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Research Article
Three new species of Peroneutypa (Diatrypaceae, Xylariales) and a first record of Eutypa camelliae in China with updated description
expand article infoXinying Mao‡, Kamran Habib§, Rizwana Zulfiqar|, Hongde Yang, Yingqian Kang‡
‡ Guizhou Medical University, Guiyang, China
§ Guizhou Medical University, Gui’an New District, China
| University of the Punjab, Lahore, Pakistan
¶ Qujing Normal University, Qujing, China

Corresponding author: Yingqian Kang ( joycekangtokyo@gmail.com )

Academic editor: Ning Jiang
© 2025 Xinying Mao, Kamran Habib, Rizwana Zulfiqar, Hongde Yang, Yingqian Kang.
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: Mao X, Habib K, Zulfiqar R, Yang H, Kang Y (2025) Three new species of Peroneutypa (Diatrypaceae, Xylariales) and a first record of Eutypa camelliae in China with updated description. MycoKeys 114: 213-238. https://doi.org/10.3897/mycokeys.114.145312
Open Access

Abstract

Diatrypaceae is a diverse family with a worldwide distribution, occurring on a wide range of hosts in terrestrial and marine environments, some of which are important plant pathogens. During a survey of ascomycete diversity in Guizhou Province, China, three new taxa within Peroneutypa are proposed based on morphological comparisons and phylogenetic analyses of combined ITS and tub2 sequences data. The newly proposed species are Peroneutypa guizhouensis, P. wanfenglinensis and P. zhujiashanesis. In addition, Eutypa camelliae was recorded for the first time from China, with an updated description. Detailed morphological descriptions, illustrations, comparative analyses, and a tabular comparison of the new species with related and similar taxa are provided.

Key words:

3 new species, diatrypaceous fungi, fungal systematics, Karst environment

Introduction

Diatrypaceae is a diverse and ecologically important family of higher ascomycetes within the order Xylariales, inhabiting a wide variety of hosts in both terrestrial and marine environments worldwide (Chlebicki 1986; Glawe and Jacobs 1987; Carmarán and Romero 1992; Carmarán et al. 2006; Trouillas and Gubler 2010; de Almeida et al. 2016; Li et al. 2023). The members of Diatrypaceae are distributed worldwide, can be found on a wide range of plant species, from economically important crops to forest trees with different life modes, functioning as saprobes, pathogens, and endophytes (Vasilyeva and Ma 2014; Dayarathne et al. 2016; Mayorquin et al. 2016; Senwanna et al. 2017; Hyde et al. 2020; Konta et al. 2020). Members of the Diatrypaceae are characterized by black or dark brown, immersed or erumpent, eustromatic or pseudostromatic stromata, 8-spored or polysporous asci with a very long pedicel and J-/J+ apical apparatus, hyaline to light brown allantoid ascospores, and a libertella-like asexual morph (Senanayake et al. 2015; Wijayawardene et al. 2017).

In Wijayawardene et al. (2022) 22 genera within Diatrypaceae have been documented. However, recent taxonomic advances have led to the addition of three more genera, Alloeutypa, Pseudoeutypa, and Stromatolinea, bringing the total to 25 recognized genera (Ma et al. 2023; Habib et al. 2024; Zhang et al. 2024). Collectively, these genera encompass approximately 1000 species. Most of the species belong to Cryptosphaeria, Diatrype, Diatrypella, Eutypa, and Eutypella (Wijayawardene et al. 2022) and are polyphyletic. The polyphyletic nature of these genera arises primarily from the phenotypic plasticity and anatomical similarities observed within Diatrypaceae, where stromatal characteristics are highly variable and often unreliable for clear species delineation (Zhu et al. 2021; Li et al. 2023; Habib et al. 2024). Previous studies have highlighted the challenges in distinguishing members of Diatrypaceae based on morphology alone, with many taxa still lacking molecular data. This absence of molecular data complicates the classification of Diatrypaceae (de Almeida et al. 2016; Shang et al. 2018; Du et al. 2022). Similar difficulties are evident in Peroneutypa. Currently, 38 species of Peroneutypa are listed in Species Fungorum (https://www.speciesfungorum.org), but molecular data are available for only 20 of these species.

Peroneutypa was established by Berlese (1902), but no type species was designated at the time. Later, Rappaz (1987) proposed P. bellula as the type species and considered the genus as a synonym of Eutypella. Carmarán et al. (2006) reinstated Peroneutypa as a distinct genus based on its ascal type and phylogenetic analyses. Peroneutypa is distinguished by its urn-shaped asci, which have a truncate apex and are wider in the middle, where ascospores tend to cluster. In contrast, Eutypella features spindle-shaped asci with ascospores that cluster and swell in the upper portion (Carmarán et al. 2006). Peroneutypa species are characterized by valsoid stroma, ascomata with long prominent necks, sessile to long-stalked asci, with truncate apices and allantoid hyaline or yellowish ascospores (Carmarán et al. 2006; Vasilyeva and Rogers 2010; Shang et al. 2017). Species of the genus are known as saprobes or pathogens and are widely distributed in terrestrial and marine habitats (Shang et al. 2017; Dayarathne et al. 2020; Du et al. 2022)

In a study focused on the diversity of ascomycetes in Guizhou, China, we identified several Diatrypaceae specimens that did not match any known species. To clarify their taxonomic status, we performed phylogenetic analyses using the internal transcribed spacer (ITS) and β-tubulin (tub2) gene regions. These analyses led to the discovery of three new species belonging to Peroneutypa. We present a brief diagnosis, descriptions, images, and phylogenetic placement of these new species.

Materials and methods

Collection and isolation

Ascomycetous fungi associated with decayed branches and twigs of various plants were collected during surveys conducted in Guizhou Province, China. All related habitat information, including details about elevation, climatic conditions, and geographical features, was recorded. The photos of the collected materials were taken using a Canon G15 camera (Canon Corporation, Tokyo, Japan). Materials were placed in paper bags and were taken to the lab for examination. To preserve the freshness of the specimens, they were dried at room temperature. Fungal isolates were obtained through single spore isolation, following the method described by Senanayake et al. (2020). Spores were observed under a Stereo Zoom microscope and transferred to potato dextrose agar (PDA; 39 g/L in distilled water, Difco potato dextrose). Cultures were incubated at 25–30 °C for 1–4 weeks with regular observations. Cultural characteristics, including mycelial color, shape, texture, and growth rate, were documented under normal light conditions.

Herbarium specimens were deposited in the Cryptogams Herbarium of the Kunming Institute of Botany, the Chinese Academy of Sciences (KUN-HKAS), and the Guizhou Provincial Key Laboratory of Agricultural Biotechnology (GZAAS).

