Research Article
Print
Research Article
Assigning Xiuguozhangia (genus incertae sedis) to Pseudoberkleasmiaceae (Dothideomycetes, Pleosporomycetidae, Pleosporales) and introducing X. broussonetiae as a novel species
expand article infoDeecksha Gomdola‡§, Rajesh Jeewon|, Eleni Gentekaki#, Ruvishika S. Jayawardena§, Kevin D. Hyde§, Fatimah Alotibi, Yong Wang¤
‡ Guizhou University, Guizhou, China
§ Mae Fah Luang University, Chiang Rai, Thailand
| Rajesh Jeewon, Réduit, Mauritius
¶ King Saud University, Riyadh, Saudi Arabia
# University of Nicosia School of Veterinary Medicine, Cyprus, Cyprus
¤ Guizhou University, Guiyang, China

Corresponding author: Yong Wang ( yongwangbis@aliyun.com )

Academic editor: Samantha C. Karunarathna
© 2024 Deecksha Gomdola, Rajesh Jeewon, Eleni Gentekaki, Ruvishika S. Jayawardena, Kevin D. Hyde, Fatimah Alotibi, Yong Wang.
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: Gomdola D, Jeewon R, Gentekaki E, S. Jayawardena R, Hyde KD, Alotibi F, Wang Y (2024) Assigning Xiuguozhangia (genus incertae sedis) to Pseudoberkleasmiaceae (Dothideomycetes, Pleosporomycetidae, Pleosporales) and introducing X. broussonetiae as a novel species. MycoKeys 111: 181-210. https://doi.org/10.3897/mycokeys.111.136202
Open Access

Abstract

Xiuguozhangia species are dematiaceous hyphomycetes that are characterised by acropleurogenous, dictyoseptate, campanulate or cheiroid, and brown to dark brown conidia that are composed of several layers of cells radiating from a protuberant basal cell, and mostly seen with appendages arising from the apical cells. The genus was introduced based on morphology to accommodate five of the six Piricaudiopsis species that exhibited holoblastic conidial ontogeny. Xiuguozhangia was referred to as Ascomycota genus incertae sedis as it was challenging to resolve its taxonomic placement based solely on the available morphological data (no DNA sequence data was previously available). In this study, we provide DNA sequence data for LSU, ITS, SSU, TEF1, and RPB2 for our isolates, collected from Broussonetia papyrifera (Moraceae) in northern Thailand. Based on morphology, we classify our isolates as Xiuguozhangia. Since they form a sister lineage to Pseudoberkleasmium, we place Xiuguozhangia in Pseudoberkleasmiaceae (Pleosporales). Within Xiuguozhangia, we establish these two isolates as a new taxon, Xiuguozhangia broussonetiae, in view of the presence of new conidiogenous cells developing from subtending cells. Xiuguozhangia broussonetiae differs from the extant species in the genus as it has longer conidiophores that are sometimes branched, comprising numerous septa, and its appendages are mostly untapered (sometimes tapering) towards the tips, a feature not observed in other Xiuguozhangia species. This is the first study to provide DNA sequence data and phylogenetic relationships for Xiuguozhangia. Furthermore, we analysed selected DNA sequence data and provided an updated phylogenetic tree incorporating all families (with representative genera) of Pleosporales.

Key words

1 new taxon, hyphomycetes, morphology, phylogeny, sequence data, taxonomy

Introduction

Xiuguozhangia was introduced by Zhang et al. (2014) based on morphological characteristics and typified with X. rosae. The genus is characterised by effuse, hairy, and brown to dark brown colonies on natural substrates. Conidiophores are erect, straight or flexuous, multiseptate, and brown to dark brown at the base, becoming pale brown towards the apex. Conidiogenous cells are monoblastic, terminal or integrated, lageniform, and pale brown, displaying zero to several percurrent proliferations. In addition, their conidia are acropleurogenous, dictyoseptate, campanulate, cheiroid and brown, comprising multiple layers of cells, either with or without appendages (Zhang et al. 2014; Liu et al. 2024b).

Initially, five Xiuguozhangia species were established, all of which are combinations of Piricaudiopsis viz., Xiuguozhangia appendiculata, X. indica, X. punicae, X. rhaphidophorae and X. rosae. Piricaudiopsis was established by Mena and Mercado (1987) based on morphology and typified with P. elegans. The new combinations of Xiuguozhangia were proposed based on their holoblastic conidial ontogeny; Piricaudiopsis was reported to have enteroblastic conidial ontogeny (Zhang et al. 2014; Dubey and Jash 2024). However, this morphological difference based on conidial ontogeny is subtle, necessitating careful consideration before applying this trait in generic delineation. Given the limited taxon sampling and analysis across both genera, coupled with the lack of DNA sequence data, it is still uncertain whether Xiuguozhangia is actually distinct from Piricaudiopsis, or if they could represent the same genus. Both genera were referred to as Ascomycota genera incertae sedis in the latest outlines by Wijayawardene et al. (2022a) and Hyde et al. (2024c) as it was difficult to establish their precise taxonomic placement without the availability of DNA sequence data. Recently, based on morphological description, a new species was added to Xiuguozhangia viz., X. macrospora, collected from dead bamboo stems in India (Dubey and Jash 2024). To date, all extant species of Xiuguozhangia have been described as saprobes and collected from China and India (Zhang et al. 2014; Dubey and Jash 2024), while no studies have reported its occurrence in Thailand.

The highlights of this study are as follows: (i) We provide DNA sequence data for multiple gene regions (LSU, ITS, SSU, TEF1, and RPB2) for our isolates, collected from Broussonetia papyrifera (Moraceae) in northern Thailand; (ii) Based on morphology, we classify our isolates as Xiuguozhangia. Since they form a sister lineage to Pseudoberkleasmium, we place Xiuguozhangia in Pseudoberkleasmiaceae (Pleosporales); (iii) Within Xiuguozhangia, these two isolates could potentially represent a new taxon, Xiuguozhangia broussonetiae in view of the presence of new conidiogenous cells developing from subtending cells. This is the first study to provide DNA sequence data and phylogenetic relationships for Xiuguozhangia. In addition, we analysed selected sequence data from GenBank and provided an updated phylogenetic tree incorporating all families (with representative genera) of Pleosporales.

Materials and methods

Sample collection and examination, material deposition, and species delimitation methods

Decaying stems of Broussonetia papyrifera (Moraceae) colonised by hyphomycetes were collected from deciduous forest in the Mae Fah Luang University Botanical Garden, Chiang Rai, Thailand during the dry, hot season; collection information was noted (Rathnayaka et al. 2024) and brought to the laboratory in paper boxes for further characterisation (Senanayake et al. 2020). Morphological characteristics were observed using a Motic SMZ 168 Series stereo-microscope. Several conidia were picked from the colonies on the substrate using a sterile needle and placed directly on fresh potato dextrose agar plates (PDA, 39 g/L) containing antibiotics (Amoxicillin, MacroPhar). Conidium germination was observed within 48 hours, and pure cultures were incubated for four weeks at 25 °C under dark conditions. Digital images of micro-morphological characters were captured using a Cannon 750D camera (Canon, Tokyo, Japan) attached to a Nikon ECLIPSE E600 compound microscope (Nikon, Tokyo, Japan) based on the bright-field microscopy technique. Photo plates were assembled using Adobe Photoshop CS6 version 2020 (Adobe Systems, USA), and measurements were made using Tarosoft® Image Frame Work (version 0.97).

The holotype specimen and ex-type living culture have been deposited in the Mae Fah Luang University Herbarium (MFLU) and Mae Fah Luang University Culture Collection (MFLUCC), respectively. Faces of Fungi and Index Fungorum numbers are given for the new species (Jayasiri et al. 2015; Index Fungorum 2024). The description and illustration of the new species have also been updated in the GMS microfungi database (https://gmsmicrofungi.org/) (Chaiwan et al. 2021).

