walled cells of textura intricata, 1.3–3.6 µm (x̄ = 2.2 µm, n = 50) in diameter. Ectal excipulum 20–95 µm (x̄ = 48 µm, n = 18) thick, thin-walled, smooth, hyaline cells of textura porrecta to textura globulosa, 1.6–4.2 µm (x̄ = 2.7 µm, n = 60). Paraphyses 25–68 × 0.9–2.0 µm (x̄ = 43 × 1.3 µm, n = 25), longer than asci, filiform, straight, aseptate, hyaline, thin-walled, rough, with slightly acute apex. Asci 35–58 × 2.2–3.60 µm (x̄ = 46 × 3.0 µm, n = 34), 8-spored, unitunicate, clavate, straight to slightly curved, inoperculate, hyaline, wall apically thickened, laterally relatively thin, slightly smooth, with an apical, amyloid pore and tapered ends, J+ in MLZ. Ascospores (85/6/2) 6.0–8.3 × 1.2–1.8 µm, (x̄ = 7.3 × 1.5 µm, n = 86), partially biseriate, filiform, aseptate, thin-walled, hyaline, rough with tapering towards the obtuse ends, partially oil guttules, subspherical, hyaline, slightly smooth. Asexual morph: Not observed.

Figure 4. 

Erioscyphella gelangheica (HKAS 135689, holotype) a–d dried ascomata on the host e a vertical section of an ascoma f excipulum g, h hairs i paraphyses and asci j–n asci (l–n asci in MLZ) o apices of asci treated with Melzer’s reagent p–u ascospores. Scale bars: 100 µm (e); 50 µm (f); 20 µm (g–n); 5 µm (o–u).

Material examined.

China • Yunnan Province, Xishuangbanna City, Menghai County, Gelanghe township, altitude 2097 m, on the decayed unidentified bark, 6 September 2022, Cuijinyi Li, LCJY1389 (HKAS 135689, holotype); • ibid., Hongli Su, SU1978 (HKAS 135695, paratype).

Notes.

Our specimens, HKAS 135689 and HKAS 135695, were grouped in a distinct clade, separated from the clade comprising E. abnormis, E. sclerotii, E. ailaoensis, E. brasiliensis and E. aseptata by 78% MLBS and 0.88 BIPP (Fig. 1). Erioscyphella gelangheica has shorter asci (35–58 µm vs. 41–104 µm), ascospores (6.0–8.3 µm vs. 39–81 µm), and paraphyses (25–68 µm vs. 52–123 µm) than those of E. abnormis (Perić and Baral 2014). Erioscyphella gelangheica differs from E. sclerotii by having long stipitate apothecia and aseptate ascospores, while E. sclerotii has short stipitate apothecia and 1–3-septate ascospores. Our species, E. gelangheica, has shorter asci (35–58 µm vs. 85–143 µm), ascospores (6.0–8.3 µm vs. 45.5–97 µm), and paraphyses (25–68 µm vs. 100–138 µm) than those of E. ailaoensis. Furthermore, E. ailaoensis has septate ascospores in contrast to the aseptate ascospores of E. gelangheica. Compared to long stipitate apothecia with white to pale yellow hairs and aseptate ascospores of E. gelangheica, E. brasiliensis has 0–1-septate ascospores and stipitate apothecia, with the stipe base often devoid of hairs and blue-black (Haines 1992). In addition to having aseptate ascospores, E. gelangheica has shorter asci (35–58 µm vs. 70–100 µm) and ascospores (6.0–8.3 µm vs. 28.5–45.6 µm) than those of E. aseptata, which has septate ascospores (Ekanayaka et al. 2019). Therefore, E. gelangheica is introduced here as a new species.

Erioscyphella tengyueica L. Luo, K.D. Hyde & C.J.Y. Li, sp. nov.

Index Fungorum: IF902531
Facesoffungi Number: FoF16392
Fig. 5

Etymology.

The epithet refers to the collection site of the type specimen.

Holotype.

HKAS 135688.

Description.