Morphological study

Macroscopic characteristics were observed under an Olympus SZ61 stereomicroscope and photographed with a Canon 700D digital camera fitted to a light microscope (Nikon Ni). The morphological characteristics of specimens were examined, and photomicrographs were taken as described in Senanayake et al. (2020). Materials were mounted in water for anatomical examination, and Melzer’s reagent was used where necessary. More than 30 ascospores and 30 asci were measured using the Tarosoft ® image framework (v. 0.9.0.7). Images were arranged using Adobe Photoshop CS6 (Adobe Systems, USA).

DNA extraction, PCR amplification, and sequencing

Genomic DNA was extracted from mycelium sourced from colonies cultured on PDA after 1–2 weeks at 25 °C, using the BIOMIGA Fungal gDNA Isolation Kit (BIOMIGA, Hangzhou City, Zhejiang Province, China). The DNA samples were stored at –20 °C. Internal transcribed spacers (ITS), and β-tubulin (tub2), were amplified by PCR with primers ITS1/ITS4 (White et al. 1990; Gardes and Bruns 1993), and Bt2a / Bt2b (Glass and Donaldson 1995; O’Donnell and Cigelnik 1997), respectively. The components of a 25 μL volume PCR mixture was: 9.5 μL of double distilled water, 12.5 μL of PCR Master Mix, 1 μL of each primer and 1 μL of template DNA. The PCR amplification was conducted as reported by Samarakoon et al. (2022). Qualified PCR products were checked through 1.5% agarose gel electrophoresis stained with GoldenView, and sent to Qingke Biotech Chongqing, China, for sequencing.

Phylogenetic analyses

The newly generated forward and reverse sequences from this study were assembled in the BioEdit v. 7.0.5 (Hall 1999) then were subjected to BLASTn search against the GenBank nucleotide database at the National Center for Biotechnology Information (NCBI) to identify closely related sequences. Sequence data of related taxa were obtained from previous publications (Long et al. 2021; Zhu et al. 2021; Li et al. 2023) and downloaded from the GenBank database (Table 1). The sequences were aligned using MAFFT v.7 online web server (Katoh et al. 2019) under default settings. Alignment was adjusted manually using BioEdit v.7.0.5.3 (Hall 1999) where necessary. The combined sequence data was used to perform maximum likelihood (ML) and Bayesian inference analysis (BI). The ML analysis was implemented in RAxML v.8.2.12 using the GTR substitution model with 1,000 bootstrap replicates (Stamatakis 2014). Bayesian inference analysis was conducted in MrBayes v. 3.2.2 (Ronquist et al. 2012) online, with Markov chain Monte Carlo (MCMC) sampling in MrBayes v.3.2.2 (Ronquist et al. 2012) used to calculate posterior probabilities (PP). Six simultaneous Markov chains were run for 1,000,000 generations, and trees were sampled every 1,000th generation. The convergence of the MCMC procedure was assessed from the effective sample size scores (all > 100) using MrBayes. The first 25% of the trees were discarded as burn-ins. The remainder was used to calculate the posterior probabilities (PPs) for individual branches. The phylogenetic tree was visualized in FIGTREE v.1.4.3 (Rambaut 2012). All analyses were run on the CIPRES Science Gateway v 3.3 web portal (Miller et al. 2010).

Table 1.
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Taxa used in the phylogenetic analyses and their corresponding GenBank accession numbers.