The new species is established based on the morphological species concept and complemented with the phylogenetic species concept (Maharachchikumbura et al. 2021; Pem et al. 2021). Features of Xiuguozhangia species are compiled, and a comparison is done to showcase the major differences across existing taxa.

DNA extraction, PCR amplification, and sequencing

Total genomic DNA was extracted from axenic cultures grown on PDA for 28 days using the BIOMIGA Fungus Genomic DNA Extraction Kit (BIOMIGA, San Diego, CA, USA), following the manufacturer’s instructions. The internal transcribed spacer (ITS), large subunit (LSU), and small subunit (SSU), and the genes for RNA polymerase 2 (RPB2) and translation elongation factor 1α (TEF1) were amplified using the following primers: ITS1/ITS4, LR0R/LR5, NS1/NS4 for ITS, LSU, and SSU, respectively (White et al. 1990); fRpb2-5F/7CR for RPB2 (Liu et al. 1999); and 728F/2218R for TEF1 (O’Donnell et al. 1998; Carbone and Kohn 1999).

The polymerase chain reaction (PCR) mixture had a final volume of 20 µL, comprising 10 µL of PCR master mix, 1 µL of the forward and reverse primers each (10 µM stock concentration), 7 µL of double-distilled water, and 1 µL of the template DNA. The PCR conditions were as follows: initial denaturation at 95 °C for 3 min; denaturation at 95 °C for 45 s; annealing at 55 °C for 50 s (ITS), 52 °C for 50 s (LSU and SSU), 58 °C for 1 min 30 s (RPB2 and TEF1); extension at 72 °C for 2 min; and final extension at 72 °C for 10 min (number of cycles = 40). Purification and bidirectional sequencing of PCR amplicons were carried out at Sangon Biotech (Shanghai) Co., Ltd., China.

Phylogenetic analyses

The raw reads were checked using DNA Baser Assembler, and ambiguous bases from the 5’ and 3’ ends were trimmed manually. Consensus sequences were generated using SeqMan (DNAStar, Madison, Wisconsin, USA). The sequences have been deposited in the NCBI GenBank database, and accession numbers for all strains are provided (Table 1). Newly obtained sequences were subjected to blast searches in NCBI, and sequences of ITS, LSU, SSU, RPB2, and TEF1 from other species were retrieved from GenBank (Table 1). Two different datasets were analysed in this study. The first dataset (Dataset 1) evaluated familial relationships and was based on a larger taxon sampling, incorporating representative genera with DNA sequence data from 92 families of Pleosporales. Diademaceae and Lizoniaceae, which also belong to Pleosporales, lack molecular data. Another dataset (Dataset 2) evaluated the phylogenetic relationships within and between Xiuguozhangia (Pseudoberkleasmiaceae) and its phylogenetically closely related genera, Pseudoberkleasmium (Pseudoberkleasmiaceae) and Hermatomyces (Hermatomycetaceae). Dataset 2 was based on multiple strains for these genera.

Table 1.
Download as
CSV
XLSX

Names, isolate numbers, and the corresponding GenBank accession numbers of taxa used in the phylogenetic analyses. Type, ex-type, and reference strains are denoted with T. The new isolates are in bold font.

