Taxonomic contributions to Pleosporales and Kirschsteiniotheliales from the Xizang Autonomous Region, China

Shu-Cheng He; Kevin D Hyde; Ruvishika S Jayawardena; Vinodhini Thiyagaraja; Dhanushka N Wanasinghe; Yun-Wei Zhao; Zhi-Yang Wang; Tong Cai; Yan-Yan Yang; F Al-Otibi; Zhuliang Yang; Qi Zhao

doi:10.1080/21501203.2025.2493072

. 2025 May 27;17(1):2493072. doi: 10.1080/21501203.2025.2493072

Taxonomic contributions to Pleosporales and Kirschsteiniotheliales from the Xizang Autonomous Region, China

Shu-Cheng He ^a,^b, Kevin D Hyde ^a,^b,^c,^✉, Ruvishika S Jayawardena ^b, Vinodhini Thiyagaraja ^a, Dhanushka N Wanasinghe ^d,^e, Yun-Wei Zhao ^a,^f, Zhi-Yang Wang ^a,^g, Tong Cai ^h, Yan-Yan Yang ^a,^b, F Al-Otibi ^c, Zhuliang Yang ^a,^✉, Qi Zhao ^a,^✉

PMCID: PMC13007439 PMID: 41878510

ABSTRACT

The Xizang Autonomous Region, China, is a key ecological security barrier, renowned for its rich biodiversity and unique resources. It supports diverse life forms, including fungi, which are vital for ecological balance and the restoration of the region’s ecosystems. This study presents the results of a microfungal survey conducted on the Plateau, focusing on the orders Pleosporales and Kirschsteiniotheliales within the class Dothideomycetes. Based on morphological characteristics and multi-gene (ITS, LSU, SSU, tef1, tub2, and rpb2) phylogeny analyses, we report the discovery of 10 new taxa, including one new genus Neotriplosphaeria and nine new species, Kirschsteiniothelia linzhiensis, K. yadongensis, Neotriplosphaeria yadongensis, Periconia linzhiensis, Tetraploa linzhiensis, Torula dingjieensis, T. yadongensis, Trichobotrys motuoensis, Triseptatospora yadongensis, and one new host record Periconia spodiopogonis. The findings provide valuable insights into the morphological and phylogenetic relationships of these fungi, highlighting the Plateau’s significant role in fungal diversity. This study enriches our understanding of fungal biodiversity in extreme environments and underscores the importance of continued exploration in this region.

KEYWORDS: 10 new taxa, Dothideomycetes, phylogeny, taxonomy, saprobic

1. Introduction

Xizang Autonomous Region, China, as an important national ecological security barrier, is one of the richest regions in the world in terms of biodiversity and unique biological resources. It is called a rare “green gene bank” and “natural science museum” in the world (Wang et al. 2023b). It provides a good living and reproduction space for all kinds of animals, plants, and fungi (Yang et al. 2017; Zhan et al. 2022). As a large group of eukaryotic organisms, fungi are a crucial part of the biodiversity of the plateau and play an important role in the ecological balance and restoration of the region (Yang et al. 2010). In the past years, especially from 2010 to 2024, researchers have carried out large-scale investigations and research on fungi in the Qinghai-Xizang Plateau, and have achieved gratifying and important results (Huang et al. 2015; Su et al. 2015; Ren et al. 2016; Timdal et al. 2016; Zhao et al. 2016; Chen et al. 2017, 2020; Shi et al. 2018; Pem et al. 2019, 2024; Phookamsak et al. 2019; Hyde et al. 2020, 2021; Boonmee et al. 2021; Chethana et al. 2021; Liu et al. 2022; Ma et al. 2023, 2025; Xu et al. 2023a, 2023b, 2024a, 2024b; He et al. 2024a; Wang et al. 2024a, 2024b; Phurbu et al. 2024). However, the remarkable species profusion within the region implies that a considerable number of fungi have yet to be identified.

Pleosporales, one of the largest orders within the class Dothideomycetes, was formally introduced by Barr (1987). It is a highly diverse order, currently encompassing 91 families and 653 genera (Hyde et al. 2024b). Members of Pleosporales are primarily characterised by their production of perithecial ascomata, which are flask-shaped fruiting bodies that typically possess a papillate, ostioles, the asexual morph being mainly coelomycetous and hyphomycetous (Hyde et al. 2013; Hongsanan et al. 2020; Pem et al. 2024). Pleosporales fungi have a global distribution in various ecosystems, ranging from terrestrial to aquatic environments (Hongsanan et al. 2020). They exhibit a high abundance in temperate regions and have been identified in soils, decaying wood, and plant debris (Hongsanan et al. 2020). In addition, some species of Pleosporales were reported as plant pathogens that can cause diseases in economically important crops, such as Alternaria, Bipolaris, Leptosphaeria, Phaeosphaeria, and Pyrenophora (Tanaka et al. 2015; El-Demerdash 2018; Jayawardena et al. 2019a, 2019b, 2022, 2025; Bhunjun et al. 2020). Kirschsteiniotheliales was introduced by Hernández-Restrepo et al. (2017) based on phylogenetic analysis to accommodate Kirschsteiniotheliaceae, with Kirschsteiniothelia as the type genus. Kirschsteiniotheliales currently encompasses one family and two incertae sedis genera (Hyde et al. 2024b).

Supported by the Second Tibetan Plateau Scientific Expedition and Research Program, seven field trips were carried out in the southeast of Xizang Autonomous Region, China (He et al. 2023, 2024b; Thiyagaraja et al. 2024) and more than 26 specimens of Pleosporales and Kirschsteiniotheliales were collected. In this paper, we used morphological data together with phylogenetic analysis of the internal transcribed spacer region (ITS), the large subunit rDNA (LSU), the beta-tubulin (tub2), the translation elongation factor 1-α (tef1) gene, and RNA polymerase II second-largest subunit (rpb2) to assess the systematic position of these specimens.

2. Materials and methods

2.1. Collection, fungal isolation, and morphology

Specimens were collected from Xizang Autonomous Region, China, following the methods of He et al. (2024b). The specimens were observed under a NIKON MODEL C-PSN Series dissecting stereo microscope (Nikon, Japan). Using a surgical needle transferred into a clean slide and photographed with the NIKON ECLIPSE Ni-U microscope imaging system (Nikon, Japan), observations of conidiophores, conidiogenous cells, and conidia were carried out. A photoplate was made using Adobe Photoshop CC 2019. The fungal structures were measured using Image Frame Work software. Pure cultures were obtained via single spore isolation using potato dextrose agar (PDA) as described in Senanayake et al. (2020) and incubated at 25 °C for one week. The pure cultures were deposited at the Kunming Institute of Botany Culture Collection (KUNCC). Specimens were deposited at the Herbarium of Cryptogams Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, China (KUN-HKAS). Facesoffungi and Index Fungorum numbers were registered following the protocol described in Jayasiri et al. (2015) and Index Fungorum in 2025, respectively.

2.2. DNA extraction, PCR amplification, and sequencing

The mycelia growing on a Potato Dextrose Agar (PDA) plate were used to extract DNA using the Trilief™ Plant Genomic DNA Kit (Tsingke Biological Technology Co., Ltd. in Beijing, China). When the fungi did not germinate or grow in culture, DNA was directly extracted using a DNA extraction kit (E.Z.N.A.® Forensic DNA kit, D3591-01, Omega Bio-Tek), following the manufacturer instructions (Telle and Thines 2008). The primer pairs ITS5/ITS4 (White et al. 1990), NS1/NS4 (White et al. 1990), LR0R/LR5 (Vilgalys and Hester 1990), T1/T22 (O’Donnell and Cigelnik 1997), EF1-983F/EF1-2218 R (Carbone and Kohn 1999), and fRPB2-5F/fRPB2-7cR (Liu et al. 1999) were used for amplification of the internal transcribed spacer region ITS1-5.8S-ITS2 (ITS), the large subunit rDNA (LSU), the beta-tubulin (tub2), the translation elongation factor 1-α (tef1) gene, and RNA polymerase II second-largest subunit (rpb2), respectively. The polymerase chain reaction (PCR) reaction was performed in a 25 μL reaction volume, comprising 21 μL PCR Mix (2× Rapid Taq Master Mix, Vazyme Biotech Co., Ltd., Nanjing, China), 1 μL of each primer, 2 μL DNA template (Table 1). The PCR products were visualised using agarose gel electrophoresis. The materials together with the targeted bands were sent to Sangon Biotech Co. Ltd., Kunming, China, for sequencing. The newly generated sequences were submitted to the GenBank (Tables 2–7).

Table 1.

Locus, primers, and PCR amplification conditions used in this study.

Locus	Primers	PCR amplification conditions
SSU	NS1/NS4	95 °C: 5 min, (95 °C: 15 s, 55 °C: 15 s, 72 °C: 15 s) × 40 cycles
ITS	ITS5/ITS4
LSU	LR0R/LR5
tef1	EF1-983F/EF1-2218R	95 °C: 5 min, (95 °C: 45 s, 52 °C: 45 s, 72 °C: 70 s) × 35 cycles
rpb2	fRPB2-5F/fRPB2-7cR	95 °C: 5 min, (95 °C: 45 s, 55 °C: 120 s, 72 °C: 50 s) × 35 cycles

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Table 2.

GenBank accession numbers of the taxa in this study of Kirschsteiniothelia.