Taxa Strain number GenBank Accession number Reference
ITS β-tubulin
Allocryptovalsa castaneae CFCC52428 MW632945 MW656393 Zhu et al. (2021)
Allocryptovalsa castaneicola CFCC52432 MW632947 MW656395 Zhu et al. (2021)
Allocryptovalsa cryptovalsoidea HVFIG02T HQ692573 NA Trouillas et al. (2011)
Allocryptovalsa elaeidis MFLUCC150707 MN308410 MN340296 Konta et al. (2020)
Allocryptovalsa polyspora MFLU 17-1218 NR153588 MG334556 Senwanna et al. (2017)
Allocryptovalsa rabenhorstii WA08CB HQ692619 HQ692523 Trouillas et al. (2011)
Allocryptovalsa rabenhorstii GMB0416 OP935171 OP938733 Li et al. (2023)
Allocryptovalsa sichuanensis HKAS107017 MW240633 MW775592 Samarakoon et al. (2022)
Allocryptovalsa xishuangbanica KUMCC21-0830 ON041128 ON081498 Maharachchikumbura et al. (2022)
Allodiatrype albelloscutata IFRD9100 OK257020 NA Li et al. (2022)
Allodiatrype arengae MFLUCC 15-0713 MN308411 MN340297 Konta et al. (2020)
Allodiatrype elaeidicola MFLUCC15-0737a MN308415 MN340299 Konta et al. (2020)
Allodiatrype elaeidis MFLUCC150708a MN308412 MN340298 Konta et al. (2020)
Allodiatrype eleiodoxae MFLU23-0357 OR571761 OR591484 Unpublished
Allodiatrype dalbergiae MFLU23-0349 OR571759 OR771026 Unpublished
Allodiatrype dalbergiae MFLU23-0350 OR571760 OR591487 Unpublished
Allodiatrype taiyangheensis IFRDCC2800 OK257021 OK345036 Li et al. (2022)
Allodiatrype thailandica MFLUCC153662 KU315392 NA Li et al. (2016)
Allodiatrype trigemina FCATAS842 MW031919 MW371289 Peng et al. (2021)
Alloeutypa flavovirens E48C AJ302457 DQ006959 Rolshausen et al. (2006)
Alloeutypa milinensis FCATAS4309 OP538689 OP557595 Ma et al. (2023)
Anthostoma decipiens JL567 JN975370 JN975407 Luque et al. (2012)
Cryptosphaeria ligniota CBS273.87 KT425233 KT425168 Acero et al. (2004)
Cryptosphaeria multicontinentalis HBPF8 KT425178 NA Trouillas et al. (2015)
Cryptosphaeria pullmanensis ATCC52655 KT425235 KT425170 Trouillas et al. (2015)
Cryptosphaeria pullmanensis HBPF24 KT425202 KT425137 Trouillas et al. (2015)
Cryptosphaeria subcutanea CBS240.87 KT425232 KT425167 Trouillas et al. (2015)
Cryptovalsa ampelina A001 GQ293901 GQ293972 Trouillas and Gubler (2010)
Cryptovalsa ampelina DRO101 GQ293902 GQ293982 Trouillas and Gubler (2010)
Diatrypasimilis australiensis ATCC MYA-3540 NR111369 NA Schoch et al. (2014)
Diatrype bullata UCDDCh 400 DQ006946 DQ007002 Rolshausen et al. (2006)
Diatrype camelliae-japonicae GMB0427 OP935172 OP938734 Li et al. (2023)
Diatrype disciformis GNA14 KR605644 KY352434 Senanayake et al. (2015)
Diatrype lancangensis GMB0045 MW797113 MW814885 Long et al. (2021)
Diatrype macowaniana Isolate D15C AJ302431 NA Acero et al. (2004)
Diatrype quercicola CFCC-52418 MW632938 MW656386 Zhu et al. (2021)
Diatrype rubi GMB0429 OP935182 OP938740 Li et al. (2023)
Diatrypella atlantica HUEFS 194228 KM396615 KR363998 de Almeida et al. (2016)
Diatrypella banksiae CPC29118 KY173402 NA Crous et al. (2013)
Diatrypella betulicola CFCC52411 MW632935 MW656383 Zhu et al. (2021)
Diatrypella delonicis MFLUCC15-1014 MH812994 MH812994 Hyde et al. (2019)
Diatrypella elaeidis MFLUCC15-0279 MN308417 MN340300 Konta et al. (2020)
Diatrypella fatsiae-japonica GMB0422 OP935184 OP938744 Li et al. (2023)
Diatrypella frostii UFMGCB 1917 HQ377280 NA Vieira et al. (2012)
Diatrypella heveae MFLUCC17-0368 MF959501 MG334557 Senwanna et al. (2017)
Diatrypella hubeiensis CFCC52413 MW632937 NA Zhu et al. (2021)
Diatrypella iranensis T KDQ18 KM245033 KY352429 Mehrabi et al. (2015)
Diatrypella longiasca T KUMCC 20-0021 MW039349 MW239658 Dissanayake et al. (2021)
Diatrypella macrospora T KDQ15 KR605648 KY352430 Mehrabi et al. (2016)
Diatrypella major ANM1947 KU320613 NA de Almeida et al. (2016)
Diatrypella pulvinata H048 FR715523 FR715495 de Almeida et al. (2016)
Diatrypella tectonae MFLUCC120172b KY283085 KY421043 Shang et al. (2017)
Diatrypella vulgaris HVFRA02 HQ692591 HQ692503 Trouillas et al. (2011)
Diatrypella yunnanensis VT01 MN653008 MN887112 Zhu et al. (2021)
Eutypa armeniacae ATCC28120 DQ006948 DQ006975 Rolshausen et al. (2006)
Eutypa astroidea CBS292.87 DQ006966 DQ006966 Rolshausen et al. (2006)
Eutypa camelliae HKAS107022 MW240634 MW775593 Samarakoon et al. (2022)
Eutypa camelliae GZAAS24-0013 PP528182 PQ301430 This study
Eutypa camelliae HKAS-107022 NR175674 MW775593 This study
Eutypa cerasi GMB0048 MW797104 MW814893 Long et al. (2021)
Eutypa consobrina F091 AJ302447 KY111596 Acero et al. (2004)
Eutypa crustata CBS210.87 AJ302448 DQ006968 Rolshausen et al. (2006)
Eutypa laevata CBS291.87 AJ302449 NA Acero et al. (2004)
Eutypa lata EP18 HQ692611 HQ692611 Trouillas et al. (2011)
Eutypa lejoplaca CBS248.87 DQ006922 DQ006974 Rolshausen et al. (2006)
Eutypa maura CBS219.87 DQ006926 DQ006967 Rolshausen et al. (2006)
Eutypa microasca BAFC51550 KF964566 KF964572 Grassi et al. (2014)
Eutypa petrakii var. hedarae BENT014 OP038000 OP079836 Unpublished
Eutypa sparsa 38023b AY684220 AY684201 Trouillas and Gubler (2004)
Eutypa tetragona CBS284.87 DQ006923 DQ006960 Rolshausen et al. (2006)
Eutypella cerviculata EL59C AJ302468 NA Acero et al. (2004)
Eutypella motuoensis FCATAS4082 OP538693 OP557599 Ma et al. (2023)
Eutypella persica IRAN 2540C KX828144 KY352451 Mehrabi et al. (2019)
Eutypella quercina IRAN2543C KX828139 KY352449 Mehrabi et al. (2019)
Eutypella semicircularis MP4669 JQ517314 NA Mehrabi et al. (2016)
Eutypella virescens CBS205.36 MH855778 MH867286 Vu et al. (2019)
Halocryptovalsa salicorniae MFLUCC 15-0185 MH304410 MH370274 Dayarathne et al. (2020)
Halodiatrype avicenniae MFLUCC 150953 KX573916 KX573931 Dayarathne et al. (2016)
Halodiatrype salinicola MFLUCC 15-1277 KX573915 KX573932 Dayarathne et al. (2016)
Kretzschmaria deusta CBS 826.72 KU683767 KU684190 U’Ren et al. (2016)
Monosporascus cannonballus CMM3646 JX971617 NA Unpublished
Monosporascus cannonballus ATCC26931 FJ430598 NA Unpublished
Neoeutypella baoshanensis HMAS255436 MH822887 MH822888 Phookamsak et al. (2019)
Paraeutypella citricolca MFLU23-0352 OR563996 NA Unpublished
Paraeutypella guizhouensis KUMCC 20-0017 MW036142 MW239661 Dissanayake et al. (2021)
Paraeutypella pseudoguizhouensis GMB0420 OP935186 OP938748 Li et al. (2023)
Pedumispora rhizophorae BCC44877 KJ888853 NA Klaysuban et al. (2014)
Peroneutypa aquilariae KUNCC-2210817 NR185767 OP572195 (Du et al. 2022).
Peroneutypa anomianthe KUNCC-2315540 PP584741 PQ046048 Dissanayake et al. (2024)
Peroneutypa anomianthe MFLU-210242 OK393705 NA de Silva et al. (2022)
Peroneutypa alsophila CBS250.87 AJ302467 NA Acero et al. (2004)
Peroneutypa comosa BAFC393 KF964568 NA Grassi et al. (2014)
Peroneutypa curvispora HUEFS136877 KM396641 KM396641 de Almeida et al. (2016)
Peroneutypa diminutiasca MFLUCC17-2144 MG873479 NA Shang et al. (2018)
Peroneutypa diminutispora HUEFS192196 KM396647 NA de Almeida et al. (2016)
Peroneutypa guizhouensis GZAAS24-0087 PQ878089 PQ876910 This study
Peroneutypa guizhouensis GZAAS24-0088 PQ878090 PQ876911 This study
Peroneutypa hainanensis GMB0424 OP935179 OP938746 Li et al. (2023)
Peroneutypa hainanensis GMB0425 OP935180 OP938747 Li et al. (2023)
Peroneutypa hongheensis KUNCC-23-16753 PP584742 PP951427 Dissanayake et al. (2024)
Peroneutypa indica NFCCI 4393 MN061368 MN431498 Dayarathne et al. (2020)
Peroneutypa kochiana F092 AJ302462 NA Carmarán et al. 2006
Peroneutypa kunmingensis HKAS 113189 MZ475070 MZ475070 Phukhamsakda et al. (2022)
Peroneutypa leucaenae T MFLU 18-0816 MW240631 MW775591 Samarakoon et al. (2022)
Peroneutypa longiasca MFLUCC170371 MF959502 MG334558 Senwanna et al. (2017)
Peroneutypa mackenziei MFLUCC16-0072 KY283083 KY706363 Shang et al. (2017)
Peroneutypa mangrovei PUFD526 MG844286 MH094409 Phookamsak et al. (2019)
Peroneutypa nayariophyti MFLU 23-0077 OQ981955 OR019690 Unpublished
Peroneutypa polysporae NFCCI4392 MN061367 MN431497 Dayarathne et al. (2020)
Peroneutypa qianensis GMB0431 OP935177 NA Li et al. (2023)
Peroneutypa qianensis GMB0432 OP935178 NA Li et al. (2023)
Peroneutypa rubiformis MFLU 17-1185 MG873477 MH316763 Shang et al. (2018)
Peroneutypa scoparia MFLUCC 17-2143 MG873477 NA Shang et al. (2018)
Peroneutypa wanfenglinensis GZAAS24-0021 PP852356 PQ301419 This Study
Peroneutypa wanfenglinensis GZAAS24-0022 PP852353 PQ301420 This Study
Peroneutypa zhujiashanesis GZAAS24-0023 PP852363 PQ301421 This Study
Peroneutypa zhujiashanesis GZAAS24-0024 PP852362 PQ301422 This Study
Pseudodiatrype hainanensis GMB0054 MW797111 MW814883 Long et al. (2021)
Quaternaria quaternata CBS278.87 AJ302469 NA Acero et al. (2004)
Quaternaria quaternata GNF13 KR605645 NA Mehrabi et al. (2016)
Stromatolinea grisea GMB4508 PQ113921 PQ115209 Habib et al. (2024)
Stromatolinea guizhouensis GMB4523 PQ113922 PQ115210 Habib et al. (2024)
Stromatolinea guizhouensis GMB4515 PQ113923 PQ115211 Habib et al. (2024)
Stromatolinea hydei GMB4509 PQ113924 PQ115212 Habib et al. (2024)
Stromatolinea linearis MFLUCC 15-0198 KU940149 MW775587 Habib et al. (2024)
Stromatolinea xishuiensis GMB4522 PQ113928 PQ115216 Habib et al. (2024)
Vasilyeva cinnamomic GMB0418 OP935174 OP938737 Li et al. (2023)
Xylaria hypoxylon CBS 122620 AM993141 KX271279 Peršoh et al. (2009)