Species Isolate/strain number LSU ITS SSU TEF1 RPB2
Acrocalymma aquatica MFLUCC 11-0208 T JX276952 JX276951 JX276953 – –
Acrocalymma pterocarpi MFLUCC 17 0926 T MK347949 MK347732 MK347840 MK360040 –
Ageratinicola kunmingensis KUMCC 21-0217 T NG_243113 NR_191196 NG_242816 – –
Aigialus grandis BCC 20000 T GU479775 – GU479739 GU479839 –
Alternaria atrobrunnea FMR 16868 T – LR537033 – LR537051 LR537044
Alternaria alternata AFTOL-ID 1610 T DQ678082 KF465761 KC584507 KC584634 KC584375
Amniculicola aquatica MFLUCC 16-1123 T MK106096 – MK106108 MK109800 –
Amorocoelophoma cassiae MFLUCC 17-2283 MK347956 MK347739 NG_065775 MK360041 MK434894
Anastomitrabeculia didymospora MFLU 20-0694 T MW412978 NR_172008 NG_073568 MW411338 –
Anastomitrabeculia didymospora MFLU 11-0236 ON077069 ON077080 ON077074 ON075063 ON075067
Angustimassarina lonicerae MFLUCC 15-0087 KY496724 KY496759 – – –
Anteaglonium gordoniae MFLUCC 17-2431 T MK347977 MK347761 MK347867 MK360042 MK434881
Anteaglonium gordoniae CD7 – OK335788 – – –
Anteaglonium latirostrum GKM1119 T GQ221874 – – GQ221937 –
Anteaglonium latirostrum GKML100Nb GQ221876 – – GQ221938 –
Aquadictyospora clematidis MFLUCC 17-2080 T MT214545 MT310592 MT226664 MT394727 MT394679
Aquastroma magniostiolata HHUF 30122 T AB807510 LC014540 AB797220 AB808486 –
Aquasubmersa japonica HHUF 30469 T NG_057138 NR_154739 NG_062426 LC194384 LC194421
Aquasubmersa japonica MFLUCC 17-2121 OP377971 OP377885 OP378047 OP473059 OP473118
Aquasubmersa japonica MFLUCC 15-0622 OP377958 OP377872 OP378036 OP473051 OP473112
Aquasubmersa mircensis MFLUCC 11-0401 T NG_042699 JX276954 NG_061141 – –
Aquihelicascus songkhlaensis MFLUCC 18-1154 T MN913692 MT627680 – MT954380 –
Aquihelicascus songkhlaensis MFLUCC 18-1273 MN913724 MT627696 MT864319 MT954369 MT878464
Aquimassariosphaeria kunmingensis KUMCC 18-1019 T MT627661 – MT864312 MT954409 MT878454
Ascocylindrica marina MD6011 T KT252905 – KT252907 – –
Ascocylindrica marina MF416 MK007123 – MK007124 – –
Astragalicola vasilyevae MFLUCC 17-0832 T MG828986 NR_157504 MG829098 MG829193 MG829248
Astrosphaeriella fusispora MFLUCC 10-0555 KT955462 – KT955443 KT955425 KT955413
Atrocalyx glutinosus DAOM: 252609 T OQ400928 OQ400918 – OQ413076 OQ413081
Atrocalyx glutinosus CHEM 2721 OQ400930 OQ400920 – – OQ413084
Bahusandhika indica GUFCC 18001 KF460274 KF460273 – – –
Bambusicola bambusae MFLUCC 11-0614 T JX442035 JX442031 JX442039 – KP761718
Berkleasmium aquaticum (Tubeufiales) MFLUCC 17-0049 T KY790432 KY790444 – KY792608 MF535268
Berkleasmium aquaticum (Tubeufiales) MFLUCC 17-0039 KY790431 KY790443 – KY792607 MF535267
Berkleasmium longisporum (Tubeufiales) MFLUCC 17-1999 T MH558825 MH558698 – MH550889 MH551012
Berkleasmium longisporum (Tubeufiales) MFLUCC 17-2002 MH558826 MH558699 – MH550890 MH551013
Bertiella fici MFLU 19-2713 T MW063223 – MW079351 MW183786 –
Bertiella fici NCYU 19-0073 MW063224 – MW079352 MW183787 –
Bipolaris adikaramae HSF070 T – MN535176 – MT548605 –
Boeremia linicola CBS 116.76 T GU237938 GU237754 – KY484705 KT389574
Boeremia linicola CBS 248.38 KT389703 KT389486 – – KT389575
Biatriospora borsei NFCCI-4245 T MK358813 MK358818 MK358811 MK330938 –
Biatriospora marina CY 1228 GQ925848 – GQ925835 GU479848 GU479823
Brevicollum hyalosporum MAFF 243400 T LC271239 LC271242 LC271236 LC271245 LC271249
Brevicollum hyalosporum MFLUCC 17-0071 MG602200 MG602204 MG602202 MG739516 –
Brevicollum versicolor HHUF 30591 T NG_058716 NR_156335 NG_065124 LC271246 LC271250
Brunneoclavispora camporesii MFLUCC 11-0001 T MN809328 MN809329 – – –
Brunneofusispora clematidis MFLUCC 17-2070 T MT214570 MT310615 MT226685 MT394629 MT394692
Camarosporium quaternatum CPC 31081 T NG_064442 NR_159756 KY929123 KY929201 –
Camarosporomyces flavigenus CBS 314.80 T GU238076 MH861266 NG_061093 – –
Camarosporidiella caraganicola MFLUCCC 14-0605 T KP711381 KP711380 KP711382 – –
Camarosporidiella melnikii MFLUCC 17-0684 T MF434250 MF434162 MF434338 MF434425 –
Capulatispora sagittiformis HHUF 29754 T NG_042319 NR_119393 NG_060997 LC001756 –
Caryospora aquatica MFLU 11-1083 T NG_059058 NR_156408 MH057850 – –
Caryospora submersa MFLUCC 18-1283 T MN913720 – – – –
Clematidis italica MFLUCC 15-0084 T KU842381 KU842380 KU842382 – –
Coelodictyosporium rosarum MFLUCC 17-0776 T MG828991 MG828875 MG829102 MG829195 –
Corylicola italica MFLU 19-0500 T MT554926 MT554925 MT554923 – MT590776
Corynespora cassiicola CBS 100822 GU301808 – GU296144 GU349052 GU371742
Corynespora torulosa CPC 15989 T KF777207 NR_145181 – – –
Crassimassarina macrospora MAFF 239606 T LC194344 LC194478 LC194302 LC194389 LC194426
Crassimassarina macrospora HHUF 30512 LC194343 LC194477 LC194301 LC194388 LC194425
Crassiperidium octosporum MAFF 242971 T LC373108 LC373096 LC373084 LC373120 LC373132
Crassiperidium octosporum MAFF 246401 LC373111 LC373099 LC373087 LC373123 LC373135
Cryptocoryneum japonicum HHUF 30482 T NG_059035 NR_153938 NG_065118 LC096144 LC194438
Cryptocoryneum pseudorilstonei CBS 113641 T NG_059036 NR_153941 LC194322 LC096152 LC194446
Cucurbitaria berberidis MFLUCC 11-0387 KC506796 – KC506800 – –
Curvularia austriaca CBS 102694 T – MN688802 – MN688856 –
Curvularia eleusinicola USJCC-0005 T – MT262877 – MT432925 –
Cylindroaseptospora leucaenicola MFLUCC 17-2424 T MK347966 NR_163333 MK347856 – –
Cyclothyriella rubronotata CBS 141486 T KX650544 NR_147651 NG_061252 KX650519 KX650574
Dacampia hookeri GZU 73897 KT383792 – – – –
Dacampia hookeri GZU 81840 KT383795 – – – –
Dacampia hookeri GZU 75980 KT383794 – – – –
Delitschia nypae MFLUCC 17-2588 T – – – MK360049 MK434878
Dendryphion fluminicola MFLUCC 17-1689 T MG208141 NR_157490 – MG207992 –
Deniquelata cassiae CMD012A T OR500088 OR500092 OR500090 OR501827 –
Dictyocheirospora bannica KH 332 T AB807513 LC014543 AB797223 AB808489 –
Didymella exigua CBS 183.55 T MH868977 MH857436 GU296147 – –
Didymella rumicicola CBS 683.79 T MH873007 KT389503 – – KT389622
Dothidotthia robiniae MFLUCC 16-1175 T MK751817 MK751727 MK751762 MK908017 MK920237
Epicoccum duchesneae CGMCC 3.18345 T KY742249 KY742095 – – MT018115
Epicoccum duchesneae CBS 218.81 MN973322 MN972935 – – MN983572
Falciformispora aquatica MFLUCC 18-0212 T MK063643 MK064216 – MK099811 –
Falciformispora tompkinsii CBS 200.79 T MH872968 MH861199 KF015639 KF015685 KF015719
Fenestella crataegi CBS 144857 T – NR_165534 – MK357555 MK357512
Fissuroma calami MFLUCC 13-0836 T MF588993 – NG_062430 MF588975 –
Flammeascoma lignicola MFLUCC 10-0128 T KT324583 KT324582 KT324584 KT324585 KT324586
Flavomyces fulophazii CBS 135761 T NG_058131 NR_137960 NG_061191 – –
Foliophoma fallens CBS 161.78 GU238074 KY940772 GU238215 – KC584502
Fuscostagonospora cytisi MFLUCC 16-0622 T KY770978 – KY770977 KY770979 –
Fuscostagonospora sasae HHUF 29106 T AB807548 AB809636 AB797258 AB808524 –
Fusculina eucalypti CBS 120083 T DQ923531 DQ923531 – – –
Fusculina eucalyptorum CBS 145083 T MK047499 NR_161140 – – –
Fusiformispora clematidis MFLUCC 17-2077 T MT214542 MT310589 MT226661 MT394725 MT394677
Gordonomyces mucovaginatus CMW 22212 T JN712552 JN712486 – – –
Halojulella avicenniae BCC 20173 T GU371822 – GU371830 GU371815 GU371786
Helminthosporiella stilbacea MFLUCC 15-0813 T MT928157 MT928159 MT928161 MT928151 –
Hermatomyces amphisporus CBS 146610 T LR812664 LR812664 – – –
Hermatomyces amphisporus CBS 146613 LR812662 LR812662 – LR812657 LR812668
Hermatomyces amphisporus CBS 146614 LR812666 LR812666 – LR812660 LR812671
Hermatomyces anomianthi MFLUCC 21-0202 T OK655817 OL413437 – OM117546 –
Hermatomyces bifurcatus CCF 5900 T LS398263 LS398263 – LS398417 LS398344
Hermatomyces bifurcatus CCF 5899 LS398262 LS398262 – LS398416 LS398343
Hermatomyces clematidis MFLUCC 17-2085 T MT214556 MT310603 MT226673 MT394735 MT394684
Hermatomyces constrictus CCF 5904 T LS398264 LS398264 – LS398418 LS398345
Hermatomyces hainanensis GZCC 23-0592 T OR091329 OR098708 – – –
Hermatomyces indicus (=H. thailandicus) MFLUCC 14-1143 T KU764692 KU144920 KU712468 – KU712488
Hermatomyces indicus (=H. thailandicus) MFLUCC 14-1144 KU764693 KU144921 KU712469 – KU712489
Hermatomyces indicus (=H. thailandicus) MFLUCC 14-1145 KU764694 KU144922 KU712470 KU872756 KU712490
Hermatomyces iriomotensis KH 361 LC194367 LC194483 – LC194394 LC194449
Hermatomyces jinghaensis HKAS 112167 T MW989519 MW989495 – MZ042642 –
Hermatomyces krabiensis (=H. chiangmaiensis) MFLUCC 16-0249 T KX525742 KX525750 – – KX525754
Hermatomyces krabiensis (=H. chiangmaiensis) MFLUCC 16-2819 KY559394 – – – –
Hermatomyces maharashtraensis NFCCI 4879 T NG_241939 NR_189384 – MZ130659 MZ130660
Hermatomyces megasporus CCF 5898 T LS398266 LS398266 – LS398420 –
Hermatomyces megasporus CCF 5897 – LS398265 – LS398419 LS398346