Taxon	Voucher	GenBank accession numbers
Taxon	Voucher	ITS	LSU	SSU
*Kirschsteiniothelia acutispora*	MFLU21-0127	OP120780	ON980758	ON980754
Kirschsteiniothelia aethiops	MFLUCC 16-1104	MH182583	MH182589	MH182615
Kirschsteiniothelia aethiops	MFLUCC 15-0424	KU500571	KU500578	KU500585
*Kirschsteiniothelia aquatica*	MFLUCC 16-1685	MH182587	MH182594	MH182618
*Kirschsteiniothelia arasbaranica*	IRAN 2508C	KX621983	KX621984	KX621985
*Kirschsteiniothelia atra*	DEN	MG602687	N/A	N/A
*Kirschsteiniothelia bulbosapicalis*	GZCC 23-0732	PQ248937	PQ248933	PQ248929
*Kirschsteiniothelia cangshanensis*	MFLUCC 16-1350	MH182584	MH182592	N/A
*Kirschsteiniothelia chiangmaiensis*	MFLUCC 23-0209	OR575473	OR575474	OR575475
*Kirschsteiniothelia crustacea*	MFLU21-0129	MW851849	MW851854	N/A
*Kirschsteiniothelia dendryphioides*	KUNCC 10431	OP626354	PQ248935	PQ248931
*Kirschsteiniothelia distoseptata*	HKAS 131382	OR714747	N/A	OR743200
*Kirschsteiniothelia dujuanhuensis*	HKAS 129177	OQ874971	OQ732682	OQ875039
Kirschsteiniothelia dushanensis	GZCC19-0415	OP377845	MW133830	MW134610
Kirschsteiniothelia ebriosa	CBSH-23379	N/A	LT985885	N/A
*Kirschsteiniothelia emarceis*	FLU10-0037	NR_138375	NG059454	N/A
*Kirschsteiniothelia extensum*	MFLU21-0130	MW851850	MW851855	N/A
*Kirschsteiniothelia fluminicola*	MFLUCC 16-1263	MH182582	MH182588	N/A
*Kirschsteiniothelia guangdongensis*	ZHKUCC 22-0233	OR164946	OR164974	N/A
*Kirschsteiniothelia guizhouensis*	GZCC 24-0034	PQ404852	PQ404856	PQ404859
*Kirschsteiniothelia inthanonensis*	MFLUCC 23-0277	OR762773	OR762781	OR764784
*Kirschsteiniothelia laojunensis*	KUN.L 88727	PP081651	PP081658	N/A
*Kirschsteiniothelia lignicola*	MFLUCC 10-0036	HQ441567	HQ441568	HQ441569
*Kirschsteiniothelia linzhiensis*	HKAS 144539*	PV484695	PQ675402	PQ675362
Kirschsteiniothelia linzhiensis	HKAS 144540*	PV484696	PQ675403	PQ675363
*Kirschsteiniothelia longiconidiophora*	KUNCC:23-13756	OR589303	OR600952	OR743201
*Kirschsteiniothelia longirostrata*	GZCC 23-0733	PQ248939	PQ248934	PQ248930
*Kirschsteiniothelia longispora*	HKAS 136268	PQ038266	PQ038273	PQ046108
*Kirschsteiniothelia nabanheensis*	HJAUP C2004	OQ023197	OQ023273	OQ023038
*Kirschsteiniothelia phoenicis*	MFLUCC 18-0216	MG859978	MG860484	MG859979
*Kirschsteiniothelia pini*	UESTCC24.0131	PP835321	PP835315	PP835318
*Kirschsteiniothelia puerensis*	ZHKUCC:22-0271	N/A	NG_242017	OP451020
*Kirschsteiniothelia ramus*	GZCC:23-0596	OR098711	OR091333	N/A
*Kirschsteiniothelia rostrata*	MFLUCC 15-0619	KY697280	KY697276	KY697278
*Kirschsteiniothelia saprophytica*	MFLUCC_23-0275	OR762774	OR762783	N/A
*Kirschsteiniothelia septemseptata*	FLU21-0126	OP120779	ON980757	ON980752
*Kirschsteiniothelia sichuanensis*	HKAS 134908	PP785368	PP784322	N/A
Kirschsteiniothelia sp.	CX79B2	OQ645272	OQ645286	OQ645279
Kirschsteiniothelia sp.	SICAUCC 23-0043	PP060659	PP057952	PP003820
Kirschsteiniothelia sp.	SICAUCC 23-0044	PP060660	PP057953	PP003821
Kirschsteiniothelia sp.	HJAUP C1209	PP505546	PP506568	PP527763
Kirschsteiniothelia sp.	HJAUP C1501	PP505547	PP506569	N/A
Kirschsteiniothelia sp.	HJAUP C1273	PP505548	PP506566	PP506565
Kirschsteiniothelia sp.	HJAUP C1313	PP505549	PP506562	PP506563
Kirschsteiniothelia sp.	KUNCC:23-13755	N/A	N/A	OR743199
*Kirschsteiniothelia spatiosum*	MFLU21-0128	OP077294	N/A	ON980753
*Kirschsteiniothelia submersa*	MFLUCC 15-0427	KU500570	KU500577	KU500584
Kirschsteiniothelia tectonae	MFLUCC 12-0050	KU144916	KU764707	N/A
*Kirschsteiniothelia thailandica*	MFLUCC 20-0116	MT985633	MT984443	MT984280
Kirschsteiniothelia thujina	JF13210	KM982716	KM982718	KM982717
*Kirschsteiniothelia vinigena*	CBSH-23378	N/A	NG075229	N/A
*Kirschsteiniothelia weiningensis*	GZCC 24-0072	PQ404851	PQ404855	PQ404858
*Kirschsteiniothelia xishuangbannaensis*	ZHKUCC 22-0220	OP289566	OP303181	OP289564
*Kirschsteiniothelia xishuiensis*	GZCC 24-0052	PQ404850	PQ404854	PQ404857
*Kirschsteiniothelia yadongensis*	HKAS 144543*	PQ684985	PV483694	PV483715
Kirschsteiniothelia yadongensis	HKAS 144544*	PQ684986	PV483695	PV483716
*Kirschsteiniothelia zizyphifolii*	MFLUCC 23-0270	OR762768	OR762776	OR764779
Homortomyces combreti	CPC 19808	JX517281	JX517291	N/A
*Homortomyces tamaricis*	MFLUCC 13–0441	NR_155161	NG_059495	N/A

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The newly generated sequences are shown in gray shade, and the type strains are marked with an asterisk (*). Other type strains are in bold. ‘N/A’ indicates that the data are not available in GenBank.

Table 3.

GenBank accession numbers of the taxa in this study of Astrosphaeriellaceae.

Taxon	Voucher	GenBank accession numbers
Taxon	Voucher	LSU	SSU	tef1
Acrocordiopsis patilii	MFLU 18-0533	MN017850	MN017916	N/A
Acrocordiopsis patilii	BCC 28166	GU479772	GU479736	N/A
Acuminatispora palmarum	MFLU:18-1068	MH390437	MH390401	MH399248
Acuminatispora palmarum	MFLU:18-1069	MH390438	MH390402	MH399249
Aquatospora cylindrica	MFLUCC 18-1287	MN913715	MT864327	MT954375
Astrosphaeriella bambusae	MFLU:12-2471	KT955461	N/A	KT955424
Astrosphaeriella fusispora	MFLUCC 10-0555	KT955462	KT955443	KT955425
Astrosphaeriella neofusispora	MFLU:11-0197	KT955463	KT955444	KT955426
Astrosphaeriella stellata	MFLU 18-2030	JN846723	JN846733	N/A
Astrosphaeriella thailandica	MFLU:11-0227	KT955465	KT955445	KT955427
Astrosphaeriella thysanolaenae	MFLU:11-0222	KT955466	KT955446	KT955428
Astrosphaeriellopsis bakeriana	CBS 115556	GU301801	N/A	GU349015
Astrosphaeriellopsis caryotae	MFLU:17-1255	NG_058605	NG_063660	MF588974
Caryospora aquatica	MFLU:11-1083	NG_059058	MH057850	N/A
Caryospora quercus	MFLU:18-2151	NG_066440	MK347869	N/A
Caryospora submersa	MFLU:18-1539	NG_073802	N/A	N/A
Mycopepon smithii	GMB0202	MH049445	MH049453	N/A
Mycopepon smithii	LGS20	MH049442	MH049450	N/A
Mycopepon smithii	SMH 1609	AF279400	AF279399	N/A
Pithomyces caryotae	MFLUCC 13-0828	MF588999	NG_063662	MF588978
Pithomyces flavus	MTCC 12224	KP814133	KP814134	N/A
Pithomyces licualae	MFLUCC 17-2031	NG_059846	NG_063661	N/A
Pithomyces esuvius	MTCC 12224	KP814136	KP814137	N/A
Pteridiospora arengae	MFLUCC 15-0289	NG_228976	ON650678	ON734017
Pteridiospora bambusae	MFLU:18-0071	MG831565	MG831566	MG833012
Pteridiospora chiangraiensis	MFLUCC 11-0162	KT955480	KT955459	KT955442
Pteridiospora javanica	MFLUCC 11-0159	KJ742940	KJ739607	KJ739605
Quercicola fusiformis	MFUCC 18-0479	MK348009	MK347898	MK360085
Quercicola guttulospora	MFLU:18-2192	MK348010	MK347899	MK360086
Triseptatospora calami	MFLUCC 15-0305	N/A	ON650705	ON734018
Triseptatospora yadongensis	HKAS 134939*	PQ675404	PQ675364	PQ671461
Triseptatospora yadongensis	HKAS 134940*	PQ675405	PQ675365	PQ671462
Xenoastrosphaeriella aquatica	CGMCC 3.20683	MZ420753	MZ420754	MZ442701
Xenoastrosphaeriella aquatica	DLUCC 0869	MZ420752	MZ420755	MZ442700
Xenoastrosphaeriella tornata	MFLUCC 11-0196	KT955467	KT955447	KT955429
Xenoastrosphaeriella tornata	KUMCC 18-0194	MT659668	MT659669	MT653597
Murispora bromicola	MFLUCC 15-0031	NG_059595	KT305996	KT305999
Murispora kazachstanica	CBS 148424	NG_088293	NG_081421	OK019410

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Table 4.

GenBank accession numbers of the taxa in this study of Dictyosporiaceae.

Taxon	Voucher	GenBank accession numbers
Taxon	Voucher	ITS	LSU	SSU	tef1
Aquadictyospora clematidis	MFLU 172080	MT310592	MT214545	MT226664	MT394727
Aquadictyospora lignicola	MFLUCC 17-1318	MF948621	MF948629	N/A	N/A
Dendryphiella paravinosa	CPC26176	KX228257	KX228309	N/A	N/A
Dendryphiella vinosa	MFLU200444	MT907477	MT907480	N/A	N/A
Dictyocheirospora aquatica	KUMCC 15-0305	KY320508	KY320513	AB787223	AB808489
Dictyocheirospora bannica	KH332	LC014543	AB807513	MH381759	N/A
Dictyocheirospora pseudomusae	yone 234	LC014550	AB807520	AB797230	AB808496
Dictyosporium alatum	ATCC 34953	NR_077171	DQ018101	DQ018080	N/A
Dictyosporium elegans	NBRC 32502	DQ018087	DQ018100	DQ018079	N/A
Dictyosporium meiosporum	MFLUCC 10-0131	KP710944	KP710945	KP710946	N/A
Dictyosporium nigroapice	MFLUCC 17-2053	MH381768	MH381777	MH381762	MH388821
Dictyosporium olivaceosporum	KH375	LC014542	AB807514	AB797224	AB808490
Dictyosporium pandanicola	MFLUCC 18-0331	MZ490792	MZ490776	N/A	MZ501208
Dictyosporium tetrasporum	KT2865	LC014551	AB807519	AB797229	AB808495
Dictyosporium tratense	MFLUCC 17-2052	MH381767	MH381776	MH381761	MF388820
Gregarithecium curvisporum	KT922	AB809644	AB807547	AB797257	AB808523
Jalapriya pulchra	MFLU17-1683	MF948628	MF948636	N/A	MF953171
Jalapriya toruloides	CBS209.65	DQ018093	DQ018104	DQ018081	N/A
Neodendryphiella brassaiopsidis	MHZU:23-0113	OR365455	OR365485	OR365491	N/A
Neodendryphiella michoacanensis	CBS:144323	NR_160583	NG_066395	N/A	N/A
Neodendryphiella tarraconensis	CBS:144324	NR_160582	NG_066394	N/A	N/A
Paradictyocheirospora tectonae	NFCCI:4878	NR_189839	NG_241935	N/A	MW854832
Pseudocoleophoma calamagrostidis	KT3284	LC014592	LC014609	LC014604	LC014614
Pseudocoleophoma flavescens	CBS178.93	N/A	GU238075	GU238216	N/A
Pseudocoleophoma polygonicola	KT731	AB809634	AB807546	AB797256	AB808522
Pseudodictyosporium elegans	CBS 688.93	MH862454	MH874101	DQ018084	N/A
Pseudodictyosporium thailandica	MFLUCC 16-0029	KX259520	KX259522	KX259524	KX259526
Trichobotrys effusa	JAUCC 6359	PP406377	PP407503	PP407508	PP405621
Trichobotrys effusus	SNC73	PP592489	PP621120	PP639248	PP761036
Trichobotrys meilingensis	JAUCC 4985	PP406380	PP407504	PP407509	PP405623
Trichobotrys motuoensis	HKAS 144531*	PQ684987	PQ675406	PQ675366	PQ671463
Trichobotrys motuoensis	HKAS 144532*	PQ684988	PQ675407	PQ675367	PQ671464
Trichobotrys sinensis	KUNCC 23-14554	OR233595	OR335805	OR501421	OR547995
Trichobotrys yunjushanensis	JAUCC 4987	PP406378	PP407506	PP407511	PP405622
Verrucoccum coppinsii	E. 00814291	NR_177147	NG_153929	NG_081399	N/A
Verrucoccum spribillei	MSC 31786	NR_177539	NG_088165	NG_087876	N/A
Vikalpa grandispora	KUNCC 22-12425	OP526638	OP526648	OP526628	OP542240
Vikalpa sphaerica	CGMCC 3.20682	OP526639	OP526649	OP526629	OP542241
Periconia igniaria	CBS 379.86	LC014585	AB807566	AB797276	AB808542
Periconia igniaria	CBS 845.96	LC014586	AB807567	AB797277	AB808543

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Table 5.

GenBank accession numbers of the taxa in this study of Periconia.