Results

Phylogenetic analyses

The combined ITS and tub2 dataset consisted of 125 ingroup strains and two outgroups: Kretzschmaria deusta (CBS 826.72) and Xylaria hypoxylon (CBS 122620). After the exclusion of ambiguously aligned regions and long gaps, the final combined data matrix contained 1,350 characters. The final ML optimization likelihood value of the best RAxML tree was −34243.792546. The tree topology derived from Maximum Likelihood (ML) analysis closely resembled that of Bayesian Inference (BI) analysis. The best-scoring RAxML tree is shown in Fig. 1.

Figure 1. 

Phylogram generated from maximum likelihood analysis (RAxML) based on combined ITS and tub2 sequences data. Bootstrap support values for maximum likelihood (ML) greater than 70% and Bayesian posterior probabilities (BPP) greater than 0.90 are displayed at the respective branches (ML/BPP). The newly described species are marked bold in red, and the new record is marked bold in green. Ex-type/type strains are indicated in black bold.

The phylogenetic tree based on BI and ML approaches confirmed the position of our new species nested within the phylogenetic branch of the genus Peroneutypa (Fig. 1). According to the phylogenetic structure of the tree, Peroneutypa formed a large clade. However, the presence of Eutypa microasca (BAFC 51550) in Peroneutypa clade renders its status polyphyletic. This suggests that the taxonomic status of Eutypa microasca should be revisited for clarification. In the phylogram, the Peroneutypa clade is represented with 4 subclades. Subclade 1 includes 12 species, with the new species P. guizhouensis forming a sister relationship with P. hainanensis, supported strongly (ML/BI = 100/1). Subclade 2 contains 10 species, including the new species P. zhujiashanesis, which appears as a sister to P. leucaenae with moderate support (ML/BI = 60/0.99). Subclade 3 consists of two species, P. indica (NFCCI 4393) and Peroneutypa kochiana (F092). Subclade 4 represents solely the new species P. wanfenglinensis, forming a distinct, well-supported clade (ML/BI = 96/0.99) at the basal position of the Peroneutypa clade. The newly generated sequences of Eutypa camelliae clustered with the type strain E. camelliae (HKAS107022). The sister branch to this clade includes E. lata (EP18) and E. americana (ATCC28120).

Taxonomy

Eutypa camelliae Samarakoon, M.C & Hyde, K.D, Fungal Diversity 112 (1), 1–88 (2022)

Index Fungorum: IF558718
Fig. 2

Description.

Saprobic on a dead branch of an unknown tree. Sexual morph: Stromata 3.1–8.3 mm diam., immersed in the bark, carbonaceous, effuse, confluent into irregularly elongated shape with diffuse margins, dark grayish to dull black surface, rarely with white dots on the surface, 20–50 loculate. Ascomata 390–620 µm in height, 190–340 µm in diameter (x̄ = 530 × 260 µm, n = 10), perithecia, coated with a white powdery substance in between, vary from globose to akin to an inverted flask, Ostiole slightly raised, conspicuous, 114–128 μm wide. Peridium 13.5–25 μm thick, two-layered, outer layer dark brown of textural angular cell, inner layer hyaline of elongated cell. Paraphyses septate, 3.5–7.2 μm (x̄ = 5.6 μm, n = 20) wide, constricted at the septa, longer than asci. Asci 50–120 × 4–6.6 μm (x̄ = 77.5 × 5.4 μm, n = 30), 8–spored clavate, with a rounded to truncate apex, J- apical rings. Ascospore 4–5.8 × 1.1–1.8 μm (x̄ = 4.93 × 1.35 μm, n = 30), overlap, allantoid, slightly curved, subhyaline, smooth, aseptate, often with a guttulae at both ends. Asexual morph: undetermined.