Hermatomyces nabanheensis KUMCC 16-0149 T KY766059 KY766058 KY766060 KY766061 –
Hermatomyces pyriformis CGMCC 3.27462 T PP491962 PP491964 – PP505452 PP505454
Hermatomyces pyriformis UESTCC 23.0441 PP491963 PP491965 – PP505453 PP505455
Hermatomyces reticulatus (=H. subiculosus) MFLUCC 15-0843 T KX259523 KX259521 KX259525 KX259527 KX259529
Hermatomyces reticulatus (=H. subiculosus) CCF 5893 LS398267 LS398267 – LS398421 LS398347
Hermatomyces sphaericoides CCF 5908 T LS398273 LS398273 – LS398427 LS398352
Hermatomyces sphaericoides CCF 5895 LS398270 LS398270 – LS398424 LS398350
Hermatomyces sphaericus PMA 116080 LS398281 LS398281 – LS398431 LS398356
Hermatomyces sphaericus PMA 116081 LS398283 LS398283 – LS398432 LS398357
Hermatomyces sphaericus PRC 4105 – LS398286 – – –
Hermatomyces sphaericus PRC 4104 – LS398278 – LS398430 LS398355
Hermatomyces sphaericus KZP 462 – LS398287 – LS398434 LS398359
Hermatomyces sphaericus PRM 946201 – LS398284 – LS398433 LS398358
Hermatomyces sphaericus PRC 4116 – LS398275 – – –
Hermatomyces sphaericus PRC 4100 LS398277 LS398277 – LS398429 LS398354
Hermatomyces sphaericus PRC 4106 LS398279 LS398279 – – –
Hermatomyces sphaericus PMA 116085 – LS398280 – – –
Hermatomyces sphaericus PMA 116082 – LS398285 – – –
Hermatomyces sphaericus PRC 4117 – LS398276 – – –
Hermatomyces sphaericus MFLUCC 17-0373 OL782061 OL782144 OL780526 – –
Hermatomyces sphaericus HKAS 112725 MW989516 MW989492 – MZ042639 MZ042636
Hermatomyces sphaericus HKAS 112166 MW989517 MW989493 – MZ042640 MZ042637
Hermatomyces sphaericus (=H. biconisporus) KUMCC 17-0183 MH260296 MH275063 MH260338 MH412771 MH412755
Hermatomyces sphaericus (=H. chromolaenae) MFLUCC 16-2818 KY559393 – – – –
Hermatomyces sphaericus (=H. pandanicola) MFLUCC 16-0251 KX525743 KX525751 KX525747 KX525759 KX525755
Hermatomyces sphaericus (=H. saikhuensis) MFLUCC 16-0266 KX525740 KX525748 – KX525756 KX525752
Hermatomyces sphaericus (=H. saikhuensis) MFLUCC 16-0267 KX525741 KX525749 – KX525757 KX525753
Hermatomyces sphaericus (=H. tectonae) MFLUCC 14-1140 KU764695 – NG_063603 KU872757 KU712486
Hermatomyces sphaericus (=H. tectonae) MFLUCC 14-1141 KU764696 KU144918 KU712466 KU872758 –
Hermatomyces sphaericus (=H. tectonae) MFLUCC 14-1142 KU764697 KU144919 KU712467 – KU712487
Hermatomyces trangensis BCC 80741 T KY790600 KY790598 KY790602 KY790606 KY790604
Hermatomyces trangensis BCC 80742 KY790601 KY790599 KY790603 KY790607 KY790605
Hermatomyces tucumanensis CCF 5912 LS398288 LS398288 – LS398435 LS398360
Hermatomyces tucumanensis CCF 5915 LS398290 LS398290 – LS398437 LS398362
Hermatomyces tucumanensis CCF 5913 LS398289 LS398289 – LS398436 LS398361
Hermatomyces turbinatus MFLUCC 21 0038 T MW989518 MW989494 – MZ042641 MZ042638
Hermatomyces verrucosus CCF 5903 T LS398292 LS398292 – LS398439 LS398364
Hermatomyces verrucosus CCF 5892 LS398291 LS398291 – LS398438 LS398363
Hongkongmyces aquaticus MFLUCC 18-1150 T MN913694 – MT864302 MT954379 –
Hypsostroma caimitalense GKM1165 T GU385180 – – – –
Hypsostroma thailandicum MFLUCC 21-0057 T MZ435867 MZ435865 – – –
Jeremyomyces labinae CBS 144617 T MK442529 MK442589 – MK442695 MK442665
Juncaceicola alpina CBS 456.84 T MH873460 MH861761 KY090699 KF253139 KF252188
Keissleriella camporesiana MFLUCC 15-0029 T MN401741 MN401745 MN401743 MN397907 –
Latorua caligans CBS 576.65 T NG_058180 – – – –
Latorua grootfonteinensis CBS 369.72 T NG_058181 – – – –
Lentimurispora urniformis MFLUCC 18-0497 MH179144 – MH179160 MH188055 –
Lentithecium clioninum HHUF 28199 T NG_059391 NR_154137 NG_064845 AB808515 –
Lentithecium pseudoclioninum HHUF 29055 T NG_059392 AB809633 NG_064847 AB808521 –
Leptosphaeria chatkalica YGS22 T MW886099 MW886101 MW886100 MW915583 –
Leptosphaerioides guizhouensis GZAAS 19-4017 T OP099529 OR225067 OR134435 – –
Leptosphaerioides guizhouensis GZAAS 19-4018 OP099530 OR225068 OR134436 – –
Leucaenicola phraeana MFLUCC 18-0472 T MK348003 MK347785 NG_065784 MK360060 MK434867
Leucaenicola aseptata MFLUCC 17-2423 T NG_066309 NR_163332 NG_065776 MK360059 MK434891
Libertasomyces myopori CPC 27354 T NG_058241 KX228281 – – –
Ligninsphaeria jonesii MFLUCC 15-0641 T KU221037 – – – –
Ligninsphaeria jonesii GZCC 15-0080 KU221038 – – – –
Lindgomyces cigarospora G619 T KX655804 KX655794 KX655805 – –
Lindgomyces ingoldianus ATCC 200398 T AB521736 NR_119938 NG_016531 – –
Longicorpus striataspora MFLUCC 18-0267 T MK035988 MK035965 MK035973 MK034428 MK034436
Longiostiolum tectonae MFLUCC 12-0562 T KU764700 KU712447 KU712459 – –
Longipedicellata aptrootii MFLU 10-0297 T KU238894 KU238893 KU238895 KU238892 KU238891
Longipedicellata megafusiformis SJ-KR4 T MZ538546 MZ538512 – MZ567090 –
Lonicericola qujingensis GMBCC1178 T OM855602 OM855593 OM855616 OM857556 –
Lophiostoma carpini CBS 147279 T MW750386 NR_173000 – MW752405 MW752384
Lophiostoma clavatum MFLUCC 18-1316 T MN274566 – MN304835 MN328901 –
Lophiotrema eburnoides MAFF 242970 T LC001707 LC001709 LC001706 LC194403 LC194458
Magnibotryascoma kunmingense HKAS 111919 T MW424785 MW424770 MW424800 MW430106 MW430113
Magnibotryascoma rubriostiolatum CBS 140734 T – KU601590 – KU601609 KU601599
Magnicamarosporium iriomotense HHUF 30125 T AB807509 AB809640 AB797219 AB808485 –
Massaria inquinans CBS 125591 T MH875187 MH863726 HQ599442 HQ599340 –
Massarina pandanicola MFLUCC 17-0596 T MG646947 MG646958 MG646979 MG646986 –
Massarioramusculicola chiangraiensis MFLUCC 17-2240 T MH040228 MH040227 MH040229 – –
Massariosphaeria clematidis MFLU 16-0174 T MT214544 MT310591 MT226663 – –
Matsushimamyces bohaniensis CBS 140592 T KR350633 KP765516 – – –
Misturatosphaeria aurantonotata GKM 1238 T NG_059927 – – GU327761 –
Montagnula acaciae MFLUCC 18-1636 T ON117298 ON117280 ON117267 ON158093 –
Montagnula aquatica MFLU 22-0171 T OP605986 OP605992 OP600504 – –
Montagnula aquatica KUNCC 23-14425 OR583116 OR583097 OR583135 OR588088 OR588107
Morosphaeria muthupetensis NFCC I4219 T MF614796 MF614795 MF614797 MF614798 –
Multilocularia bambusae MFLUCC 11-0180 T KU693438 KU693446 KU693442 – –
Multiseptospora thailandica MFLUCC 11-0183 T KP744490 KP744447 KP753955 KU705657 –
Murilentithecium lonicerae MFLUCC 18-0675 T MK214373 MK214370 MK214376 MK214379 –
Murispora galii MFLUCC 13-0819 T KT709175 KT736081 KT709182 KT709189 –
Muritestudina chiangraiensis MFLUCC 17-2551 T MG602248 MG602247 MG602249 MG602251 MG602250
Neoastrosphaeriella phoenicis MFLUCC 18-1477 T MN712339 MN735995 MN699324 MN744232 MN744233
Neobambusicola magnoliae HKAS 107122 T ON870389 ON878076 ON870914 – –
Neocamarosporium goegapense CPC 23676 T KJ869220 KJ869163 – – –
Neocamarosporium halophilum RSM70 T – OR242722 – OR289930 –
Neocamarosporium halophilum BBB RMS57 OR297952 OR297950 – OR339881 –
Neohelicascus aquaticus KUMCC 19-0107 T MT627662 MT627719 MT864314 MT954384 –
Neolophiostoma pigmentatum MFLUCC 10-0129 T KT324588 KT324587 KT324589 KT324590 –
Neomassaria fabacearum MFLUCC 16-1875 T KX524145 – NG_061245 KX524149 –
Neomassaria formosana NTUCC 17-007 T MH714756 – MH714759 MH714762 MH714765
Neomassaria hongheensis KUMCC 21-0344 T OL423113 OL477614 OL423115 OL754594 OL754595
Neomassarina chromolaenae MFLUCC 17-1480 T MT214466 MT214372 MT214419 MT235785 MT235822
Neomassarina pandanicola MFLUCC 16-0270 T MG298946 MG298945 MG298947 – –
Neomassarina thailandica MFLU 11-0144 T NG_059718 NR_154244 – – –
Neomassarina thailandica MFLUCC 17-1432 MT214467 MT214373 MT214420 – –
Neooccultibambusa thailandensis MFLUCC 16-0274 T MH260308 MH275074 MH260348 MH412780 MH412758
Neophaeosphaeria phragmiticola KUMCC 16-0216 T MG837009 – NG_065735 MG838020 –
Neophaeosphaeria livistonae NCYUCC 19-0393 T OQ437387 OQ437390 OQ437393 – –
Neoplatysporoides aloes CPC 36068 T MN567619 NR_166316 – – –
Neopyrenochaeta annellidica MFLU 11-1105 T MT183502 MT185538 – – –
Neopyrenochaeta cercidis MFLUCC 18-2089 T MK347932 MK347718 MK347823 – MK434908
Neopyrenochaeta chiangraiensis MFLUCC 13-0881 T MT183503 MT185539 MT214975 MT454041 –
Neopyrenochaetopsis hominis UTHSC: DI16-238 T LN907381 LT592923 – – LT593061
Neoroussoella chiangmaiensis MFLU 22-0205 T OQ065735 OQ065738 OQ065736 OQ186448 OQ186450
Neothyrostroma encephalarti CPC 35999 T MN567613 MN562105 – MN556831 –
Neotorula aquatica MFLUCC 15-0342 T KU500576 KU500569 KU500583 – –
Neotorula submersa KUMCC 15-0280 T KX789217 KX789214 – – –
Nigrograna italica MFLU 23-0139 T OR538591 OR538590 – OR531366 OR531365
Occultibambusa bambusae MFLUCC 13-0855 T KU863112 KU940123 – KU940193 KU940170
Occultibambusa jonesii GZCC 16-0117 T KY628322 – KY628324 KY814756 KY814758
Ochraceocephala foeniculi CBS 145654 T MN516774 MN516753 MN516743 MN520149 MN520145
Ohleria modesta CBS 141480 – KX650563 KX650513 KX650534 KX650583
Ohleria modesta WU 36870 – KX650562 – KX650533 KX650582