Taxon	Voucher	GenBank accession numbers
Taxon	Voucher	SSU	ITS	LSU	tef1
Periconia algeriana	CBS 321.79	N/A	MH861212	MH872979	N/A
Periconia alishanica	KUMCC 19-0174	N/A	MW063167	MW063231	MW183792
Periconia alishanica	NCYUCC 19-0186	N/A	MW063166	MW063230	MW183791
Periconia ananasi	MFLUCC:21-0155	OL606142	OL753685	OL606153	OL912946
Periconia aquatica	MFLUCC16-0912	N/A	KY794701	KY794705	KY814760
Periconia arecacearum	MFLU 19-0803	PP639222	PP592462	PP621090	PP828795
Periconia artemisiae	KUMCC 20-0265	MW448658	MW448657	MW448571	MW460898
Periconia atropurpurea	CBS 381.55	N/A	MH857524	MH869061	N/A
Periconia banksiae	CBS 129526	N/A	JF951147	NG_064279	N/A
Periconia byssoides	MFLUCC17-2292	MK347858	MK347751	MK347968	MK360069
Periconia byssoides	MFLUCC18-1553	MK347914	MK347806	MK348025	MK360068
Periconia caespitosa	LAMIC 110/16	N/A	MH051906	MH051907	N/A
Periconia calamagrostidicola	CBS H-25342	N/A	NR_197928	PP791460	PP780620
Periconia celtidis	MFLU 19-2784	N/A	NR_174830	NG_079543	N/A
Periconia chengduensis	UESTCC 22.0140	OP956046	OP955977	OP956002	OP961443
Periconia chiangraiensis	MFLU 21-0280	OL606143	OL753686	OL606154	OL912947
Periconia chimonanthi	KUMCC 20-0266	MW448656	NR_176752	MW448572	MW460897
Periconia circinata	CBS 263.37	N/A	MW810265	MH867413	MW735660
Periconia citlaltepetlensis	ENCB 140251	N/A	MH890645	MT625978	N/A
Periconia citlaltepetlensis	IOM 325319.2	N/A	MT649221	MT649216	N/A
Periconia cookei	UESTCC 22.0134	OP956037	OP955968	OP955993	N/A
Periconia cortaderiae	MFLUCC 15-0457	KX986345	KX965732	KX954401	KY310703
Periconia cynodontis	CGMCC 3.23927	OP909920	OP909925	OP909921	OP961434
Periconia cyperacearum	CPC 32138	N/A	MH327815	MH327851	N/A
Periconia delonicis	MFLUCC 17-2584	MK347832	N/A	MK347941	MK360071
Periconia didymosporum	MFLU 15-0058	KP761738	KP761734	KP761731	KP761728
Periconia digitata	CBS 510.77	AB797271	LC014584	AB807561	AB808537
Periconia elaeidis	MFLUCC 17-0087	MH108551	MG742713	MH108552	N/A
Periconia endophytica	ZHKUCC 23-0995	PP277722	OR995582	OR995588	PP025968
Periconia epilithographicola	MFLUCC21-0153	OL606144	OL753687	OL606155	OL912948
Periconia festucae	CGMCC 3.23929	OP956042	OP955973	OP955998	OP961439
Periconia floridana	CBS 150884		NR_197917	NG_244042
Periconia genistae	CBS 322.79	N/A	MH861213	MH872980	N/A
Periconia homothallica	CBS 139698	AB797275	AB809645	NG_059397	AB808541
Periconia igniaria	CBS 298.66	N/A	MH858798	MH870438	N/A
Periconia igniaria	CBS 583.66	N/A	MH858888	MH870553	N/A
Periconia imperatae	UESTCC 22.0145	OP956048	OP955979	OP956004	OP961445
Periconia kunmingensis	HKAS:102239	OR225814	MH892346	MH892400	MH908963
Periconia lateralis	CBS 292.36	N/A	MH855804	MH867311	N/A
Periconia linzhiensis	HKAS 144526*	PQ675368	PQ684989	PQ675408	PQ671465
Periconia linzhiensis	HKAS 144525*	PQ675369	PQ684990	PQ675409	PQ671466
Periconia macrospinosa	CPC 22898	KP184080	KP183999	KP184038	N/A
Periconia minutissima	MFLUCC 15-0245	N/A	KY794703	KY794707	N/A
Periconia muchuanensis	HKAS 135150	PQ066546	PQ067783	PQ067698	PQ278553
Periconia neobrittanica	CBS 146062	N/A	MN562149	MN567656	N/A
Periconia neominutissima	CBS 149514	N/A	NR_189522	NG_242103	N/A
Periconia palmicola	MFLUCC 14-0400	MN648319	N/A	MN648327	MN821070
Periconia penniseti	CGMCC 3.23928	OP956040	OP955971	OP955996	OP961437
Periconia philadelphiana	CPC 42854	N/A	OQ628486	OQ629068	N/A
Periconia prolifica	CBS 209.64	N/A	NR_160097	MH870050	N/A
Periconia pseudobyssoides	UESTCC 22.0135	OP956034	OP955965	OP955990	OP961431
Periconia pseudobyssoides	UESTCC 22.0147	OP956044	OP955975	OP956000	OP961441
Periconia pseudodigitata	CBS 139699	AB797274	LC014591	AB807564	AB808540
Periconia sahariana	CBS 320.79	N/A	MH861211	MH872978	N/A
Periconia salina	MFLU 19-1235	MN017912	MN047086	MN017846	N/A
Periconia shannanensis	HKAS 134945	PP968558	PP968552	PP968555	PP226770
Periconia sichuanensis	CGMCC 3.25598	PQ066545	PQ067867	PQ067697	PQ278551
Periconia spodiopogonis	CGMCC 3.23932	OP956032	OP955963	OP955988	OP961429
Periconia spodiopogonis	HKAS 135635*	PQ675370	PP952765	N/A	N/A
Periconia submersa	MFLUCC 16-1098	N/A	KY794702	KY794706	KY814761
Periconia thailandica	MFLUCC 17-0065	KY753889	KY753887	KY753888	N/A
Periconia thysanolaenae	KUMCC 20-0262	NG_081407	MW442967	MW444850	MW460896
Periconia variicolor	CBS 120374	N/A	DQ336713	N/A	N/A
Periconia verrucosa	UESTCC 22.0136	OP956035	OP955966	OP955991	OP961432
Periconia wurfbainiae	ZHKUCC 23-0999	PP277726	OR995586	OR995592	PP025972
Periconia yangjiangensis	ZHKUCC 23-0997	PP277724	OR995584	OR995590	PP025970
Massarina cisti	CBS 266.62	FJ795490	LC014568	AB807539	AB808514
Massarina pandanicola	MFLU:18-0004	NG_065725	MG646958	NG_064488	MG646986

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Table 6.

GenBank accession numbers of the taxa in this study of Tetraplosphaeriaceae.

Taxon	Voucher	GenBank accession numbers
Taxon	Voucher	SSU		ITS	LSU
Aquatisphaeria thailandica	MFLUCC 21-0025	MW890967	MW890969		MW890763
Aquatisphaeria thailandica	DLUCC B151	MW890968	N/A		MW890764
Byssolophis sphaerioides	IFRDCC 2053	GU296140	N/A		GU301805
Ernakulamia cochinensis	MFLUCC 18-1237	MT864326	MT627670		MN913716
Ernakulamia krabiensis	MFLUCC 18-0237	MK347880	MK347773		MK347990
Ernakulamia tanakae	NFCCI 4615	N/A	MN937229		MN937211
Neotriplosphaeria yadongensis	HKAS 144527*	PQ675371	PQ684991		PQ675410
Neotriplosphaeria yadongensis	HKAS 144528*	PQ675372	PQ684992		PQ675411
Parapolyplosphaeria thailandica	MFLU 15-3273	N/A	KU248766		KU248767
Polyplosphaeria fusca	KT1616	AB524463	AB524789		AB524604
Polyplosphaeria guizhouensis	GZCC 23-0598		OR427327		OR438888
Polyplosphaeria hainanensis	GZCC 23-0599	OR438285	OR427323		OR438889
Polyplosphaeria nabanheensis	KUMCC 16-0151	MH260352	MH275078		MH260312
Polyplosphaeria pandanicola	KUMCC 17-0180	MH260353	MH275079		MH260313
Pseudopolyplosphaeria guizhouensis	GZCC 19-0247	OR134445	N/A		N/A
Pseudotetraploa bambusicola	CGMCC 3.20939	ON332923	ON332915		ON332933
Pseudotetraploa curviappendiculata	HC:4930	AB524467	AB524792		AB524608
Pseudotetraploa longissima	HC:4933	AB524471	AB524796		AB524612
Pseudotetraploa rajmachiensis	NFCCI 4618	N/A	MN937222		MN937204
Pseudotetraploa yunnanensis	KUNCC 10464	N/A	OR449073		OR438891
Quadricrura bicornis	CBS 125427	AB524472	AB524797		AB524613
Quadricrura meridionalis	CBS 125684	AB524473	AB524798		AB524614
Shrungabeeja aquatica	MFLUCC 18-0664	N/A	MT627722		MT627663
Shrungabeeja fluviatilis	GZCC 20-0505	OP377989	OP377804		OP377903
Shrungabeeja longiappendiculata	BCC 76463	KT376471	KT376474		KT376472
Shrungabeeja vadirajensis	MFLUCC 17-2362	N/A	MT627681		MN913685
Tetraploa aquatica	MFLU 19-0995	N/A	MT530448		MT530452
Tetraploa aristata	CBS 996.70	AB524486	AB524805		AB524627
Tetraploa bambusae	KUMCC 21-0844	ON077073	ON077078		ON077067
Tetraploa cylindrica	KUMCC 20-0205	MT893203	MT893205		MT893204
Tetraploa cylindrica	ZHKUCC 22-0087	ON555690	ON555689		ON555688
Tetraploa dashaoensis	KUMCC 21-0010	OL473556	OL473549		OL473555
Tetraploa dwibahubeeja	NFCCI 4621	N/A	MN937226		MN937208
Tetraploa endophytica	CBS 147114	N/A	KT270279		MW659165
Tetraploa hainanensis	GZCC 23-0601	OR438286	OR427325		OR438892
Tetraploa juncicola	CBS 149046	N/A	ON603780		ON603800
Tetraploa lignicola	KUNCC 10794	ON422300	ON422286		ON422294
Tetraploa *linzhiensis*	HKAS 144535*	PQ675373	PQ684993		PQ675412
Tetraploa linzhiensis	HKAS 144536*	PQ675374	PQ684994		PQ675413
Tetraploa nagasakiensis	KT 1682	AB524489	AB524806		AB524630
Tetraploa obpyriformis	KUMCC 21-0011	OL473557	OL473558		OL473554
Tetraploa oryzae	MFLU 23-0195	OR458352	OR438372		OR438841
Tetraploa palmae	HKAS 115638	PP639245	N/A		PP621116
Tetraploa phoenicis	HKAS 115675	PP639246	PP592486		PP621117
Tetraploa pseudoaristata	NFCCI 4624	N/A	MN937232		MN937214
Tetraploa puzheheiensis	KUMCC 20-0151	N/A	MT627744		MT627655
Tetraploa sasicola	KT 563	AB524490	AB524807		AB524631
Tetraploa scheueri	CY112	N/A	HQ607964		N/A
Tetraploa sp.	HKAS 131566	N/A	PP768923		PP768935
Tetraploa thailandica	MFLUCC 21-0030	MZ413274	MZ412518		MZ412530
Tetraploa thrayabahubeeja	NFCCI 4627	N/A	MN937235		MN937217
Tetraploa wurfbainiae	MHZU 23-0231	OR626043	OR626039		OR626041
Tetraploa yakushimensis	KT 1906	AB524491	AB524808		AB524632
Tetraploa yunnanensis	MFLUCC 19-0319	MT864341	MT627743		MN913735
Triplosphaeria acuta	KT 1170	AB524492	AB524809		AB524633
Triplosphaeria cylindrica	KT 1800	AB524494	AB524810		AB524635
Triplosphaeria maxima	KT 870	AB524496	AB524812		AB524637
Murispora aquatica	MFLU 19-0990	MN325077	MN325085		MN325075
Murispora cardui	MFLU 15-2248	NG_063589	NR_171721		KT709176

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Table 7.

GenBank accession numbers of the taxa in this study of Torula.