Figure 2. 

Eutypa camelliae a, b stromata on dead branch c cross section of a stroma showing perithecia d, e vertical sections of ascomata f ostiole g peridium h paraphyses i–k asci l, m ascospores n germinating ascospore p culture on PDA. Scale bars: 1 mm (a–d); 100 μm (e, f); 10 μm (i–o).

Culture characteristics.

Colonies on PDA reach 60 mm in diameter after seven days at 28 °C. They are cottony, moderately dense, fluffy aerial mycelium, white from above and pale yellowish from below. Mycelium is composed of branched, septate, smooth-walled, hyaline hyphae.

Specimens examined.

China • Guizhou Province, Kaili City, Leigongshan State Reserve (108°11'47"E, 26°22'43"N), altitude 1664 m, on a dead branch of an unknown tree, 23 August 2023, Xin Y Mao & Y.Q. Kang, LGS19 (GZAAS24-0012, KUN-HKAS133146, strain number GZCC 24-0187). GenBank accession numbers (ITS: PP528179; tub2: PQ301429). Guizhou Province, Libo County, MaoLan National Nature Reserve (108°4'9"E, 25°17'8"N), altitude 694 m, on dead branches of an unknown tree, 22 March 2022, Xin Y Mao & Y.Q. Kang, LBML10 (GZAAS24-0013, KUN-HKAS133147; strain number GZCC 24-0188). GenBank accession numbers (ITS: PP528182; tub2: PQ301430).

Notes.

The sequence of our collection GZAAS24-0012 clustered with Eutypa camelliae in the phylogenetic tree, and ITS sequence BLAST searches also confirmed a 100% match with E. camelliae (HKAS 107022). The holotype description of this species was based on immature stromata, no asci or ascospores were observed in the material and the isolates were obtained from internal tissue of the stromata (Samarakoon et al. 2022). Our study represents the first report of Eutypa camelliae from China and provides the complete anatomical details, including mature stromata with asci and ascospores.

Peroneutypa guizhouensis X.Y. Mao, K. Habib & Y.Q. Kang, sp. nov.

Index Fungorum: IF903236
Fig. 3

Etymology.

The epithet refers to the name of the province from where the samples were collected.

Type.

China • Guizhou Province, Guiyang City, Panlongshan Forest Park. (106°49'18"E, 26°44'58"N), altitude 1242.1 m, on branch of an unidentified plant, 8 June 2024. Xin Y Mao & Y.Q. Kang, PLS29 (Holotype GZAAS24-0087; ex-type cultures GZCC 24-0296; Isotype KUN-HKAS 145344). GenBank accession numbers (ITS: PQ878089; tub2: PQ876910).

Description.

Saprobic on dead branches of an unidentified plant. Sexual morph: Stromata 0.5–1.5 mm in diameter, immersed in the host surface, ostiolar canals protruding through the bark, poorly developed, solitary, rarely gregarious, 1–4 locules, usually two, arranged irregularly, dark brown to black, glabrous, circular to irregular in shape, Ascomata (excluding neck) perithecia 400–720 μm high, 400–600 µm diam. (x̄ = 650 × 400 μm, n = 20), immersed in a stroma, black, globose to sub-globose, each has an individual ostiole with a long neck. Ostiolar canals: erumpent, smooth, 300–570 (x̄ = 435 μm) in length, cylindrical, smooth, curved at the apex. Peridium 48–56 μm (x̄ = 52.4 μm) thick, composed of two layers, outer layer dark brown to black, cells thick-walled, texture angularis, inner layers hyaline, cells flattened. Paraphyses 3–5.8 μm (x̄ = 4.9 μm, n = 20) wide, wider at the base, long, septate, smooth-walled. Asci 16–33 × 3.6–6.8 μm (x̄ = 24.1 × 5.0 μm, n = 30), unitunicate, 8-spored, clavate, apically truncates, with a J- apical ring. Ascospore 2.2–4.7 × 1.1–1.8 μm (x̄ = 3.3 × 1.4 μm, n = 30), overlapping, allantoid, subhyaline, smooth, aseptate, strongly curved, with 1–2 small guttules. Asexual morph: undetermined.

Figure 3. 

Peroneutypa guizhouensis (Holotype GZAAS24-0087) a–c stromata surface view d vertical section of ascomata e ostiole f peridium g paraphyses h numerous ascospores i-j asci k, l ascospores m germinating ascospore n culture on PDA. Scale bars: 1 mm (a–d); 100 μm (e–g); 10 μm (h–o).

Culture characteristics.

Colonies growing fast on PDA, reach 55 mm in 1 week at 28 °C, effuse, thin towards the edge, from above at first white, becoming dirty white at the edge after 2 weeks, from below brownish at the center, the rest white.

Additional specimens examined.

China • Guizhou Province, Zunyi county, Dashahe Natural Reserve (107°34'19"E, 29°7'32"N) altitude: 1900 m, on branches of an unidentified plant, 26 April 2024; Xin Y Mao & Y.Q. Kang, XHP01 (Paratype GZAAS24-0088, Isotype KUN-HKAS 145343, ex-paratype cultures GZCC 24-0297).). GenBank accession numbers (ITS: PQ878090; tub2: PQ876911).

Notes.

Peroneutypa guizhouensis is morphologically and phylogenetically like P. hainanensis, mainly due to its strongly curved ascospores. However, P. guizhouensis can be distinguished by its longer ostiolar necks (300–570 μm vs. 105–420 μm), smaller asci (16–33 μm in length, x̄ = 24.1 × 5.0 μm vs. 28.5–40 μm, x̄ = 33.5 × 5.5 μm), and significantly smaller ascospores (2.2–4.7 × 1.1–1.8 μm vs. 5.0–7.3 × 1–2 μm) (Li et al. 2023).

In addition to Peroneutypa hainanensis, P. guizhouensis shares similarities with P. diminutiasca, P. curvispora, and P. qianensis due to its strongly curved ascospores.

Compared to P. diminutiasca, P. guizhouensis has significantly longer ostiolar necks (300–570 μm vs. 105–280 μm), a thicker peridium (48–56 μm vs. 15–32 μm), and smaller ascospores (2.2–4.7 × 1.1–1.8 μm vs. 3.1–5.9 × 1.3–2.2 μm) (Shang et al. 2018). Peroneutypa curvispora differs from P. guizhouensis in having much longer ostiolar necks (400–800 μm vs. 300–570 μm), smaller asci (9–16.5 × 4–6 μm vs. 16–33 μm), and the absence of paraphyses (vs. present) (de Almeida et al. 2016).