Omania hydei SQUCC 13750 T MW077155 MW077146 MW077162 MW075772 MW276077
Paraconiothyrium kelleni CBS 149290 T NG_229027 NR_185757 OP348926 OP328919 –
Paradictyoarthrinium aquatica MFLUCC 16-1116 T NG_064501 NR_158861 – – –
Paradictyoarthrinium diffractum MFLUCC 13-0466 KP744498 KP744455 KP753960 – KX437764
Paradictyocheirospora tectonae AMH 10301 T MW854647 MW854646 – MW854832 –
Paralophiostoma hysterioides PUFNI 17617 MT912850 MN582758 MN582762 – MT926117
Paraleptosphaeria polylepidis MA 57843 T – NR_119469 – – –
Paraleptosphaeria polylepidis APA-2999 MK795717 MK795714 MK795720 MK831009 –
Paramonodictys globosa HKAS 129169 T OR091331 OR139016 – OR494045 OR494048
Parapyrenochaeta protearum CBS 131315 JQ044453 JQ044434 – – LT717683
Periconia delonicis MFLUCC 17-2584 T NG_068611 – NG_065770 – MK434901
Periconia pseudodigitata KT 1395 T AB807564 LC014591 AB797274 – –
Phaeomycocentrospora xinjangensis CGMCC 3.20479 T OK256190 OK256193 – – –
Phaeoseptum mali MFLUCC 17-2108 T MK625197 MK659580 – MK647990 MK647991
Phaeoseptum terricola MFLUCC 10-0102 T MH105779 MH105778 MH105780 MH105781 MH105782
Phaeosphaeria oryzae CBS 110110 T KF251689 KF251186 GQ387530 – KF252193
Pleomonodictys capensis CBS 968.97 T KY853521 MH862684 – – –
Pleomonodictys capensis DLUCC 1323 MZ420757 MZ420742 – – MZ442696
Pleomonodictys descalsii FMR 12716 T KY853522 KY853461 – – –
Plenodomus changchunensis CCMJ 5011 T OL897174 OL996123 OL984031 – –
Plenodomus changchunensis CCMJ 5012 OL966928 OL996124 OL984032 – OL944508
Pleopunctum clematidis MFLUCC 17-2091 T MT214573 MT310618 – MT394632 MT394693
Pleopunctum thailandicum MFLUCC 21-0039 T MZ198896 MZ198894 – MZ172461 –
Polyschema sclerotigenum UTHSC DI14-305 T KP769976 KP769975 – – –
Prosthemium alni MFLUCC 17-0240 T KY815013 KY797636 – KY815019 –
Prosthemium intermedium HHUF 30063 T AB553778 AB554108 – – –
Pseudoasteromassaria aquatica MFLUCC 18-1397 T MN913721 MT627674 MT864322 MT954378 –
Pseudoastrosphaeriella longicolla MFLUCC 11-0171 T KT955476 – – KT955438 KT955420
Pseudoastrosphaeriella thailandensis MFLUCC 11-0144 T KT955478 – KT955457 KT955440 KT955416
Pseudoberkleasmium acaciae MFLUCC 17-2590 T NG_066316 NR_163343 NG_065782 MK360073 –
Pseudoberkleasmium chiangmaiense MFLUCC 17-1809 T MK131260 MK131259 – MK131261 –
Pseudoberkleasmium chiangmaiense MFLU 21-0290 OM065940 OM066271 OM065948 OM102996 OM102997
Pseudoberkleasmium chiangmaiense MFLUCC 17-2088 MT214585 MT310630 MT226698 MT394643 MT394699
Pseudoberkleasmium chiangmaiense DLUCC 1655 MZ420759 MZ420744 MZ420749 MZ442693 –
Pseudoberkleasmium chiangraiense MFLUCC 21-0154 T OL584200 OL584189 OL606408 OL912943 OL697401
Pseudoberkleasmium chiangraiense MFLUCC 21-0161 OL584201 OL584190 OL606409 OL912944 OL697402
Pseudoberkleasmium chiangraiense MFLUCC 21-0162 OL584205 OL584191 OL606410 OL912945 OL697403
Pseudoberkleasmium pandanicola KUMCC 17-0178 T MH260304 MH275071 MH260344 – –
Pseudochaetosphaeronema chiangraiense MFLU 21-0083 T MZ457922 MZ457923 – MZ476770 –
Pseudochaetosphaeronema chiangraiense UESTCC 23.0065 OR253260 OR253108 – OR251160 –
Pseudochaetosphaeronema irregulare CGMCC 3.22458 T OQ758163 OQ798972 OQ758194 OQ809057 OQ809023
Pseudochaetosphaeronema irregulare CGMCC 3.22461 OQ758162 OQ798971 OQ758193 OQ809056 OQ809022
Pseudocoleodictyospora sukhothaiensis MFLUCC 12-0554 T KU764710 KU712440 NG_062416 – KU712493
Pseudocoleodictyospora thailandica MFLUCC 12-0565 T KU764701 NR_154337 NG_062417 – KU712494
Pseudocoleophoma heteropanacicola ZHKUCC 23-0880 T OR365486 OR365456 – OR700204 –
Pseudolophiotrema elymicola HHUF 28984 T LC194381 LC194505 LC194339 LC194418 LC194473
Pseudomassarina clematidis MFLU 16-0493 T MT214586 MT415397 MT226699 MT394644 MT394700
Pseudopyrenochaeta lycopersici FMR 15746 T EU754205 NR_103581 NG_062728 – LT717680
Pseudopyrenochaeta terretris FMR 15327 T LT623216 LT623228 – – LT623287
Pseudotetraploa rajmachiensis NFCCI 4618 T MN937204 MN937222 – – –
Pseudoxylomyces elegans KT 2887 AB807598 LC014593 AB797308 AB808576 –
Profundisphaeria fusiformispora GZAAS 20-4010 T – – OR134442 OR140432 OR146942
Profundisphaeria fusiformispora GZAAS 20-4012 OR209667 – OR134443 OR140433 –
Pyrenochaetopsis leptospora CBS 101635 T GQ387627 JF740262 NG_063097 MF795881 LT623282
Pyrenochaetopsis tabarestanensis IBRC:M 30051 T KF803343 NR_155636 NG_065034 – –
Quadricrura bicornis HHUF 30023 T AB524613 AB524797 AB524472 AB524828 –
Quercicola fusiformis MFLUCC 18-0479 T MK348009 MK347790 MK347898 MK360085 MK434864
Quercicola guttulospora MFLUCC 18-0481 T MK348010 MK347791 MK347899 MK360086 –
Quixadomyces cearensis HUEFS 238438 T MG970695 NR_160606 – – –
Roussoella bambusarum GMBC 0316 T ON479892 ON479891 – ON505017 ON505012
Roussoella guttulata MFLUCC 20-0102 T MT734818 MT734821 – MW022188 MW022187
Rubroshiraia bambusae HKAS 102255 T MK804658 MK804678 MK804704 MK819218 –
Rubroshiraia bambusae HKAS 102256 MK804659 MK804679 MK804705 MK819219 –
Salsuginea phoenicis MFLU 19-0015 T MK405280 – – MK404650 –
Salsuginea ramicola KT 2597.2 GU479801 – GU479768 GU479862 GU479834
Salsuginea rhizophorae MFLU 18-0540 T MN017851 – MN017917 – –
Seltsamia ulmi CBS 143002 T MF795794 MF795794 MF795794 MF795882 MF795836
Septoriella chlamydospora MFLUCC 15-0177 T KU163654 KU163658 KU163655 – –
Septoriella hibernica CBS 145371 T MK540036 MK539966 – – MK540097
Setoarthopyrenia chromolaenae MFLUCC 17-1444 T MT214438 MT214344 MT214392 MT235768 MT235805
Shiraia bambusicola GZAAS 2.0708 T KC460982 GQ845414 – – –
Shiraia bambusicola HKAS 102267 MK804657 MK804677 MK804703 MK819217 MK819237
Sporormurispora atraphaxidis MFLUCC 17-0742 T MG829083 MG828971 MG829183 – –
Sporormurispora paulsenii MFLUCC 17-1957 T MK966143 – MK963075 – MN023029
Stagonospora forlicesenensis MFLUCC 15-0054 T KX655547 KX655557 KX655552 KX655562 –
Stagonospora imperaticola MFLUCC 15-0026 KY706133 KY706143 KY706138 KY706146 KY706149
Stemphylium clematidis MFLUCC 14-0937 T MT214583 MT310628 MT226696 – –
Stemphylium carpobroti CPC 38637 T MW175395 MW175355 – – –
Striatiguttula nypae MFLUCC 18-0265 T MK035992 MK035969 MK035977 MK034432 MK034440
Striatiguttula phoenicis MFLUCC 18-0266 T MK035995 MK035972 MK035980 MK034435 MK034442
Subglobosporium tectonae MFLUCC 12-0393 T KU764703 KU712445 KU712464 – KU712485
Subglobosporium tectonae MFLUCC 12-0390 KU764702 KU712446 KU712463 – KU712495
Sublophiostoma thailandica MFLUCC 11-0207 T KX534212 MW136257 KX534218 KX550077 MW088714
Sublophiostoma thailandica MFLUCC 11-0185 KX534216 MW136275 KX534222 KX550080 MW088718
Submersispora variabilis MFLUCC 17-2360 T MN913682 MT627683 MT864310 – –
Submersispora variabilis N-KR15 MZ538561 MZ538527 MZ538575 MZ567103 MZ567114
Sulcatispora acerina HHUF 30449 T LC014610 LC014597 LC014605 LC014615 –
Sulcatispora berchemiae HHUF 29097 T AB807534 AB809635 AB797244 AB808509 –
Sulcosporium thailandica MFLUCC 12-0004 KT426563 MG520958 KT426564 – –
Teichospora trabicola C134 T KU601591 KU601591 – KU601601 KU601600
Tetraploa aquatica MFLU 19-0995 T MT530452 MT530448 – – –
Tetraploa aquatica MFLU 19-0996 MT530453 MT530449 MT530454 – –
Thyridaria acaciae CBS 138873 T NG_058127 KP004469 – – –
Thyridaria broussonetiae CBS 141481 T KX650568 KX650568 KX650515 KX650539 KX650586
Thyrostroma jaczewskii MFLUCC 18-0787 T MK765857 MK765856 MK765858 – –
Torula camporesii KUMCC 19-0112 T MN507402 MN507400 MN507401 MN507403 MN507404
Torula pluriseptata MFLUCC 14-0437 T KY197855 MN061338 KY197862 KY197875 KY197869
Trematosphaeria grisea CBS 332.50 T NG_057979 NR_132039 NG_062930 KF015698 KF015720
Trematosphaeria pertusa CBS 122368 T NG_057809 NR_132040 FJ201991 KF015701 FJ795476
Tubeufia abundata (Tubeufiales) MFLUCC 17-2024 T MH558894 MH558769 – MH550961 MH551095
Tubeufia aquatica (Tubeufiales) MFLUCC 16-1249 T KY320539 KY320522 – KY320556 MH551142
Tubeufia hainanensis (Tubeufiales) GZCC 22-2015 T OR030835 OR030842 – OR046679 OR046685
Tubeufia hainanensis (Tubeufiales) GZCC 23-0589 OR066421 OR066414 – OR058860 OR058853
Tzeanania taiwanensis NTUCC 17-005 T MH461120 MH461123 MH461126 MH461130 MH461128
Tzeanania taiwanensis NTUCC 17-006 MH461121 MH461124 MH461127 MH461131 MH461129
Wicklowia aquatica CBS 125634 T MH875044 OM322822 GU266232 – GU371813
Wicklowia fusiformispora N-KR1 T MZ538567 MZ538533 MZ538576 MZ567108 –
Xenomassariosphaeria clematidis MFLUCC 14-0923 T MT214571 MT310616 – MT394630 –
Xenomassariosphaeria rosae MFLUCC 15-0179 T MG829092 – MG829192 – –
Xenopyrenochaetopsis pratorum CBS 445.81 T GU238136 MH861363 NG_062792 – KT389671
Xiuguozhangia broussonetiae MFLUCC 24-0258 T PQ137419 PQ137417 PQ137421 PQ488461 PQ488459
Xiuguozhangia broussonetiae MFLUCC 24-0259 PQ137420 PQ137418 PQ137422 – PQ488460
Zopfia rosatii CBS 427.62 T NG_066246 NR_160090 – – –