Taxon	Voucher	GenBank accession numbers
Taxon	Voucher	ITS	LSU	SSU	rpb2	tef1
Torula acaciae	CPC 29737	KY173471	KY173560	N/A	KY173594	N/A
Torula aquatica	MFLUCC 16-1115	MG208167	MG208146	N/A	MG207977	N/A
Torula aquatica	DLUCC 0550	MG208166	MG208145	N/A	MG207976	MG207996
Torula breviconidiophora	KUMCC 18-0130	MK071670	MK071672	MK071697	N/A	MK077673
Torula calceiformis	HKAS 125551	OP751055	OP751053	OP751051	OQ630510	OQ630512
Torula calceiformis	HKAS 125552	OP751054	OP751052	OP751050	OQ630511	OQ630513
Torula camporesii	KUMCC 19-0112	MN507400	MN507402	MN507401	MN507404	MN507403
Torula chiangmaiensis	KUMCC 16-0039	MN061342	KY197856	KY197863	N/A	KY197876
Torula chinensis	UESTCC 22-0085	OQ127986	OQ128004	OQ127995	N/A	N/A
Torula chromolaenae	KUMCC 16-0036	MN061345	KY197860	KY197867	KY197873	KY197880
Torula chromolaenae	MFLUCC 20-0237	MW412524	MW412518	MW412515	MW422161	MW422158
Torula dingjieensis	HKAS 144529*	PQ684995	PQ675414	PQ675375	PQ671471	PQ671467
Torula dingjieensis	HKAS 144530*	PQ684996	PQ675415	PQ675376	PQ671472	PQ671468
Torula fici	CBS 595.96	KF443408	KF443385	KF443387	KF443395	KF443402
Torula fici	KUMCC 16-0038	MN061341	KY197859	KY197866	KY197872	KY197879
Torula gaodangensis	MFLUCC 17-0234	MF034135	NG_059827	MF034134	N/A	N/A
Torula goaensis	MTCC 12620	NR_159045	NG_060016	N/A	N/A	N/A
Torula herbarum	CPC 24414	KR873260	KR873288	N/A	N/A	N/A
Torula hollandica	CBS 220.69	NR_132893	NG_064274	KF443389	KF443393	KF443401
Torula hydei	KUMCC 16-0037	MN061346	MH253926	MH253928	N/A	N/A
Torula lancangjiangensis	HKAS 112709	NR_175706	NG_081516	NG_078759	MW729780	MW729785
Torula lancangjiangensis	B03	MZ538529	MZ538563	N/A	N/A	MZ567104
Torula longan	ZHKUCC 22-0121	OR194035	OR194027	OR194032	OR228535	OR228537
Torula longan	ZHKUCC 22-0122	OR194036	OR194028	OR194033	OR228536	OR228538
Torula luguhuensis	KUNCC 22-12427	OQ729758	OQ947766	N/A	OQ999002	OQ999004
Torula mackenziei	MFLUCC 13-0839	MN061344	KY197861	KY197868	KY197874	KY197881
Torula mackenziei	HKAS 112705	MW723058	MW879525	MW774581	N/A	N/A
Torula masonii	CBS 245.57	NR_145193	NG_058185	N/A	N/A	N/A
Torula masonii	MFLUCC 20-0239	MW412523	MW412517	MW412514	MW422160	MW422157
Torula phytolaccae	ZHKUCC 22-0107	ON611796	ON611800	ON611798	ON660879	ON660881
Torula phytolaccae	ZHKUCC 22-0108	ON611795	ON611799	ON611797	ON660878	ON660880
Torula pluriseptata	MFLUCC 14-0437	MN061338	KY197855	KY197862	KY197869	KY197875
Torula polyseptata	MFLUCC 17-1495	MT214382	MT214476	MT214427	MT235830	MT235791
Torula sichuanensis	UESTCC 22-0087	OQ127981	OQ127999	OQ127990	N/A	N/A
Torula snbmersa	UESTCC 22-0086	OQ127985	OQ128003	OQ127994	N/A	N/A
Torula suae	CGMCC 3.24259	OP359406	OP359415	OP369300	OP476730	OP471618
Torula sundara	UESTCC 22-0125	OQ127984	OQ128002	OQ127993	N/A	N/A
Torula sundara	UESTCC 22-0088	OQ127983	OQ128001	OQ127992	N/A	N/A
Torula sundara	MFLU 21-0089	OM276824	OM287866	N/A	N/A	N/A
Torula thailandica	MFLU:19-2856	MN907426	MN907427	MN907428	N/A	N/A
Torula yadongensis	HKAS 144533*	PQ684997	PQ675416	PQ675377	PQ671473	PQ671469
Torula yadongensis	HKAS 144534*	PQ684998	PQ675417	PQ675378	PQ671474	PQ671470
Cylindrotorula indica	NFCCI:4836	MT339444	MT339442	N/A	MT321490	MT321492
Cylindrotorula indica	NFCCI:4837	MT339445	MT339443	N/A	MT321491	MT321493

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2.3. Phylogenetic analyses

The raw sequences were assembled using SeqMan (Katoh et al. 2005). Comparison with GenBank data was conducted to identify their close relatives. The assembled sequences were compared with GenBank data to identify their closest related taxa. Each gene matrix was aligned separately using MAFFT v. 6.8 (Katoh et al. 2005). The aligned datasets were then manually edited with BioEdit v. 7.0.9 (Hall 1999). SequenceMatrix v. 1.7.8 (Vaidya et al. 2011) was used to combine these datasets. The combined alignment was subsequently used for maximum likelihood (ML) and Bayesian inference (BI) analyses.

The ML analysis was performed using RAxML-HPC2 on XSEDE v. 8.2.10 on the CIPRES Science Gateway v. 3.3 (Miller et al. 2010) with default parameters. The adjustments were made by setting the bootstrap iterations to 1,000, and the substitution model was defined as GTR+GAMMA+I. MrModeltest v. 2.3 was utilised to determine the optimal model for each gene. The BI analysis was conducted using MrBayes v. 3.2.6 (Ronquist et al. 2012). Six simultaneous Markov Chain Monte Carlo (MCMC) chains ran for 500,000 generations, and trees were sampled every 1,000th generation until the standard deviation of the split frequencies dropped below 0.01. The first 25% of generations were discarded as the burn-in phase (Huelsenbeck and Ronquist 2001). The remaining ones were used to calculate Posterior probabilities (PP). The resulting phylogenetic tree was visualised using FigTree v. 1.4.0 and further edited using Adobe Illustrator 2020 and Adobe Photoshop CC 2019.

3. Results

Kirschsteiniotheliales Hern-Restr., R.F. Castañeda, Gené & Crous

Kirschsteiniotheliaceae Boonmee & K.D. Hyde

For Kirschsteiniothelia, 56th taxa were used (Table 2) to assess the taxonomic placement of new isolates using combined three loci (SSU: 1–926 bp; LSU: 927–1,714 bp; ITS: 1,715–2,168 bp), with Homortomyces combreti and H. tamaricis (Wijayawardene et al. 2014) as the outgroup. The ML and BI methods were used to obtain the phylogenetic results (Figure 1). The total character number of combined sequences was 2,168 including gaps. The matrix had 909 distinct alignment patterns, with 22.74% of undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.229197, C = 0.252822, G = 0.299630, T = 0.218352; substitution rates: AC = 1.435134, AG = 2.861902, AT = 0.965679, CG = 1.486506, CT = 6.142113, GT = 1.000000; gamma distribution shape parameter α = 0.532956. The best-scoring RAxML tree had a final likelihood value of −15,870.140997. The ML tree topology was similar to the one inferred from BI analysis. Our specimens Kirschsteiniothelia linzhiensis (HKAS 144539 & HKAS 144540) and K. yadongensis (HKAS 144543 & HKAS 144544) formed distinct monophyletic clades with K. aqutica and K. phoenicis with support value of (100% ML/1.00 PP), respectively.

Figure 1. — Maximum likelihood tree of *Kirschsteiniothelia* from a four-locus dataset (ITS, SSU, and LSU). RAxML bootstrap support values (ML ≥ 70%) and Bayesian posterior probability (PP ≥ 0.90) are shown at the nodes (ML/PP). The scale bar indicates 0.05 changes per site. *Homortomyces combreti* (CPC 19808) and *H. tamaricis* (MFLUCC 13-0441) were selected as outgroup taxa. The type strains are in bold while the newly generated sequences are indicated in red.

Kirschsteiniothelia D. Hawksw., Bot. J. Linn. Soc. 91: 182 (1985)

Kirschsteiniothelia was introduced by Hawksworth (1985) and was initially classified under Pleosporaceae with Kirschsteiniothelia aethiops as the type species, which was later moved to Pleomassariaceae by Barr (1987) based on morphology. Molecular analyses by Kodsueb et al. (2006) further indicated that K. aethiops did not belong to Pleosporaceae. Later, Boonmee et al. (2012) proposed a new family, Kirschsteiniotheliaceae (Pleosporales), based on morphological and phylogenetic analyses (ITS, SSU, and LSU) to accommodate Kirschsteiniothelia (Boonmee et al. 2012). Subsequently, Kirschsteiniothelia was placed in the newly proposed order Kirschsteiniotheliales (Dothideomycetes) by Hernández-Restrepo et al. (2017). Morphologically, the sexual morph of Kirschsteiniothelia is characterised by brown, semi-immersed, subglobose ascomata, cylindrical-clavate, bitunicate, 8-spored asci, brown ascospores with smooth walls and usually have 1–2-septate (Hawksworth 1985; Yang et al. 2010; Xu et al. 2023a; Louangphan et al. 2024), and the asexual morph is characterised by macronematous, branched or unbranched, erect, septate, conidiophores, terminal, determinate, monophialidic, conidiogenous cells and cylindric-obclavate, septate, brown to dark brown conidia (Boonmee et al. 2012; Bao et al. 2018). The genus Kirschsteiniothelia currently comprises 43 species (Senanayake et al. 2023; Xu et al. 2023a; Louangphan et al. 2024; Tang et al. 2024). The members of Kirschsteiniothelia are commonly saprobic on decaying wood (Senanayake et al. 2023). Kirschsteiniothelia species are globally distributed and found in a wide range of habitats across both temperate and tropical regions (Suetrong et al. 2009; Boonmee et al. 2012; Jayawardena et al. 2022). In this study, we introduce two new species from the Xizang Autonomous Region, China, based on phylogenetic and morphological analyses.

Kirschsteiniothelia linzhiensis S.C. He, Q. Zhao & K.D. Hyde, sp. nov. Figure 2

MycoBank number: MB857898; Facesoffungi number: FoF17072.

Etymology – Referring to the collecting site of the type specimen, Linzhi City, Xizang Autonomous Region, China.

Description – Saprobic on unidentified decaying wood. Asexual morph: Hyphomycetous. Colonies effuse, scattered, erect, and brown. Conidiophores 77–200 × 6–8 μm ( $\overset{ˉ}{x}$ = 140 × 7 μm, n = 20), macronematous, mononematous, erect, simple, straight or flexuous, unbranched, cylindrical, smooth, thick-walled, septate, pale brown to brown. Conidiogenous cells monoblastic, terminal, cylindrical, brown conidia developing at the apex. Conidia 31–43 × 6–10 μm ( $\overset{ˉ}{x}$ = 36 × 8 μm, n = 30), solitary, acrogenous dry, obovoid, guttulate, thick-walled, 3–4-septate, straight or curved, pale brown to brown, with a gelatinous sheath at the apex. Sexual morph: Not observed.

Culture characteristics – germinating within 24 h on PDA at 25 °C, slow growing, reaching 1.1–1.3 cm after 10 days incubation, above colony grey, reverse black, entire, convex, surface smooth, dense, no pigment.

Host – On unknown substrate.

Known distribution – China, Xizang Autonomous Region, Linzhi City, Chayu County.

Material examined – China, Xizang Autonomous Region, Linzhi City, Chayu County (28°54′N, 96°98′E, 1,878 m), on unknown wood, 13 August 2023, Shucheng He, WZY20 (KUN-HKAS 144539, holotype) – ex-type, KUNCC24-18549.

Notes – Kirschsteiniothelia linzhiensis formed a monophyletic clade K. aquatica with 100% ML and 1.00 PP support. Morphologically, K. linzhiensis resembles K. aquatica in having macronematous, simple, straight or flexuous, unbranched, thick-walled, smooth, septate conidiophores, and solitary, septate conidia (Bao et al. 2018). However, K. linzhiensis differs from K. aquatica in having obovoid, guttulate, and pale brown to brown conidia with a gelatinous sheath (Bao et al. 2018). The nucleotide differences for ITS, LSU, and SSU between K. linzhiensis and K. aquatica are 36/762 (4.7%, without gaps), 6/750 (0.8%, without gaps), and 5/740 (0.7%, without gaps), respectively. Based on morphological and phylogenetic analyses, we propose a new species K. linzhiensis from Linzhi City, Xizang Autonomous Region, China.

Kirschsteiniothelia yadongensis S.C. He, Q. Zhao & K.D. Hyde, sp. nov. Figure 3

MycoBank number: MB857899; Facesoffungi number: FoF17073.

Etymology – Referring to the collecting site of the type specimen, Yadong County, Shigatse City, Xizang Autonomous Region, China.