Saprobic on the dead twigs. Sexual morph: Apothecia superficial, scattered to partly gregarious, 0.16–0.68 mm in diameter, 0.3–0.7 mm high when dry, discoid to cupulate, shortly stipitate, externally covered with short, white hairs. Discs concave, surface slightly rough, white. Margin slightly involute, white, covered with white hairs. Receptacle cupulate, concolorous, clothed entirely with short, white hairs. Stipe 0.09–0.23 mm in diameter, 0.18–0.4 mm long when dry, cylindrical, solitary, concolorous with the receptacle, clothed with white hairs. Hairs 45–95 × 2.8–6.9 µm (x̄ = 68 × 4.8 µm, n = 10), clavate to cylindrical, straight or curved, septate, hyaline, thin-walled, less covered with hyaline granules, obtuse apex. Hymenium 65–115 µm (x̄ = 87 µm, n = 12), concave, surface slightly rough, light white in dry. Medullary excipulum 18–33 µm (x̄ = 26 µm, n = 18), thick, comprising hyaline, thin-walled, poorly developed cells of textura globulosa, 1.4–3.5 µm (x̄ = 2.4 µm, n = 50) in diameter. Ectal excipulum 11–30 µm (x̄ = 21 µm, n = 18) thick, comprising thick-walled, smooth, light yellowish cells of textura oblita to textura porrecta, 1.2–3.8 µm (x̄ = 2.2 µm, n = 60). Paraphyses 65–97 × 1.8–3.0 µm (x̄ = 86 × 2.5 µm, n = 25), longer than asci, filiform, straight, aseptate, hyaline, thin-walled, narrow lanceolate, smooth, less covered with hyaline granules, with slightly obtuse apex. Asci 60–80 × 6.0–9.3 µm (x̄ = 70 × 7.6 µm, n = 34), 8-spored, clavate, straight to slightly curved, inoperculate, hyaline, unitunicate, slightly smooth, with an apical, amyloid pore and rounded ends, croziers absent at the basal septum, J+ in MLZ. Ascospores (80/6/2) 25–31.5 × 1.6–5.5 µm, (x̄ = 27.7 × 3.3 µm, n = 80), overlapping biseriate, filiform, aseptate, thin-walled, hyaline, rough with tapering towards obtuse ends, filled with oil guttules or with 1–2- large oil guttules. Asexual morph: Not observed.

Figure 5. 

Erioscyphella tengyueica (HKAS 135688, holotype) a–c dried ascomata on the host d a vertical section of an ascoma e excipulum f paraphyses g–j asci (i, j asci in MLZ) k ascospores. Scale bars: 100 µm (d); 50 µm (e, f); 20 µm (g–j); 10 µm (k) .

Material examined.

China • Yunnan Province, Tengchong City, Tengyue Street, altitude 1983.3 m, on the decayed unidentified twig, 21 August 2022, Cuijinyi Li, LCJY1171 (HKAS 135688, holotype); • ibid., altitude 1774 m, on the decayed unidentified twig, 18 August 2022, Le Luo, Ly255 (HKAS 135693, paratype).

Notes.

Our specimens, HKAS 135688 and HKAS 135693, were grouped into a distinct clade, separated from E. papillaris (TNS-F-81272) by 91% MLBS and 0.98 BIPP (Fig. 1). Erioscyphella tengyueica differs from E. papillaris by having aseptate ascospores and aseptate paraphyses, while the latter has septate paraphyses and aseptate or one-septate (rarely two-septate) ascospores. Our species differs from E. otanii by having longer asci (60–80 µm vs. 34–38.8 µm), longer ascospores (34–38.8 µm vs. 12.3–14.6 µm) and aseptate paraphyses, in contrast to the septate paraphyses of E. otanii (Tochihara and Hosoya 2022). Therefore, E. tengyueica is introduced here as a new species.

Discussion

In China, the diversity of Leotiomycetes is substantial due to varied climates and ecosystems in the country (Li et al. 2022; Su et al. 2023; Guo et al. 2024; Luo et al. 2024; Zhang et al. 2024). Ongoing research continues to uncover new species and understand their roles in ecosystem functioning, highlighting the importance of preserving fungal diversity for ecological health and agricultural sustainability (Li et al. 2022; Su et al. 2023; Lestari and Chethana 2024; Luo et al. 2024).

Based on the previous research on the phylogeny, morphology and ecology of Lachnaceae (Hosoya et al. 2010), delimiting generic boundaries in Lachnaceae can no longer be defined by morphological characteristics alone (Li et al. 2022; Tochihara and Hosoya 2022; Su et al. 2023). The latest concept for Erioscyphella was proposed as the typical taxa characterized by the hair structures (the swollen apices, apical anamorph material/resinous material) and ascospore length (Tochihara and Hosoya 2022). This revision signifies a departure from the previously emphasized characteristics, highlighting the dynamic nature of fungal taxonomy (Tochihara and Hosoya 2022).