– Data unavailable.

Single gene datasets were aligned using MAFFT version 7 by applying the default setting (https://mafft.cbrc.jp/alignment/server/) (Katoh et al. 2019) and trimmed using trimAl (Capella-Gutiérrez et al. 2009). The trimmed datasets were concatenated using SequenceMatrix (Vaidya et al. 2011). Maximum likelihood (ML) phylogeny was conducted in the IQ-TREE webserver (https://iqtree.cibiv.univie.ac.at) using the default parameters and 1000 ultrafast bootstrap replicates (Nguyen et al. 2015). The nucleotide substitution model for each DNA marker was automatically generated. The Bayesian information criterion (BIC) selection results were as follows: Dataset 1 – invgamma for ITS, LSU, RPB2 and TEF1, and gamma for SSU; Dataset 2 – gamma for ITS and SSU, and invgamma for LSU, RPB2 and TEF1.

Bayesian inference (BI) was carried out in MrBayes on XSEDE (version 3.2.7a) in the online CIPRES Portal (https://www.phylo.org/portal2) (Huelsenbeck et al. 2001; Ronquist and Huelsenbeck 2003; Miller et al. 2010). Markov chain Monte Carlo (MCMC) sampling was applied to obtain posterior probabilities (PP). Four Markov chains were run simultaneously for 50,000,000 and 5,000,000 generations for datasets 1 and 2, respectively, with trees sampled every 100th generation. Burn-in was set to 20% and the remaining 80% were used to compute the PP of the consensus trees. Phylogenetic trees were visualised in FigTree version 1.4.4 (Rambaut and Drummond 2014).