Description – Saprobic on decaying wood. Asexual morph: Hyphomycetous. Colonies immersed on the substrate, effuse, composed of septate, unbranched, brown to black. Conidiophores 92–146 × 4–8.5 μm ( $\overset{ˉ}{x}$ = 120 × 7 μm, n = 20), macronematous, mononematous, erect, simple, straight or flexuous, unbranched, smooth, thick-walled, septate, pale brown to brown. Conidiogenous cells monoblastic, terminal, cylindrical, brown, terminal developing into conidia. Conidia 61–97 × 16–21 μm ( $\overset{ˉ}{x}$ = 77 × 19 μm, n = 20), solitary, acrogenous, simple, obclavate, smooth, guttulate, thick-walled, 6–8-septate, brown to black brown, lighter apex with a hayline gelatinous sheath. Sexual morph: Not observed.

Culture characteristics – germinating within 24 h on PDA at 25 °C, reaching 2.5–3.0 cm after 20 days incubation, above and reverse both black with white margin, filamentous, raised, surface smooth, medium dense, no pigment.

Host – On unknown substrate.

Known distribution – China, Xizang Autonomous Region, Shigatse City, Yadong County.

Material examined – China, Xizang Autonomous Region, Shigatse City, Yadong County (27°63′N, 88°92′E, 3,707 m), on unknown wood of a deciduous tree, 29 July 2023, Shucheng He, ZYW119 (KUN-HKAS 144543, holotype) – ex-type, KUNCC24-18543.

Notes – Phylogenetically, K. yadongensis formed a monophyletic clade with K. phoenicis (92% ML/1.00 PP, Figure 1). Kirschsteiniothelia phoenicis, which is saprobic on Phoenix paludosa (Arecaceae) and was reported from Thailand, was described by Hyde et al. (2018) based on its sexual morphic characteristics and phylogenetic results (ITS, SSU, and LSU). Because Kirschsteiniothelia phoenicis only reports the sexual morph (Hyde et al. 2018), and our isolates only report the asexual morph, there is no morphological comparison. It is distinguishable K. yadongensis from K. phoenicis based on morphology (Hyde et al. 2018). However, K. yadongensis differs from K. phoenicis by 49/510 (9.6%, without gaps) ITS, 11/810 (1.3%, without gaps) LSU, and 13/805 (1.6%, without gaps) SSU differences in the nucleotides comparisons. Thus, we propose a new species K. yadongensis from Shigatse City, Xizang Autonomous Region, China.

Astrosphaeriellaceae Phook. & K.D. Hyde

The SSU+LSU+tef1 dataset includes sequences from 38 samples representing 31 species in the GenBank database (Table 3). Murispora kazachstanica and Murispora bromicola are selected as outgroups following Crous et al. (2021). After comparison, the LSU, SSU, and tef1 were assembled (SSU = 1–908 bp; LSU = 909–1,689 bp; tef1 = 1,690–2,561 bp), and the phylogenetic trees of ML and BI were constructed. The phylogenetic analysis results showed that the total number of combined sequence sites was 2,561 including gaps. The matrix had 654 distinct alignment patterns, with 17.40% of undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.244259, C = 0.245957, G = 0.280553, T = 0.229231; substitution rates: AC = 1.245609, AG = 3.886107, AT = 1.345370, CG = 1.060643, CT = 13.468140, GT = 1.000000; gamma distribution shape parameter α = 0.426349. The best-scoring RAxML tree had a final likelihood value of −11,241.554985. Triseptatospora yadongensis (HKAS 134939 and HKAS 134939) formed a distinct clade with T. calami with support value of (86% ML/0.96 PP, Figure 4).

Figure 4. — Maximum likelihood tree of Astrosphaeriellaceae from a three-locus dataset (SSU, LSU, and *tef*1). RAxML bootstrap support values (ML ≥ 70%) and Bayesian posterior probability (PP ≥ 0.90) are shown at the nodes (ML/PP). The scale bar indicates 0.03 changes per site. *Murispora kazachstanica* (CBS 148424) and *M. bromicola* (MFLUCC 15-0031) were selected as outgroup. The type strains are in bold while the newly generated sequences are indicated in red.

Triseptatospora Konta & K.D. Hyde. Mycosphere 14(1): 117 (2023)

Triseptatospora (Astrosphaeriellaceae, Pleosporales), a monotypic genus, was introduced by Konta et al. (2023) to accommodate the type species Triseptatospora calami. Triseptatospora was collected from Calamus sp. in the terrestrial habitat of Thailand. Morphologically, only the sexual morph has been reported. The genus is characterised by scattered to gregarious, superficial, raised, subglobose to dome-shaped, clustered, black, ostiolate ascomata, 6–8-spored, bitunicate, short pedicels asci, fusiform, 3-septate, hyaline, guttulate, mucilaginous sheath ascospores (Konta et al. 2023).

Triseptatospora yadongensis S.C. He, Jayaward. & Q. Zhao, sp. nov. Figure 5

MycoBank number: MB 857900; Facesoffungi number: FoF17074.

Etymology – Referring to the collecting site of the type specimen, Yadong County, China.

Description – Saprobic on unknown wood. Asexual morph: Hyphomycetous. Synnemata 270–430 × 20–27 μm ( $\overset{ˉ}{x}$ = 360 × 24 μm, n = 10), on host surface, synnematous, synnemata composed of conidiophores, erect, solitary, straight, terminal branched, closely compacted hyphae, with brown to dark brown stems, terminal developing into conidiogenous cells and conidia. Conidiophores 280–340 × 2.8–3.5 μm ( $\overset{ˉ}{x}$ = 325 × 3 μm, n = 10), straight, unbranched, smooth, thick-walled, rough-walled, septate, brown, terminal developing into conidiogenous cells. Conidiogenous cells 14–28 × 10–17 μm ( $\overset{ˉ}{x}$ = 19 × 13 μm, n = 10), monoblastic, integrated, terminal, determinate, connected at the base of conidia, brown. Conidia 55–71 × 32–50 μm ( $\overset{ˉ}{x}$ = 63 × 40 μm, n = 20), dictyospores, solitary, acrogenous, simple, papillate, rugose, thick-walled, rough-walled, 4–6 transverse septa, 7–9 longitudinal septa, yellow-brown, constricted at the septa. Sexual morph: Not observed.

Culture characteristics – Conidia germinating within 20 h on PDA at 25 °C, reaching 2.5–3 cm after 20 days incubation, colony surface pale olivaceous-grey, reverse grey green, dense, circular, slightly raised, smooth, entire, hairy, wrinkled folded, pigmentation not produced on PDA.

Host – On unknown substrate.

Known distribution – China, Xizang Autonomous Region, Shigatse City, Yadong County.

Material examined – China, Xizang Autonomous Region, Shigatse City, Yadong County (27°23’N, 89°01′E, 1,687 m), 25 July 2023, Shucheng He, CT20 (KUN-HKAS 134939, holotype) – ex-type: KUNCC 24-17789. ibid. Shucheng He, CT24 (KUN-HKAS 134940, paratype) – ex-paratype: KUNCC24-17790.

Notes – The isolates formed a monophyletic clade with T. calami (Figure 4). Morphologically, our collections have distinct synnemata, which were first reported in Astrosphaeriellaceae (Liu et al. 2018; Wanasinghe et al. 2018; Dong et al. 2020; Crous et al. 2021; Luo et al. 2022; Konta et al. 2023). The isolates are similar to Kostermansinda in morphology, have synnematous and dictyospores, acrogenous, papillate conidia, however, the conidia of Kostermansinda have multiple (12–13 vs. 7–9) longitudinal septa (Boedijn 1960; Rifai 1968). The phylogenetic position of Kostermansinda has not been resolved as they lack DNA-based sequence data. Comparison of LSU, mtSSU, rpb2, and tef1 sequences in the NCBI database revealed that they belong to the orders Lecanorales, Monoblastiales, and Tubeufiales, respectively (Hosoya and Yano 2020). Species in Triseptatospora are known only by sexual morphs (Konta et al. 2023), the nucleotide differences of SSU and tef1 between our isolates and Triseptatospora calami are 3/793 (0.4%, without gaps) and 47/956 (5%, without gaps), respectively (Konta et al. 2023). Based on this evidence, following Chethana et al. (2021), we have retained our isolate in Triseptatospora and propose a new species, T. yadongensis, from Shigatse City, Xizang Autonomous Region, China.

Dictyosporiaceae Boonmee & K.D. Hyde

The combined ITS, LSU, SSU, and tef1 datasets comprise 40 taxa, with Periconia igniaria (Su et al. 2023) as the outgroup taxa (Table 4). The dataset consisted of 3,967 total characters, including gaps (LSU: 1–826 bp, SSU: 827–1,751 bp, ITS: 1,752–2,271 bp, tef1: 2,272–3,099 bp). The matrix had 818 distinct alignment patterns, with 21.01% of undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.243068, C = 0.241628, G = 0.269153, T = 0.246152; substitution rates: AC = 1.684401, AG = 3.847836, AT = 2.349320, CG = 0.793813, CT = 9.946496, GT = 1.000000; gamma distribution shape parameter α = 0.457026. The best-scoring RAxML tree with a final likelihood value of −15,213.255591 is presented in Figure 6. The tree topology obtained from ML analysis is similar to the one inferred from BI analysis. Based on the phylogenetic analyses of the combined ITS, LSU, and SSU sequence data, the novel species Trichobotrys motuoensis (HKAS 144531 and HKAS 144532) is closely related to T. sinensis with 100% ML/0.97 PP statistical support (Figure 6).

Figure 6. — Maximum likelihood tree of Dictyosporiaceae reconstructed from a four-locus dataset (LSU, SSU, ITS, and *tef*1). RAxML bootstrap support values (ML ≥ 70%) and Bayesian posterior probability (PP ≥ 0.90) are shown at the nodes (ML/PP). The scale bar indicates 0.03 changes per site. *Periconia igniaria* (CBS 379.86 and CBS 845.96) was selected as outgroup. The type strains are in bold while the newly generated sequences are indicated in red.

Trichobotrys Penz. & Sacc., Malpighia 15(7–9): 245 (1902) [1901]

Trichobotrys was introduced by Penzig and Saccardo (1902) to accommodate the type species Tr. pannosus. Later, Tr. pannosus was treated as a synonym of Tr. effusa (Morgan-Jones et al. 1987; D’Souza and Bhat 2001). Trichobotrys has eight epithets in Index Fungorum (2025, accessed on 7 May 2025) and among them, only four species (Tr. effusa, Tr. meilingensis, Tr. sinensis, and Tr. yunjushanensis) have been molecularly confirmed (Phookamsak et al. 2024; Zhang et al. 2024). The genus is known solely by its asexual morph, which is characterised by solitary or clustered in a small group, scattered to gregarious, globose to subglobose, ostiolate, minute papilla conidiomata, dark brown to black, effuse colonies, macronematous, mononematous, brown, verruculose or echinulate conidiophores, smooth, fertile, often unciform lateral branches, with sterile, setiform apex, polyblastic, integrated, terminal or discrete, ellipsoidal, spherical or subspherical conidiogenous cells and catenated, spherical, brown, aseptate, verruculose or echinulate conidia (D’Souza and Bhat 2001; Phookamsak et al. 2024; Zhang et al. 2024). Species of Trichobotrys were reported in China and Java, from both aquatic and terrestrial habitats, where they are saprobic on Arecaceae and Poaceae (Brachiaria mutica and Bamboo) (D’Souza and Bhat 2001; Phookamsak et al. 2024; Zhang et al. 2024).

Trichobotrys motuoensis S.C. He, Q. Zhao & K.D. Hyde, sp. nov. Figure 7

MycoBank number: MB857901; Facesoffungi number: FoF17075.

Etymology – Referring to the collecting site of the type specimen, Motuo County, China.

Description – Asexual morph: Hyphomycetous. Colonies on the Synthetic Low Nutrient Agar (SNA), effuse, branched, dark brown, hairy. Conidiophores 800–1,100 μm long, 3–5 μm wide ( $\overset{ˉ}{x}$ = 4 μm, n = 20), dichotomously branched, mononematous, simple, flexuous, branched or occasionally loosely branched, verruculose, thick-walled, rough-walled, echinulate, septate, pale brown to brown. Conidiogenous cells 14–20 × 4–4.6 μm ( $\overset{ˉ}{x}$ = 17 × 4 μm, n = 20), polyblastic, integrated, terminal, hyaline to pale brown, verruculose, thick-walled, denticulate. Conidia 5–6 × 5–6 μm ( $\overset{ˉ}{x}$ = 5.5 × 5.5 μm, n = 30), catenate, acropleurogenous, solitary, simple, dry, verruculose, echinulate, globose, thin-walled, smooth-walled, aseptate, brown. Sexual morph: Not observed.