Compared to P. qianensis, P. guizhouensis differs in having longer ostiolar necks (300–570 μm vs. 105–420 μm), larger asci (16–33 × 3.6–6.8 vs. 16.5–20.5 × 4–6 μm), and smaller ascospores (2.2–4.7 × 1.1–1.8 μm vs. 4.5–6.3 × 1.5–0.3 μm) and presence of paraphyses (vs. lack) (Li et al. 2023).

These morphological differences (Table 2), combined with phylogenetic evidence, highlight the distinctiveness of P. guizhouensis and confirm its status as a new species.

Table 2.
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Comparison of new taxa with closely related species.

Species Stromata (mm wide) Ascomata (µm) Ostiolar canal (µm) Peridium (µm) Paraphyses (μm) Asci (µm) Ascospores (µm) Country Host References
P. guizhouensis 0.5–1.5 400–720 × 400–600 300–570 long 48–56 3–5.5 16–33 × 3.6–6.8, J– apical ring, short pedicellate 2.2–4.7 × 1.1–1.8, strongly curved China Unknown tree branch This Paper
P. wanfenglinensis 1.9–2.5 330–620 × 280–520 120–140 long 20–45 3.5–6 20–30.5 × 3–5, J– apical ring, long pedicellate 3–4.2 × 1–2, slightly curved China Betula platyphylla This Paper
P. zhujiashanesis 1–2.8 550–910 × 400–570 µm 50–145 long 24–42 3.5–6 23–31 × 3.5–7, J– apical ring, long pedicellate 3.5–5 × 1–1.5, slightly curved China Unknown tree branch This Paper
P. indica N/A 375 × 202 100–350 long, 15–35 1–2 42 × 3.5, short pedicellate, J– apical ring 5.5 × 1.3, slightly curved India Suaeda monoica Dayarathne et al. 2020
P. curvispora 0.6–3 300–700 400–800 long N/A Absent 9–16.5 × 4–6, long pedicellate 3–5 × 1–2, strongly curved Brazil Unidentified plant de Almeida et al. (2016)
P. diminutiasca 1.2–1.4 75–220 × 99–340 193 × 48 15–32 4–7 12–33 × 2.8–5, J− apical ring, long pedicellate, 4.2 × 1.7 µm, slight to moderately curved China, Thailand Unidentified wood Shang et al. 2018; Du et al. 2022
P. hainanensis 0.4–0.7 350–600 × 130–300 105–420 × 80–120 45–65 N/A 28.5–40 × 3.5–6.5, J− apical ring 5.0–7.3 × 1–2, strongly curved China Unidentified plant Li et al. 2023
P. kochiana N/A 150 Neck not prominent N/A N/A 18–28 long, J+ apical ring 4.5–6 × 1.5–2 slightly curved Russia, Spain Atriplex halimus Acero et al. 2004; Carmarán et al. 2006
P. leucaenae N/A 655 × 525 275–350 long 22–43 3.2–7 wide, septate 33 × 4.2, J+ apical ring, long pedicellate 2.9–3.7 × 0.9–1.3 slightly curved Thailand Leucaena leucocephala Samarakoon et al. 2022
P. qianensis 1.5–2 320–540 × 175–290 105–420 × 80–120 45–65 N/A 16.5–20.5 × 4–6, J− apical ring 4.5–6.3 × 1.5–0.3, slightly curved China Unidentified plant Li et al. 2023

Peroneutypa wanfenglinensis X.Y. Mao, K. Habib & Y.Q. Kang, sp. nov.

Index Fungorum: IF902676
Fig. 4

Etymology.

The epithet refers to the name of the location (Wan Feng Lin State Reserve), where the type specimen was collected.

Type.

China • Guizhou Province, Xingyi City, Wan Feng Lin State Reserve (104°55'28"E, 24°59'26"N), altitude 896.8 m, on dead branches of Betula platyphylla, 28 May 2022, Xin Y Mao & Y.Q. Kang, WFL02 (Holotype GZAAS24-0021; Isotype KUN-HKAS133157, ex-type cultures GZCC 24-0196). GenBank accession numbers (ITS: PP852356; tub2: PQ301419).

Description.

Saprobic on decaying branches of Betula platyphylla. Sexual morph: Stromata 1.90–2.5 mm in diameter, interior, solitary to gregarious, with 1–4 perithecia, immersed, erumpent by a long ostiolar canal, dark brown to black, surface glabrous, shape circular to irregular, arranged irregularly. Ascomata (excluding necks) 330–620 μm high, 280–520 µm diam. (x̄ = 540 × 320 μm, n = 10), immersed in a stroma, black, monostichous to distichous, circular to oval, each has an individual ostiole with a short neck. Ostiolar canals erumpent, smooth, 120–140 μm (x̄ = 135 μm, n = 10) long, arch-shaped, sulcate, and curved at the apex. Peridium 20–45 μm (x̄ = 31.85 μm) thick, composed of two layers, outer layer brown to dark, cells thick-walled, texture angularis, inner layers hyaline, cells flattened, texture angularis. Paraphyses septate, slightly swollen at the septa, 3.5–6 μm (x̄ = 5.4 μm, n = 20) wide. Asci 20–30.5 × 3–5 μm (x̄ = 25 × 4 μm, n = 30), unitunicate, 8-spored, clavate, with apically rounded to truncate ends, with a J- apical ring. Ascospore 3–4.2 × 1–2 μm (x̄ = 3.6 × 1.4 μm, n = 30), overlapping, allantoid, subhyaline, smooth, aseptate, with 1–2 oil droplets. Asexual morph: undetermined.

Figure 4. 

Peroneutypa wanfenglinensis (Holotype GZAAS24-0021) a surface view of stromata b cross section of a stroma showing perithecia c, d vertical sections of ascomata e ostioles f peridium g paraphyses h–k asci l, m ascospores n germinating ascospore o culture on PDA. Scale bars: 1 mm (a–c); 10 μm (d, e); 100 μm (f–o).

Culture characteristics.

Colonies growing fast on PDA, reach 60 mm in 1 week at 28 °C, effuse, velvety to hairy, nearly circular, dense towards the edge, fluffy aerial mycelium, appear white from above and pale from below. Mycelium is composed of branched, septate, smooth-walled, hyaline hyphae.

Additional specimens examined.