Genetic distances

To corroborate the phylogenetic placement and evolutionary relationships of the new taxon, intra- and inter-generic genetic distances were computed in MEGA-X by applying the Kimura 2-parameter substitution model, gamma distribution, and pairwise deletion options (Tamura et al. 2013).

Results

Phylogenetic analyses

Blast searches of LSU, ITS, SSU, TEF1 and RPB2 sequences indicated that the two isolates are highly similar to various genera in Pleosporales, including Atrocalyx, Hermatomyces, Lophiotrema and Pseudoberkleasmium.

Dataset 1 consisted of 4250 characters (LSU = 1–846, ITS = 847–1355, SSU = 1356–2359, TEF1 = 2360–3257, and RPB2 = 3258–4250), which was analysed to depict relationships at a higher taxonomic level for Xiuguozhangia (Fig. 1). Outgroup taxa were selected from Tubeufiales. The log-likelihood of the consensus tree (Fig. 1) was -143487.839. The average standard deviation of split frequencies at the end of the total MCMC generations converged to 0.0092.

Figure 1. 

Maximum likelihood analysis (IQ-tree) based on a combined dataset of LSU, ITS, SSU, TEF1, and RPB2 sequences of all families (with representative genera) of Pleosporales. Bootstrap support values (ML ≥ 80%) and Bayesian posterior probabilities (PP ≥ 0.95) are given above the branches or near the nodes as ML/PP. Hyphens (--) indicate bootstrap support values below 80% for ML and posterior probabilities below 0.95. The tree is rooted with Tubeufia abundata (MFLUCC 17-2024), T. aquatica (MFLUCC 16-1249), T. hainanensis (GZCC 22-2015 and GZCC 23-0589), Berkleasmium aquaticum (MFLUCC 17-0049 and MFLUCC 17-0039) and B. longisporum (MFLUCC 17-1999 and MFLUCC 17-2002) (Tubeufiales). Type, ex-type, and reference strains are denoted with T. Our isolates are in bold font. The different colour blocks indicate the families to which the taxa belong.

Dataset 2 comprised 4285 characters (LSU = 1–833, ITS = 834–1344, SSU = 1345–2350, TEF1 = 2351–3272, and RPB2 = 3273–4285), and was used to infer the inter-generic relationships of Xiuguozhangia, which comprised multiple strains for each genus (Fig. 2). Based on the results from dataset 1, four taxa belonging to Anteagloniaceae (Pleosporales) were selected as outgroups as they are phylogenetically closely related to Pseudoberkleasmiaceae and Hermatomycetaceae. The log-likelihood of the consensus tree (Fig. 2) was -17829.366. The average standard deviation of split frequencies at the end of the total MCMC generations converged to 0.0031.

Figure 2. 

Maximum likelihood analysis (IQ-tree) based on the combined LSU, ITS, SSU, TEF1 and RPB2 sequences of Xiuguozhangia, Pseudoberkleasmium and Hermatomyces, generated from dataset 2. Bootstrap support values (ML ≥ 80%) and Bayesian posterior probabilities (PP ≥ 0.95) are given above the branches or near the nodes as ML/PP. Hyphens (--) indicate bootstrap support values below 80% for ML and posterior probabilities below 0.95. The tree is rooted with Anteaglonium gordoniae (MFLUCC 17-2431 and CD7) and A. latirostrum (GKM1119 and GKML100Nb) (Anteagloniaceae, Pleosporales). Type, ex-type, and reference strains are denoted with T. The new isolates are in bold font. The different colour blocks indicate the families to which the taxa belong.

The two strains of Xiuguozhangia broussonetiae (MFLUCC 24-0258 and MFLUCC 24-0259) grouped with 100% ML and 1.00 PP support. This subclade formed a separate sister lineage to Pseudoberkleasmium with 100% ML and 1.00 PP support (Figs 1, 2). Single and combined gene trees from datasets 1 and 2 indicate that our species is phylogenetically most closely related to Pseudoberkleasmium, and its placement in Pseudoberkleasmiaceae is supported by maximum bootstrap support and posterior probability in both combined data trees.