Culture characteristics – on PDA at 25 °C, reaching 6.0–6.2 cm after 45 days incubation, above colony pale yellow, reverse yellow edge with a black centre, irregular, raised, umbonate, surface rough, dense, no pigment.

Host – On unknown substrate.

Known distribution – China, Xizang Autonomous Region, Linzhi City, Motuo County.

Material examined – China, Xizang Autonomous Region, Linzhi City, Motuo County (29°25′N, 95°20′E, 792 m), on unidentified substrate, 6 July 2022, Shucheng He, HSC944 (KUN-HKAS 144531, holotype) – ex-type – KUNCC24-18551.

Notes – The phylogenetic tree (LSU, SSU, ITS, and tef1) showed that Tr. motuoensis was clustered with Tr. sinensis and Tr. meilingensis (Figure 6). Morphologically, Tr. motuoensis resembles Tr. sinensis and Tr. meilingensis in having acropleurogenous, verruculose, echinulate, globose conidia. However, Tr. motuoensis has larger conidia compared to Tr. sinensis (5.5–5.5 µm vs. 2.8–2 µm), and smaller conidia compared to Tr. meilingensis (5.5–5.5 µm vs. 7–13 µm) (Phookamsak et al. 2024; Zhang et al. 2024). Trichobotrys sinensis and Tr. meilingensis have both been found in aquatic environments in China. Trichobotrys sinensis is saprobic on Brachiaria mutica (Poaceae), while Tr. meilingensis is saprobic on decaying bamboo (Phookamsak et al. 2024; Zhang et al. 2024). The nucleotide differences for ITS, LSU, SSU, and tef1 between Tr. motuoensis and Tr. sinensis are 6/480 (1.2%, without gaps), 2/770 (0.3%, without gaps), 2/763 (0.3%, without gaps), and 15/870 (1.7%, without gaps), respectively. Similarly, the nucleotide differences for ITS, LSU, SSU, and tef1 between Tr. motuoensis and Tr. meilingensis are 6/501 (1.2%, without gaps), 2/800 (0.2%, without gaps), 3/752 (0.4%, without gaps), and 11/774 (1.4%, without gaps), respectively. Therefore, we introduce Tr. motuoensis as a new species, from Motuo County, Xizang Autonomous Region, China.

Periconiaceae Nann.

The combined LSU, SSU, ITS, and tef1 dataset comprises 68 taxa, with Massarina cisti (Tanaka et al. 2015) and Massarina pandanicola (Tibpromma et al. 2018) designated as the outgroup taxon (Table 5). The dataset consists of 3,118 total characters including gaps (LSU: 1–823 bp, SSU: 824–1,770 bp, tef1: 1,771–2,618 bp, ITS: 2,619–3,118 bp). The matrix had 802 distinct alignment patterns, with 25.80% of undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.236633, C = 0.256358, G = 0.270666, T = 0.236343; substitution rates: AC = 1.797398, AG = 2.963500, AT = 2.015171, CG = 1.254214, CT = 10.715473, GT = 1.000000; gamma distribution shape parameter α = 0.515216. The best-scoring RAxML tree had a final likelihood value of −15,110.354168. Both the ML and BI analyses yielded similar tree topologies, and only the tree inferred from ML analysis is shown (Figure 8). Our isolates (HKAS 144526 and HKAS 144527) formed a well-supported, distinct clade with Periconia didymosporum (100% ML/1.00 PP, Figure 8). In addition, we identify P. spodiopogonis as a new host record for China.

Figure 8. — Maximum likelihood tree of *Periconia* reconstructed from a four-locus dataset (ITS, SSU, LSU, and *tef*1). RAxML bootstrap support values (ML ≥ 70%) and Bayesian posterior probability (PP ≥ 0.90) are shown at the nodes (ML/PP). The scale bar indicates 0.02 changes per site. *Massarina cisti* (CBS 266.62) and *M. pandanicola* (MFLU 18-0004) were selected as outgroup taxa. The type strains are in bold while the newly generated sequences are indicated in red.

Periconia Tode, Fung. mecklenb. sel. (Lüneburg) 2: 2 (1791)

Periconia (Periconiaceae, Pleosporales) was introduced by Tode (1791) with P. lichenoides as the type species. This genus primarily comprises species known for their asexual morphs, characterised by macronematous, mononematous conidiophores, and globose to ellipsoidal conidia (Phookamsak et al. 2019; Hongsanan et al. 2020; Liao et al. 2024). The sexual morph is characterised by immersed to erumpent, scattered or aggregated, subglobose to globose ascomata, hyaline, 8-spored, bitunicate, cylindrical, septate, guttulate, ascospores with an entire sheath (Tanaka et al. 2015; Yang et al. 2022; Su et al. 2023; Sun et al. 2025). Periconia species are mostly saprophytes and endophytes, though some are plant pathogens such as P. circinata and P. macrospinosa (Kolomiets et al. 2008; Sarkar et al. 2019) and are widely distributed in terrestrial habitats. The genus includes 234 records (Index Fungorum 2025, accessed on 7 May 2025). These fungi have a cosmopolitan distribution and inhabit various environments, including soil, decaying plant material, and living plant tissues. Notable species include P. byssoides, P. digitate, and P. minutissima, found on diverse hosts such as grasses, legumes, and daisies (Hyde et al. 2020; Crous et al. 2023).

Periconia linzhiensis S.C. He, Q. Zhao & K.D. Hyde, sp. nov. Figure 9

MycoBank number: MB857902; Facesoffungi number: FoF17076.

Etymology – Referring to the collecting site of the type specimen, Linzhi City, Xizang Autonomous Region, China.

Description – Saprobic on dead bamboo. Asexual morph: Hyphomycetous. Colonies on the substrate, effuse, erect, brown to dark brown, hairy. Conidiophores 315–427 × 16–21 μm ( $\overset{ˉ}{x}$ = 355 × 18 μm, n = 20), macronematous, mononematous, erect, simple, straight or flexuous, unbranched, thick and smooth-walled, septate, brown to dark brown. Conidiogenous cells polyblastic, terminal, subglobose, smooth to verruculose, hyaline to brown. Conidia 14–24 × 16–25 μm ( $\overset{ˉ}{x}$ = 21 × 20 μm, n = 30), catenate, acrogenous, globose, simple, simple, verruculose, thick-walled, rough-walled, aseptate, pale brown when young; dark brown when mature. Sexual morph: Not observed.

Culture characteristics – Germinating within 24 h on PDA at 25 °C, reaching 5.5–6.0 cm after 30 days incubation, above colony brown black with grey centre; reverse brown, circular undulate, umbonate, surface rough, mycelia dense, no pigment.

Host – Bamboo.

Known distribution – China, Xizang Autonomous Region, Linzhi City, Motuo County.

Material examined – China, Xizang Autonomous Region, Linzhi City, Motuo County (29°35′N, 95°14′E, 1,730 m), on bamboo, 7 August 2023, Shucheng He, SBB30 (KUN-HKAS 144526, holotype) – ex-type, KUNCC25-19123.

Notes – Periconia linzhiensis formed a monophyletic clade with P. didymosporum with 100% ML/1.00 PP statistical support (Figure 8). Periconia linzhiensis and P. didymosporum were recorded from terrestrial environments and are saprophytic on Bamboo (Yang et al. 2022). However, P. didymosporum (= Bambusistroma didymosporum) was reported based on the sexual morph (Yang et al. 2022). The asexual morph of Periconia linzhiensis was reported. Periconia linzhiensis differed from P. didymosporum by 16/510 (3.2%, without gaps) ITS, 5/750 (0.6%, without gaps) LSU, 1/750 (0.1%, without gaps) SSU, and 7/810 (0.8%, without gaps) tef1 differences in the nucleotides. Thus, we propose P. linzhiensis as a new species.

Periconia spodiopogonis Y.P. Chen & Z.H. Lu, Journal of Fungi 9: 18 (2023) Figure 10

MycoBank number: MB847469; Facesoffungi number: FoF17077.

Description – Saprobic on dead bamboo. Asexual morph: Hyphomycetous. Colonies on the substrate, effuse, brown to dark brown, hairy. Conidiophores 430–720 × 16–27 μm ( $\overset{ˉ}{x}$ = 590 × 22 μm, n = 20), macronematous, mononematous, erect, simple, straight or flexuous, branched at the apex, thick-walled, smooth-walled, septate, dark brown. Conidiogenous cells polyblastic, integrated, terminal, subglobose, smooth to verruculose, brown. Conidia 17–22 × 11–15 μm ( $\overset{ˉ}{x}$ = 20 × 12.6 μm, n = 30), acrogenous, ellipsoidal to cylindrical, simple, verruculose, guttulate, thick-walled, rough-walled, aseptate, pale brown to dark brown. Sexual morph: Not observed.

Material examined – China, Xizang Autonomous Region, Linzhi City, Motuo County (29°17′N, 95°15′E, 1,550 m) on bamboo, 9 August 2023, Shucheng He, SBB04 (HKAS 135635).

Known distribution – China (Su et al. 2023).

Known hosts – Spodiopogon (Su et al. 2023), Bamboo (this study).

Notes – In the combined phylogenetic analyses (LSU, ITS, and SSU), our isolate was grouped together with P. spodiopogonis, having 100% ML/1.00 PP statistical support. Periconia spodiopogonis was initially described as a saprobe on Spodiopogon in China (Su et al. 2023). There were no differences in the ITS and SSU base sequences between our isolate and ex-type strain of P. spodiopogonis. Morphologically, our isolate (Figure 10) shares similar characteristics with P. spodiopogonis in conidiophores (590 × 22 vs. 595 × 15 μm) that are appressed to the substrate, macronematous, mononematous, erect, straight or flexuous and the conidia (20 × 12.6 vs. 20 × 12.5 μm) are acrogenous, ellipsoidal to cylindrical, verruculose, guttulate and aseptate (Su et al. 2023). Thus, based on phylogenetic analysis and morphological characteristics, we identified our isolate as P. spodiopogonis, which is a new host record in China.

Tetraplosphaeriaceae Kaz. Tanaka & K. Hiray

In order to clarify the taxonomic status of four taxa, based on these four gene fragments, 58 fungal sequences of Tetraplosphaeriaceae were downloaded from the GenBank databases (Table 6), with Murispora aquatica and Murispora cardui as the outgroup (Bao et al. 2019). According to the LSU-SSU-ITS sequence data, the three genes were spliced (LSU = 1–821 bp; SSU = 822–1,747 bp; ITS = 1,748–2,289 bp). The total number of combined sequence sites is 2,289 including gaps. The matrix had 586 distinct alignment patterns, with 17.18% of undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.247845, C = 0.241156, G = 0.282371, T = 0.228628; substitution rates: AC = 4.006945, AG = 4.409444, AT = 2.264925, CG = 1.075482, CT = 11.088939, GT = 1.000000; gamma distribution shape parameter α = 0.563830. The best-scoring RAxML tree had a final likelihood value of −11,459.593875. The ML tree topology was similar to the one inferred from BI analysis. The ML tree topology was similar to the one inferred from BI analysis. In the phylogenetic tree, each genus formed a separate clade, Neotriplosphaeria yadongensis forming a monophyletic clade in Tetraplosphaeriaceae, our isolates formed a monophyletic clade with Tetraploa nagasakiensis (Figure 11). Tetraplosphaeriaceae was established by Tanaka et al. (2009) to accommodate Polyplosphaeria, Pseudotetraploa, Quadricrura, Tetraplosphaeria (the type genus), and Triplosphaeria. It is characterised by massarina-like asexual morphs, immersed to superficial, glabrous, brown, cylindrical to clavate ascomata, and fusiform to cylindrical, narrowly, 8-spored asci, septate, hyaline to pale brown, with appendage-like sheath ascospores (Tanaka et al. 2009; Tian et al. 2024). The anamorphs of Tetraplosphaeriaceae are tetraploa-like hyphomycetes, micronematous, macronematous, erect, septate conidiophores, monoblastic, terminal conidiogenous cells, cylindrical to obpyriform, comprising 3–8 columns, verrucose, with 2–8-setose conidia (Tanaka et al. 2009). Based on morphological and molecular evidence (ITS, LSU, and SSU), Ariyawansa et al. (2015) introduced Shrungabeeja to Tetraplosphaeriaceae. Subsequently, Aquatisphaeria, Byssolophis, and Ernakulamia were introduced into the Tetraplosphaeriaceae (Delgado et al. 2017; Pem et al. 2019; Li et al. 2021). Recently, Zhang et al. (2023) introduced a new genus, Pseudopolyplospha, into Tetraplosphaeriaceae, which now consists of 10 genera (Zhang et al. 2023; Hyde et al. 2024b).