Despite advances in molecular techniques and the integration of morphological and ecological data, our analysis reveals persistent challenges in clarifying species boundaries. Particularly for paraphyletic members of long-spored Lachnum within Erioscyphella, the utilization of the UNITE Species Hypotheses (SH) system analysis based on ITS gene fragment, alongside traditional methods, has not provided definitive resolutions, indicating the ongoing complexity in species delineation (Tochihara and Hosoya 2022). Phylogenetic relationships in Erioscyphella gradually became clear as more fresh collections were identified with sequence data (Li et al. 2022; Su et al. 2023). Almost all species of Erioscyphella have stable phylogenetic positions with strong statistical support, except for E. bambusina, which lacks sequences in public databases.

Nearly all Erioscyphella species occupy stable phylogenetic positions supported by strong statistical evidence, except for E. bambusina, which lacks sequence data in public databases. We compared the morphology of E. bambusina with our newly established species (Dennis 1954; Su et al. 2023). Erioscyphella bambusina has septate, shorter ascospores, and septate paraphyses, in contrast to the multi-septate, longer ascospores, and aseptate paraphyses of E. ailaoensis. Erioscyphella gelangheica differs from E. bambusina by having smaller apothecia and aseptate paraphyses, while the latter has larger apothecia and septate paraphyses. Erioscyphella baimana has aseptate hairs and aseptate paraphyses, while E. tengyueica has white discs, and J- in MLZ with and without 3% KOH pretreatment. In contrast to these two, E. bambusina has cream to pale yellow discs, pore blued in MLZ, and septate paraphyses.

Most records of Erioscyphella are from the tropics (Spooner 1987; Haines and Dumont 1983; Tochihara and Hosoya 2022). Although sample collections have extended to subtropic, temperate and cold-temperate regions (Bien and Damm 2020; Tochihara and Hosoya 2022), collected samples are still scarce. Further, it was found that the lack of available sequences of these species led to Lachnaceae and Helotiaceae taxa exhibiting paraphyletic characters in the ITS-LSU phylogeny (Johnston et al. 2019; Quandt and Haelewaters 2021). Although Johnston et al. (2019) solved this issue based on phylogenetic analyses of up to 15 concatenated genes across 279 specimens, obtaining gene sequences for multiple loci for Lachnaceae taxa is still one of the critical problems.

Continued interdisciplinary research efforts are warranted to refine our understanding of fungal taxonomy within the Lachnaceae. Future studies should explore novel methodologies, such as high-throughput sequencing and ecological niche modeling, to elucidate species boundaries and evolutionary relationships more comprehensively. Additionally, comprehensive taxonomic revisions, including detailed examination of type specimens and expanded sampling, will be crucial for resolving taxonomic ambiguities and advancing fungal systematics.

In conclusion, our study contributes to the growing body of knowledge on fungal taxonomy and highlights the need for an integrated approach combining molecular, morphological, and ecological data to address the complexities inherent in delineating generic boundaries and species relationships within the Lachnaceae family.

Acknowledgements

Le luo thanks Dr Shaun Pennycook for his valuable assistance with the Latin binomial nomenclature.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by the Second Tibetan Plateau Scientific Expedition and Research (STEP) Program (Grant No. 2019QZKK0503) and the Chinese Research Fund (Project No. E1644111K1) under the initiative “Flexible Introduction of the High-Level Expert Program” at the Kunming Institute of Botany, Chinese Academy of Sciences. The authors extend their appreciation to the Researchers Supporting Project number (RSP2024R114), King Saud University, Riyadh, Saudi Arabia.

Author contributions

Supervision: KWTC. Writing - original draft: LL, FA. Writing - review and editing: QZ, KWTC, KDH. Collection: LL, HLS, CJYL, VL.

Author ORCIDs

Le Luo https://orcid.org/0009-0006-7213-0498

Kandawatte W. Thilini Chethana https://orcid.org/0000-0002-5816-9269

Qi Zhao https://orcid.org/0000-0001-8169-0573

Hong-Li Su https://orcid.org/0000-0001-9071-7635

Cui-Jin-Yi Li https://orcid.org/0000-0002-2805-7071

Vinodhini Thiyagaraja https://orcid.org/0000-0002-8091-4579

Fatimah Al-Otibi https://orcid.org/0000-0003-362

Kevin D. Hyde https://orcid.org/0000-0002-2191-0762

Data availability

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

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