China • Guizhou Province, Zunyi City, Chishui Zhuhai National Forest Park (105°99'14"E, 28°47'19"N), altitude 838 m, on branches of an unidentified plant, 21 July 2023. Xin Y Mao, CSZH01 (Paratype GZAAS24-0022; KUN-HKAS133156; ex-paratype cultures, GZCC 24-0197). GenBank accession numbers (ITS: PP852353; tub2: PQ301420).

Notes.

BLAST results reveal that Peroneutypa wanfenglinensis is closely related to P. kochiana. However, P. wanfenglinensis differs morphologically from P. kochiana in having smaller ascospores (3–4.2 × 1.0–1.9 μm vs. 4.5–6 × 1.5–2 μm), larger ascomata (330–620 μm high, 280–520 μm diam. vs. 150 μm diam.), and asci with a J- (non-amyloid) apical ring, compared to the J+ (amyloid) apical ring in P. kochiana (Carmarán et al. 2006). Sequence analysis also indicates a notable difference between P. wanfenglinensis and P. kochiana, showing a relatively low ITS similarity of 91%.

In terms of ascomata size and apical ring, Peroneutypa wanfenglinensis is more like P. indica. However, their ascus and ascospore dimensions can differentiate the two species. Peroneutypa indica has longer asci (35–47 μm vs. 20–30.5 μm) and ascospores (4–8 μm vs. 3–4.2 μm) compared to P. wanfenglinensis (Dayarathne et al. 2020).

In terms of ascomata and ascospore dimensions, Peroneutypa wanfenglinensis is comparable to P. leucaenae. However, P. leucaenae can be distinguished by its significantly longer ostiolar neck (275–350 μm vs. 120–140 μm) and larger asci (average 33 × 4.2 μm vs. average 25 × 4 μm). Additionally, P. leucaenae is characterized by a J+ (amyloid) apical ring, contrasting with the J− (non-amyloid) apical ring observed in P. wanfenglinensis (Du et al. 2022).

These distinct morphological features (Table 2), together with their distinct phylogenetic position, support the recognition of P. wanfenglinensis as a new species.

Peroneutypa zhujiashanesis X.Y. Mao & Y.Q. Kang, sp. nov.

Index Fungorum: IF902672
Fig. 5

Etymology.

The epithet refers to the name of the location where the type specimen was collected, Zhujiashan National Forest Park.

Type.

China • Guizhou Province, Douyun City, Weng‘an County, Zhujiashan National Forest Park (107°38'35"E, 26°58'35"N), altitude 848 m, on branches of an unidentified plant, 14 February 2022. Xin Y Mao & Y.Q. Kang, ZJS14 (Holotype GZAAS24-0023; KUN-HKAS133155; ex-type GZCC 24-0198). GenBank accession numbers (ITS: PP852363; tub2: PQ301421).

Description.

Saprobic on decaying branches of an unknown tree. Sexual morph: Stromata 1–2.8 mm in diameter, immersed in the host surface, with necks conspicuously protruding through the bark, erumpent through an ostiolar canal, solitary to gregarious,1–3 locules, mostly solitary, arranged irregularly, dark brown to black, glabrous, circular to irregular in shape, arranged irregularly, delimited by a black zone in host tissues. Ascomata (excluding neck) immersed in a stroma, dark brown to black, perithecia 550–910 μm high, 400–570 µm diam. (x̄ = 790 × 520 μm, n = 10), black, single to aggregated, globose to sub-globose, each has an individual ostiole with a long neck. Ostiolar canals erumpent, smooth, 50–145 μm (x̄ = 134.69 μm) in length, cylindrical, and curved at the apex. Peridium 24–42 μm (x̄ = 32.73 μm) thick, composed of two layers, outer layer dark brown to black, cells thick-walled, texture angularis, inner layers hyaline, cells flattened, texture angularis. Paraphyses 3.5–6 μm (x̄ = 5.4 μm, n = 20) wide, wider at the base, long, septate, smooth-walled, constricted at septa. Asci 23–31 × 3.5–7 μm (x̄ = 27.5 × 5.5 μm, n = 30), unitunicate, 8-spored, clavate, apically truncates, with a J- apical ring. Ascospore 3.5–5 × 1–1.5 μm (x̄ = 4.2 × 1.3 μm, n = 30), overlapping, allantoid, subhyaline, smooth, aseptate, with 1–2 small guttules. Asexual morph: undetermined.

Figure 5. 

Peroneutypa zhujiashanesis (Holotype GZAAS24-0023) a, b stromata on dead branch c transverse section of ascomata d, e vertical section of ascomata f ostioles g peridium h paraphyses i–k asci l, m ascospores n germinating ascospore o culture on PDA. Scale bars: 1 mm (a–d); 100 μm (e–f); 5 μm (i–o).

Culture characteristics.

Colonies grow fast on PDA, reach 60 mm in 1 week at 28 °C, effuse towards the edge, from above at first white, becoming dirty white at the edge after 2 weeks, from below black at center, the rest white.

Additional specimens examined.

China • Guizhou Province, Anlong county, Xianheping National Forest Park (105°36'26"E, 24°58'39"N) altitude: 1298 m, on branches of an unidentified plant, 30 May 2022; Xin Y Mao & Y.Q. Kang, XHP01 (Paratype GZAAS24-0024; KUN-HKAS-133158; ex-paratype GZCC 24-0199). GenBank accession numbers (ITS: PP852362; tub2: PQ301422).

Notes.

Phylogenetically, Peroneutypa zhujiashanesis is closely related to P. leucaenae. Morphologically, it also shares similarities with P. leucaenae in terms of ascomata size and the shape and size of paraphyses. However, P. zhujiashanesis can be distinguished from P. leucaenae by its smaller asci (23–31 × 3.5–7 μm vs. 30–37 × 3.8–4.5 μm) and longer ascospores (3.5–5 μm vs. 2.9–3.7 μm) (Samarakoon et al. 2022). Additionally, P. zhujiashanesis has smaller ostiolar necks (50–145 μm) and a J– apical ring, whereas P. leucaenae has longer ostiolar necks (275–350 μm) and an amyloid apical ring.

Morphologically, Peroneutypa zhujiashanesis is also like P. diminutiasca in ascospore size and presence of a J– subapical ring. However, P. diminutiasca differs by having smaller ascomata (147–218 μm in diameter), with longer ostiolar neck (average 193 μm vs 134.6 μm), and possessing 1–10 locules per ascomata (vs 1–3 loculate, mostly single) (Du et al. 2022).

Based on these morphological differences (Table 2) and phylogenetic evidence, we introduce our collection as a new species, Peroneutypa zhujiashanesis.