Figure 11. — Maximum likelihood tree of Tetraplosphaeriaceae reconstructed from a three-locus dataset (LSU, SSU, and ITS). RAxML bootstrap support values (ML ≥ 70%) and Bayesian posterior probability (PP ≥ 0.90) are shown at the nodes (ML/PP). The scale bar indicates 0.03 changes per site. *Murispora aquatica* (MFLU 19-0990) and *Murispora cardui* (MFLU 15-2248) were selected as outgroup taxa. The type strains are in bold while the newly generated sequences are indicated in red.

Neotriplosphaeria S.C. He, Q. Zhao & K.D. Hyde, gen. nov.

MycoBank number: MB857903; Facesoffungi number: FoF17078.

Etymology – Named after its conidial similarity to Triplosphaeria.

Description – Saprobic on dead stems of bamboo. Asexual morph: Colonies effuse, medium dense, hairy, black. Conidiophores absent. Conidiogenous cells mononematous, monoblastic, integrated, terminal or intercalary, determinate, and cylindrical. Conidia solitary, septate, brown to dark brown, ovoid to obclavate, consisting of 2–4 columns of cells, multi-septate in each column, smooth, thick-walled, mostly with 2 to 4 apical appendages. Appendages cylindrical, solitary, unbranched, guttulate, septate, wide at the base, pale brown to brown, multi-septate, straight or flexuous, smooth-walled.

Neotriplosphaeria yadongensis S.C. He, Q. Zhao & K.D. Hyde, sp. nov. Figure 12

MycoBank number: MB857904; Facesoffungi number: FoF17079.

Etymology – Refers to the collecting site of the type specimen, Yadong County, China.

Description – Saprobic on dead stems of bamboo. Asexual morph: Colonies effuse, medium dense, hairy, black. Conidiophores absent. Conidiogenous cells mononematous, monoblastic, integrated, terminal or intercalary, determinate, cylindrical. Conidia 42–60 × 18–27 μm ( $\overset{ˉ}{x}$ = 49 × 22 μm, n = 30), solitary, septate, brown to dark brown, ovoid to obclavate, consisting of 2–4 columns of cells, 6–9-septate in each column, smooth, thick-walled, mostly with 2–4 apical appendages. Appendages 33–74 × 3–4 μm ( $\overset{ˉ}{x}$ = 55 × 3.5 μm, n = 30), cylindrical, solitary, unbranched, guttulate, 4–6-septate, wide at the base, pale brown to brown, straight or flexuous, smooth-walled. Sexual morph: Not observed.

Host – Bamboo.

Known distribution – China, Xizang Autonomous Region, Shigatse City, Yadong County.

Material examined – China, Xizang Autonomous Region, Shigatse City, Yadong County (27°13′N, 80°00′E, 1,824 m), on bamboo, 24 July 2023, Shucheng He, CT10 (KUN-HKAS 144527, holotype).

Notes – Neotriplosphaeria yadongensis formed a distinct clade within Tetraplosphaeriaceae. Its morphology fits well with the generic concept of Tetraplosphaeriaceae with cylindrical to obpyriform, comprising 3–8 columns, verrucose, with 2–8 appendages conidia (Tanaka et al. 2009). Morphologically, the conidia of Neotriplosphaeria consist of 2–4 columns of cells with 2–4 apical appendages, while the conidia of Triplosphaeria consist of only three columns of cells with three apical appendages (Tanaka et al. 2009). Neotriplosphaeria has ovoid to obclavate conidia with 2–4 appendages of similar length, while Quadricrura has globose to subglobose conidia with long appendages that are usually 1–2 appendages, and short that are 4–5 appendages (Tanaka et al. 2009). Based on phylogenetic and morphological analyses, we propose a new genus, Neotriplosphaeria, within Tetraplosphaeriaceae to accommodate the new species N. yadongensis from Yadong County, Xizang Autonomous Region, China.

Tetraploa Berk. & Broome, Ann. Mag. nat. Hist., Ser. 2 5: 459 (1850)

Tetraploa was introduced by Berkeley and Broome (1850) and was later assigned as the type genus of Tetraplosphaeriaceae (Tanaka et al. 2009). Tetraploa is the largest genus in Tetraplosphaeriaceae (Tang et al. 2023; Zhang et al. 2023), with 44 epithets currently listed in Index Fungorum (accessed on 25 August 2024). The sexual morph is characterised by immersed to erumpent, globose to subglobose, glabrous ascomata, fissitunicate, cylindrical to clavate, short-stalked asci with eight ascospores, and narrowly fusiform, hyaline, smooth ascospores with a narrow mucilaginous appendage-like sheath (Tanaka et al. 2009). The main characteristics of the asexual morph are cylindrical, brown, verrucose, aseptate conidia with four setose appendages at the apex (Tanaka et al. 2009; Tang et al. 2023; Zhang et al. 2023). Species of Tetraploa were commonly found as endophytes and saprobes on plant leaves and stems in both terrestrial and aquatic environments (Tanaka et al. 2009; Ariyawansa et al. 2015; Delgado et al. 2017; Pem et al. 2019; Hongsanan et al. 2020). They were documented on bamboo, other herbaceous plants, and in decaying wood and soil (Dong et al. 2020; Li et al. 2021). Notably, Te. aristata was reported as a pathogen affecting a variety of plants and humans (Markham et al. 1990; Hyde et al. 2013).

Tetraploa linzhiensis S.C. He, Q. Zhao & K.D. Hyde, sp. nov. Figure 13

MycoBank number: MB857905; Facesoffungi number: FoF17080.

Etymology – Referring to the collecting site of the type specimen, Linzhi city, China.

Description – Saprobic on dead stems of Elsholtzia sp. Asexual morph: Hyphomycetous Colonies effuse, sparse, black. Conidiophores absent. Conidiogenous cells monoblastic, integrated, determinate. Conidia 25–45 × 19–30 μm ( $\overset{ˉ}{x}$ = 39 × 23 μm, n = 30), solitary, cylindrical to obovate, septate, brown to dark brown, consisting of 1–3 columns of cells, 3–6-septate in each column, verruculose, thick-walled, rough-walled, with 1–3 apical appendages. Appendages 180–320 × 1–1.2 μm ( $\overset{ˉ}{x}$ = 260 × 1.1 μm, n = 30), cylindrical, solitary, unbranched, wide at the base, pale brown to brown, hyaline at the apex, 6–9-septate, straight, guttulate, smooth-walled, long appendages, short appendages 102–188 × 7–9 μm ( $\overset{ˉ}{x}$ = 137 × 8 μm, n = 30). Sexual morph: Not observed.

Culture characteristics – Germinating within 24 h on PDA at 25 °C, reaching 1.8–2 cm after 15 days incubation, above colony grey, reverse dark grey with black edge, entire, undulate, umbonate, surface smooth, mycelia dense, no pigment.

Host – Elsholtzia sp. (Lamiaceae).

Known distribution – China, Xizang Autonomous Region, Linzhi City, Chayu County.

Material examined – China, Xizang Autonomous Region, Linzhi City, Chayu County (28°54′N, 96°98′E, 1,878 m), on Elsholtzia sp., 13 August 2023, Shucheng He, ZYW49 (KUN-HKAS 144535, holotype) – ex-type culture in KUNCC24-18547.

Notes – The phylogenetic tree showed Te. linzhiensis was clustered with Te. cylindrica and Te. nagasakiensis with 100% ML and 1.00 PP support (Figure 11). Morphologically, Te. linzhiensis resembles Te. cylindrica and Te. nagasakiensis in having cylindrical to obovate, brown, multi columns of cells, verruculose, thick-walled, rough-walled conidia (Tanaka et al. 2009; Hyde et al. 2013). However, Te. linzhiensis mostly consists of three columns of cells and three appendages conidia, while Te. cylindrica and Te. nagasakiensis mostly consist of four columns of cells and four appendages conidia (Tanaka et al. 2009; Hyde et al. 2013). Tetraploa cylindrica and Te. nagasakiensis are both saprobic on Poaceae (Sasa kurilensis and bamboo) in Japan (Tanaka et al. 2009). The nucleotide differences for ITS, LSU, and SSU between Te. linzhiensis and Te. cylindrica are 10/505 (2%, without gaps), 3/748 (0.4%, without gaps), and 3/755 (0.4%, without gaps), respectively. Similarly, the nucleotide differences for ITS, LSU, and SSU between Te. linzhiensis and Te. nagasakiensis are 18/490 (3.6%), 2/732 (0.3%), and 3/782 (0.4%), respectively. Therefore, based on phylogenetic and morphological analyses, we propose a new species, Te. linzhiensis, from Linzhi City, Xizang Autonomous Region, China.

Torulaceae Corda

The combined ITS, LSU, SSU, tef1, and rpb2 dataset comprised 43 taxa, with Cylindrotorula indica (Boonmee et al. 2021) as the outgroup taxa (Table 7). The dataset consisted of 3,920 total characters, including gaps (LSU: 1–806 bp, SSU: 807–1,715 bp, ITS: 1,716–2,214 bp, tef1: 2,215–3,080 bp, rpb2: 3,081–3,920 bp). The matrix had 842 distinct alignment patterns, with 26.55% of undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.247197, C = 0.259213, G = 0.270214, T = 0.223376; substitution rates: AC = 2.116465, AG = 4.272528, AT = 1.684889, CG = 1.309028, CT = 9.872550, GT = 1.000000; gamma distribution shape parameter α = 0.521756. The best-scoring RAxML tree with a final likelihood value of −14,127.258641 is presented in Figure 14. The tree topology obtained from ML analysis is similar to the one inferred from BI analysis. The best-scoring RAxML tree is presented in (Figure 14). Our specimens Torula dingjieensis (HKAS 144529 & HKAS 144530) and T. yadongensis (HKAS 144533 & HKAS 144534) formed distinct monophyletic clades with T. fici and T. calceiformis with support value of 87% ML/0.92 PP and 95% ML/0.97 PP, indicating they are closely related.

Figure 14. — Maximum likelihood tree of *Torula* reconstructed from a five-locus dataset (LSU, SSU, ITS, *tef*1, and *rpb*2). RAxML bootstrap support values (ML ≥ 70%) and Bayesian posterior probability (PP ≥ 0.90) are shown at the nodes (ML/PP). The scale bar indicates 0.03 changes per site. *Cylindrotorula indica* (NFCCI 4836 and NFCCI 4837) was selected as outgroup. The type strains are in bold while the newly generated sequences are indicated in red.

Torula Pers., Ann. Bot. (Usteri) 15: 25 (1795)

Torula was introduced by Persoon (1795) with the type species of To. herbarum. Most species of Torula were found in terrestrial and freshwater habitats on submerged wood or dead branches of Asteraceae, Cyperaceae, and Ranunculaceae in Asia, Europe, North America, and other regions (Crous et al. 2015; Hyde et al. 2017; Hongsanan et al. 2020; Jayawardena et al. 2022). Torula is the biggest genus in Torulaceae with 542 epithets recorded in Index Fungorum (24 November 2024). However, only 30 species were molecularly confirmed (Hyde et al. 2023; Tian et al. 2023; Wang et al. 2023a). Torula was documented solely in its asexual morph (Jayawardena et al. 2022; He et al. 2024b).

Torula dingjieensis S.C. He, Q. Zhao & K.D. Hyde, sp. nov. Figure 15

MycoBank number: MB857906; Facesoffungi number: FoF17081.

Etymology – Referring to the collecting site of the type specimen, Dingjie County, China.

Description – Saprobic on dead stems of Ageratina adenophora. Asexual morph: Colonies effuse, medium dense, black. Conidiophores semi-macronematous, mononematous, straight or flexuous, unbranched or occasionally loosely branched, verruculose, thick-walled, rough-walled, doliiform to subcylindrical, septate consisting of 1–4 cells or reduced to conidiogenous cells, pale brown to brown. Conidiogenous cells polyblastic, terminal, pale brown to brown, verruculose, thick-walled, doliiform to cupulate. Conidia 11–29 × 6–22 μm ( $\overset{ˉ}{x}$ = 20 × 9 μm, n = 30), phragmospores, solitary, simple, dry, acrogenous, monilioid, verruculose, thick-walled, rough-walled, 0–4-septate, brown to dark brown. Sexual morph: Not observed.

Culture characteristics – germinating within 24 h on PDA at 25 °C, reaching 5.8–6.2 cm after 25 days incubation, above colony brown, reverse dark brown, both above and reverse with pale yellow edge, entire, undulate, umbonate, surface rough, mycelia dense, no pigment.

Host – Ageratina adenophora (Asteraceae).

Known distribution – China, Xizang Autonomous Region, Shigatse City, Dingjie County.

Material examined – China, Xizang Autonomous Region, Shigatse City, Dingjie County (27°54′N, 87°22′E, 2,240 m), on Ageratina adenophora, 4 July 2022, Shucheng He, HSC820 (KUN-HKAS 144529, holotype) – ex-type, KUNCC23-17478.

Notes – The phylogenetic tree showed that our isolates (HKAS 144529 and HKAS 144530) To. dingjieensis clustered with To. calceiformis (Figure 14). Torula calceiformis is saprobic on unidentified wood form Guizhou, China (Hyde et al. 2023). Our isolates (HKAS 144529 and HKAS 144530) were collected from Ageratina adenophora in Shigatse, Xizang Autonomous Region, China. Morphologically, To. dingjieensis resembles To. calceiformis in having semi-macronematous, mononematous, verruculose conidiophores, phragmospores, and monilioid, verruculose, multi-septate conidia. Compared to To. dingjieensis, To. calceiformis has conidia with additional septa (0–8-septate vs. 0–4-septate) and longer conidia (L/W ratio: 3 vs. 2.2) (Hyde et al. 2023). The nucleotide differences for ITS, LSU, SSU, tef1, and rpb2 between To. dingjieensis and To. calceiformis are 7/480 (1.5%, without gaps), 3/750 (0.4%, without gaps), 2/807 (0.2%, without gaps), 5/710 (0.7%, without gaps), and 21/885 (2.4%, without gaps), respectively (Hyde et al. 2023). Therefore, based on phylogenetic and morphological analyses, we propose a new species, To. dingjieensis, from Shigatse City, Xizang Autonomous Region, China.

Torula yadongensis S.C. He, Q. Zhao & K.D. Hyde, sp. nov. Figure 16

MycoBank number: MB857907; Facesoffungi number: FoF17082.

Etymology – Referring to the collecting site of the type specimen, Yadong County, China.

Description – Saprobic on dead stems of Artemisia sp. Asexual morph: Colonies effuse on nature substrate, medium dense, scattered, velutinous, black. Conidiophores macronematous, mononematous, straight or flexuous, branched of conidia chain, verruculose, thick-walled, rough-walled, doliiform to subcylindrical, septate, terminal reduced to conidiogenous cells, conidia chain hyaline or golden to dark brown. Conidiogenous cells monoblastic, terminal, pale brown to dark brown, verruculose, thick-walled, rough-walled, doliiform to cupulate. Conidia 8.4–14.6 × 4.3–6.2 μm ( $\overset{ˉ}{x}$ = 12 × 6 μm, n = 30), didymospores, solitary, simple, pleuroacrogenous, monilioid, verruculose, thick-walled, rough-walled, 0–2-septate, golden to dark brown. Sexual morph: Not observed.

Culture characteristics – Germinating within 24 h on PDA at 25 °C, reaching 3.8–4.0 cm after 20 days incubation, both above and reverse white, entire, flat, surface smooth, mycelia dense, no pigment.

Host – Artemisia sp. (Asteraceae).

Known distribution – China, Xizang Autonomous Region, Shigatse City, Yadong County.

Material examined – China, Xizang Autonomous Region, Shigatse City, Yadong County (27°29′N, 88°54′E, 2,960 m), on Artemisia sp., 3 July 2022, Shucheng He, HSC720 (KUN-HKAS 144533, holotype) – ex-type KUNCC23-17229.

Notes – The new species was isolated from Artemisia sp. in Yadong County, China. Phylogenetic analyses (LSU-SSU-ITS-tef1-rpb2) showed To. yadongensis is closely related to To. fici, which was collected from leaves of Ficus religiosa (Moraceae) in Cuba. The phylogenetic tree showed To. yadongensis and To. fici were close relatives with 87% ML/0.92 PP support (Figure 14). Torula yadongensis resembles To. fici with intact or cupulate conidiogenous cells that produce chains of rough-walled conidia (He et al. 2024c). Torula yadongensis has shorter (L/W ratio: 2 vs. 3.6) conidia than To. fici. The nucleotide differences for ITS, LSU, SSU, tef1, and rpb2 between To. yadongensis and To. fici were 10/489 (2%, without gaps), 21/780 (2.7%, without gaps), 5/991 (0.5%, without gaps), 18/798 (2.2%, without gaps), and 10/808 (1.2%, without gaps), respectively (Crous et al. 2015). Therefore, based on phylogenetic and morphological analyses, we propose a new species, To. yadongensis, from Yadong County, Xizang Autonomous Region, China.

4. Discussion

The Xizang Autonomous Region, China, is known for its extreme environmental conditions such as low temperatures, high UV radiation, and low oxygen levels (Chen et al. 2020; Wang et al. 2021). This provides a distinct habitat fostering the evolution of novel fungal taxa (Phurbu et al. 2024). In this study, we collected 19 specimens from the Xizang Autonomous Region, China, and identified a new genus, nine new species, and one new host record. Based on morphological characteristics and phylogenetic analyses, these taxa have been placed in two orders of Dothideomycetes: Astrosphaeriellaceae, Periconiaceae, Tetraplosphaeriaceae, Torulaceae, and one incertae sedis genus in Pleosporales (Zhang et al. 2012; Hongsanan et al. 2020), and Kirschsteiniotheliaceae in Kirschsteiniotheliales (Hernández-Restrepo et al. 2017).

Triseptatospora (Astrosphaeriellaceae, Pleosporales) was introduced by Konta et al. (2023). Species of Astrosphaeriellaceae were mainly found as sexual morph (Kuo and Goh 2019; Konta et al. 2023). Triseptatospora yadongensis has distinct synnemata, which is the key feature distinguishing it from other species in the Astrosphaeriellaceae (Liu et al. 2018; Wanasinghe et al. 2018; Dong et al. 2020; Crous et al. 2021; Luo et al. 2022; Konta et al. 2023). However, in the phylogenetic analysis, based on the genetic distance was not sufficient to provide strong evidence for the delineation of a new genus (Figure 4). Therefore, we retained our isolates in Triseptatospora, and for the first time, introduced a species from the Xizang Autonomous Region, China, into the genus (Konta et al. 2023), and proposed a new species, T. yadongensis. Periconia is the largest genus in Periconiaceae (Hyde et al. 2024b). Earlier members of this genus were placed in Massarinaceae. Based on phylogenetic analysis, Tanaka et al. (2015) suggested that Periconiaceae is a family distinct from Massarinaceae and Cai et al. (2024) first reported Periconia species in the Xizang Autonomous Region, China, based on morphology and phylogenetic analyses. In this study, we also introduce a new species and a new host record from the region. Tetraplosphaeriaceae was established by Tanaka et al. (2009) and consists of 10 genera (Zhang et al. 2023; Hyde et al. 2024b). In this study, based on phylogenetic and morphological analyses, we introduce a new genus, Neotriplosphaeria, to accommodate N. yadongensis, and a new species, Te. linzhiensis. Torula, is the most speciose genus in the Torulaceae with 542 epithets (He et al. 2024c). However, most species are not supported by molecular data (He et al. 2024c). Torulaceae currently has six genera introduced, namely, Cylindrotorula, Dendryphion, Neopodoconis, Neotorula, Rutola, and Torula (Qiu et al. 2022; Wang et al. 2023b). There is controversy over the division of some genera. Qiu et al. (2022) synonymised Sporidesmioides and Rostriconidium with Neopodoconis, Wang et al. (2023b) accepted Neopodoconis as a synonym of Rostriconidium and pointed out that there is insufficient evidence to synonymize Sporidesmioides with Neopodoconis. He et al. (2024c) proposed Torula longiconidiophora (Tian et al. 2023) as a synonym of Torula sundara (Jayawardena et al. 2022) based on phylogenetic and morphological analysis while investigating the relationships among species in the genus Torula (He et al. 2024c). Species of Torula are mainly distributed in Asia, Europe, North America (Jayawardena et al. 2022; Dong et al. 2023; He et al. 2024c). Both To. dingjieensis and To. yadongensis collected from Xizang Autonomous Region, China, add to the growing evidence that this region has a rich and largely unexplored fungal diversity. Trichobotrys, an incertae sedis genus, was placed in Pleosporales (Hyde et al. 2024b). To date, Trichobotrys includes four species supported by molecular data and morphological evidence, namely Tr. effusus, Tr. meilingensis, Tr. sinensis, and Tr. yunjushanensis (D’Souza and Bhat 2001; Phookamsak et al. 2024; Zhang et al. 2024). We constructed a phylogenetic tree (Figure 6) of Trichobotrys based on five genes (LSU, SSU, ITS, and tef1), and the results showed that Trichobotrys formed a similar topology to Phookamsak et al. (2024) and Zhang et al. (2024), with Gregarithecium having strong statistical support (100% ML/1.00 PP) in Dictyosporiaceae. Therefore, we propose the placement of Trichobotrys in Dictyosporiaceae, which further supports the conclusion proposed by Zhang et al. (2024). Kirschsteiniotheliales is a small order within Dothideomycetes (Hyde et al. 2024b). Species of Kirschsteiniothelia in China are primarily reported in the provinces of Guizhou, Hainan, Sichuan, and Yunnan (Tang et al. 2024; Xiao et al. 2025). In this study, we first report two species of Kirschsteiniothelia from Xizang Autonomous Region, China, including two new species K. linzhiensis and K. yadongensis.

The Xizang Autonomous Region, with its extreme environmental conditions, is a significant but underexplored area for fungal biodiversity. This study focused on microfungi from the Plateau, particularly within the orders Pleosporales and Kirschsteiniotheliales, both of which belong to the class Dothideomycetes. Fungal diversity plays a crucial role in the Xizang Autonomous Region by enhancing biodiversity understanding and ecosystem health (Hyde et al. 2024a; Phurbu et al. 2024; He et al. 2025). Xizang Autonomous Region, China, continues to be a rich source of fungal biodiversity. Further exploration of this unique environment is likely to uncover even more undiscovered fungal diversity.

Funding Statement

This study was supported by the Second Tibetan Plateau Scientific Expedition and Research Program [2024QZKK02010303], the Second Qinghai-Tibet Plateau Scientific Expedition and Research (STEP) Program [2019QZKK0503], the Survey of Wildlife Resources in Key Areas of Tibet [ZL202303601], the Chinese Academy of Sciences Key Technical Talents [2021-5-E12671], the Chinese Research Fund [E1644111K1] titled “Flexible introduction of the high-level expert program, Kunming Institute, Institute of Botany, Chinese Academy of Sciences”, and the Researchers Supporting Project of King Saud University, Riyadh, Saudi Arabia [RSP2025R114].

Disclosure statement

No potential conflict of interest was reported by the author(s).

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PERMALINK

Taxonomic contributions to Pleosporales and Kirschsteiniotheliales from the Xizang Autonomous Region, China

Shu-Cheng He

Kevin D Hyde

Ruvishika S Jayawardena

Vinodhini Thiyagaraja

Dhanushka N Wanasinghe

Yun-Wei Zhao

Zhi-Yang Wang

Tong Cai

Yan-Yan Yang

F Al-Otibi

Zhuliang Yang

Qi Zhao

ABSTRACT

1. Introduction

2. Materials and methods

2.1. Collection, fungal isolation, and morphology

2.2. DNA extraction, PCR amplification, and sequencing

Table 1.

Table 2.

Table 3.

Table 4.

Table 5.

Table 6.

Table 7.

2.3. Phylogenetic analyses

3. Results

Figure 1.

Figure 2.

Figure 3.

Figure 4.

Figure 5.

Figure 6.

Figure 7.

Figure 8.

Figure 9.

Figure 10.

Figure 11.

Figure 12.

Figure 13.

Torulaceae Corda

Figure 14.

Figure 15.

Figure 16.

4. Discussion

Funding Statement

Disclosure statement

References

ACTIONS

PERMALINK

RESOURCES

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