Discussion

The taxonomy of the Diatrypaceae has long been challenging, with unstable generic boundaries that lack strong morphological or phylogenetic support. Current classifications often fail to reflect the evolutionary relationships among these fungi accurately. Our phylogenetic analyses, based on ITS and β-tubulin sequences, corroborate previous findings (Shang et al. 2017; Dissanayake et al. 2021; Zhu et al. 2021; Long et al. 2021; Li et al. 2023) and reveal that several genera, such as Cryptosphaeria, Diatrype, Diatrypella, and Eutypa, are not monophyletic and contain multiple problematic clades.

The phylogeny of the family reveals many clades that may represent distinct genera, suggesting that the current classification is too simplistic. A comprehensive revision involving extensive sampling and a combination of taxonomic methods (integrating morphology, molecular data, chemical profiles and genomic information) is essential to resolve these complex relationships and achieve a more natural classification for the Diatrypaceae. Peroneutypa exemplifies the unresolved taxonomic and phylogenetic issues that are common within Diatrypaceae.

Phylogenetically, Peroneutypa species form a well-supported clade (Fig. 1). However, the inclusion of Eutypella microasca within this clade suggests its polyphyletic nature. In our investigation, we found that molecular data did not correlate well with morphological characteristics. Species that are phylogenetically close differed significantly in morphology. For instance, P. polysporae, characterized by multispored asci, clustered with P. mangrovei, which possesses eight-spored asci. Similarly, the newly described species Peroneutypa guizhouensis clustered with P. hainanensis, both of which possess strongly curved ascospores. However, other strongly curved ascospore-bearing species, such as P. curvispora, P. diminutiasca, P. obesa, and P. qianensis, are phylogenetically distant. Another example is Peroneutypa zhujiashanesis, which has a J- apical ring, clustering with P. leucaenae, a species that possesses a J+ apical ring. Only six species in the genus are known to have a J+ apical ring (P. alsophila, P. comosa, P. exigua, P. iranica, P. kochiana, and P. leucaenae), and they are not closely clustered phylogenetically.

These analyses suggest that the genus Peroneutypa exhibits a complex evolutionary history, where phylogenetic relationships are not always reflected in morphological traits. The discordance between molecular and morphological data underscores the need for a comprehensive integrative approach. The use of additional genetic regions, such as LSU, SSU, rpb2, and tef-1α, should be explored to achieve more accurate phylogenetic analyses. Coupled with detailed morphological studies, these efforts will enable the precise delineation of species boundaries and provide deeper insights into the evolutionary relationships within this genus. The inclusion of newly described species in future studies, such as those introduced in this research, will continue to refine the phylogeny and enhance understanding of the genus’s diversity and evolutionary patterns, and contribute to a more robust and accurate classification system.

Peroneutypa species are mostly reported from plant species belonging to the families Thymelaeaceae, Rubiaceae, Moraceae, Fabaceae, and Euphorbiaceae, among others (Du et al. 2022) and primarily associated with woody angiosperms, particularly trees and shrubs, and are rarely found on herbaceous plants. To date, there are no reports of Peroneutypa species occurring on gymnosperms. Among angiosperms, Peroneutypa species are mainly recorded on dicots, with the exception of P. scoparia and P. bellula, which have been documented on monocots.

Within this genus, species can be distinguished based on several morphological traits, including the size of the ascomata, ostiolar canal length, asci, and ascospores. Additional characteristics, such as the shape of the asci and ascospores, the reaction of the ascus apex in Melzer’s reagent, and the presence or absence of paraphyses, are also used in differentiating species (Shang et al. 2017; Du et al. 2022). Peroneutypa polymorpha and P. rubiformis are unique within the genus for having larger asci, measuring more than 40 µm in length. Most other species in the genus have smaller asci. Similarly, ascospore size is generally consistent across the genus, ranging from 3–6 × 1–2 µm, except P. polysporae, which has notably larger ascospores measuring up to 9 × 1.8 µm.

Ostiolar canal length is another key characteristic for species delimitation. Species such as P. coffea, P. comosa, P. curvispora, P. cylindrica, P. cyphelioides, P. exigua, P. komonoensis, P. macroceras, P. philippinarum, and P. variabilis all have longer ostiolar canals (> 500 µm), distinguishing them from other species in the genus that have shorter canals (< 400 µm). Curved ascospores have been observed in P. curvispora, P. diminutiasca, P. hainanensis, P. obesa and P. qianensis, another prominent differentiating feature within the species.

Overall, the morphological variability in Peroneutypa requires using a combination of these traits for accurate species identification.

Acknowledgements

The authors would like to thank the Key Laboratory of Microbiology and Parasitology of Education Department, Guizhou, Medical University. Engineering - Research. Center of the Utilization, for Characteristic-pharmaceutical Resources in Southwestern,” Ministry of Education Guizhou University, School of Food and Drug Manufacturing Engineering, Guizhou Institute of Technology, Guizhou Academy of Agricultural Sciences.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This research was supported by Support Fund: High-level Innovation Talent Project of Guizhou Province (GCC[2022]036-1); The Science and Technology Department of Guizhou Province innovation and project grant (QKHJC-ZK[2022]YB297),111 Project (D20009);China-Ukraine Intergovernmental Exchange Project (8); National Natural Science Foundation of China (NSFC;no.32060034/no.32460051); International Science and Technology Cooperation Base of Guizhou Province ([2020]4101); Scientists Workstation Guizhou Province KXJZ[2024]009; Guizhou Key Laboratory (ZDSYS[2023]004); Talent Base Project of Guizhou Province, China [RCJD2018-22]; Major Science and Technology Projects of China Tobacco [No.110202101048(LS-08)]; Foundation of Key Laboratory of Microbiology and Parasitology of Education Department, Guizhou (QJJ [2022] 019); Ministry of Education Project(07150120711).

Author contributions

Yingqian Kang conceived and designed the experiments and performed the experiment. Xinying Mao analyzed the data and wrote the manuscript. Hongde Yang dealt with some of the sequences. Kamran Habib and Rizwana Zulfiqar reviewed and polished the language and approved the final version of the manuscript. All authors contributed extensively to the study presented in the manuscript.

Author ORCIDs

Xinying Mao https://orcid.org/0000-0003-0684-3124

Kamran Habib https://orcid.org/0000-0003-2572-0306

Data availability

The datasets generated during and/or analyzed during the current study are available in the MycoBank repository (included in the manuscript) and GenBank (included in Table 1). Also, the datasets generated during and/or analyzed during the current study are available from the corresponding author upon reasonable request.

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