Genetic distances

Based on the generic relationship depicted in Figs 1, 2, we computed the group mean distances between Hermatomyces spp. (group 1), Pseudoberkleasmium spp. (group 2) and Xiuguozhangia spp. (group 3) across ITS and LSU markers. The difference in the genetic distances across both markers is given in Table 2.

Table 2.
Download as
CSV
XLSX

Group mean genetic distances (%) in Hermatomyces spp., Pseudoberkleasmium spp., and Xiuguozhangia spp. across ITS (511 bp) and LSU (833 bp).

DNA markers Groups Group 1: Hermatomyces spp. Group 2: Pseudoberkleasmium spp.
ITS Group 1: Hermatomyces spp. 0 –
Group 2: Pseudoberkleasmium spp. 9.052 0
Group 3: Xiuguozhangia spp. 10.62834205 13.25008949
LSU Group 1: Hermatomyces spp. 0 –
Group 2: Pseudoberkleasmium spp. 3.1615 0
Group 3: Xiuguozhangia spp. 4.1404 3.8669

Taxonomy

Xiuguozhangia broussonetiae Gomdola, Jayaward. & K.D. Hyde, sp. nov.

Index Fungorum: IF901943
Facesoffungi Number: FoF16323
Fig. 3

Holotype

MFLU 24-0227.

Etymology

The specific epithet refers to the host genus, Broussonetia, from which the species was isolated.

Description

Saprobic on decaying stems of Broussonetia papyrifera. Sexual morph not observed. Asexual morph on substrate. Hyphomycetous. Colonies on the substrate effuse, hairy, olivaceous to dark brown, appearing velvety due to numerous conidiophores. Mycelium semi-immersed or immersed, composed of septate, branched, smooth, hyaline or pale brown to brown hyphae. Conidiophores (430–)550–750(–890) µm long (x– = 681 µm, n = 10), 15–24(–34) µm wide (x– = 21.8 µm, n = 10) at the base, 11–14(–16) µm wide (x– = 12.3 µm, n = 10) in the middle, 7–10 µm(–12) wide (x– = 7.8 µm, n = 10) at the apex, rarely branched, macronematous, mononematous, differentiated, smooth, thick-walled, erect, straight or flexuous, brown to dark brown, wider at the base and ocasionally conical at the apex, comprising 12–17 septa. Conidiogenous cells 5–12 µm long × 4–9 µm wide (x– = 7.4 × 6.2 µm, n = 10), holoblastic, enteroblastic, integrated, smooth-walled, ovoid to ampulliform, hyaline or brown to dark brown, occurring terminally or intercalary on conidiophores, with up to four successive percurrent proliferations, with new conidiogenous cells developing on subtending cells. Conidia 25–40 µm long × 30–60 µm wide (x– = 35 × 47 µm, n = 30), width measured between the two extremities of the apices, solitary, dictyospored and cheirospored, fan-shaped to cheiroid, lenticular in edge view, occasionally apically 2–3-lobed, dark brown to olivaceous brown, dictyoseptate, with up to 15 columns of cells radiating from a protuberant basal cell 2–3 µm wide, septa obscured by dark pigmentation, and with 1–3 apical appendages (rarely without appendages). Apical appendages (2–)6–16(–20) µm long (x– = 10.9 µm, n = 10), 4–5 µm wide (x– = 4.4 µm, n = 10) at the base, 4–5 µm wide (x– = 4.1 µm, n = 10) at the apex, mostly short and untapered, sometimes long and tapering, arising from the sides of the outermost rows of cells of the conidium, pale brown to brown, becoming hyaline and rounded at the tips, and consisting of 1–5 septa.

Culture characteristics

On PDA, colony circular with lobate to crenated margin, reaching 25 mm diam. after 28 days incubated at 25 °C, greyish white to olivaceous brown from center to edge, fluffy, raised to convex, penetrating the media and displaying a dark brown colour around the media.

Material examined

Thailand • Chiang Rai Province, Mae Fah Luang University Botanical Garden, on decaying stems of Broussonetia papyrifera (Moraceae), 19 May 2023, D. Gomdola, F2-A (MFLU 24-0227, holotype), ex-type MFLUCC 24-0258.

Additional material examined

Thailand • Chiang Rai Province, Mae Fah Luang University Botanical Garden, on decaying stems of Broussonetia papyrifera (Moraceae), 19 May 2023, D. Gomdola, F2-B (MFLU 24-0228), living culture MFLUCC 24-0259.

GenBank accession numbers

MFLUCC 24-0258; ITS = PQ137417, LSU = PQ137419, SSU = PQ137421, RPB2 = PQ488459 and TEF1 = PQ488461; MFLUCC 24-0259; ITS = PQ137418, LSU = PQ137420, SSU = PQ137422 and RPB2 = PQ488460.

Notes

Our two isolates (MFLUCC 24-0258 and MFLUCC 24-0259) group together with 100% ML and 1.00 PP support, and this subclade is sister to Pseudoberkleasmium species with 100% ML and 1.00 PP support (Figs 1, 2).

Figure 3. 

Xiuguozhangia broussonetiae (MFLUCC 24-0258) A substrate (Broussonetia papyrifera) B, C colonies on the substrate D–F conidiophores and attachment of conidia G–I conidiogenous cells with percurrent proliferations J–O, Q, R conidia bearing appendages P top view of a conidium S top view of colonies on PDA after 7 days T top and reverse colonies on PDA after 28 days. Scale bars: 100 µm (B, C); 50 µm (D–F); 10 µm (G–I, P); 20 µm (J–O, Q, R).

A close comparison of the morphological characters across the existing Xiuguozhangia species is collated and presented in Table 3 to support the establishment of the new species, X. broussonetiae.

Table 3.
Download as
CSV
XLSX

Phenotypic comparison of Xiuguozhangia species. Features that depict X. broussonetiae from other Xiuguozhangia species are in bold.

Species and characters X. appendiculata X. broussonetiae X. indica X. macrospora X. punicae X. rhaphidophorae X. rosae
Colonies on the natural substrates Features Effuse, hairy Effuse, appearing velvety due to numerous conidiophores Effuse, hairy Effuse, hairy, growing in association with species of lichen Effuse, hairy Effuse, hairy Effuse, hairy
Colour Olivaceous to dark brown Olivaceous to dark brown Olivaceous to dark brown Olivaceous to dark brown Olivaceous to dark brown Olivaceous to dark brown Olivaceous to dark brown
Conidiophores Size (µm) Up to 620 µm long, 18–35 µm wide at the base, 13–18 µm wide in the middle, 7–11.5 µm wide at the apex (430–)550–750(–890) µm long, 15–24(–34) µm wide at the base, 11–14(–16) µm wide in the middle, 7–10 µm(–12) wide at the apex Up to 530 µm long, 13–15 µm wide at the base, 10–12 µm wide in the middle, 3.6–5 µm wide at the apex 160–340(–570) µm long, 12–21 µm wide at the base, 11–14 µm wide in the middle, 6–7.5 µm wide at the apex Up to 550 µm long, 20–30 µm wide at the base, 10–17 µm wide in the middle, 5–8 µm wide at the apex Up to 630 µm long, 18–25 µm wide at base, 10–15 µm wide in the middle, 4–7 µm wide at apex Up to 730 µm long, 12–20 µm wide at the base, 7–14 µm wide in the middle, 5–9 µm wide at the apex
Features Mononematous, erect, straight or flexuous, unbranched, smooth, thick-walled, comprising 4–8 septa Macronematous, mononematous, erect, straight or flexuous, rarely branched, smooth, thick-walled, wider at the base and ocasionally conical at the apex, comprising 12–17 septa Mononematous, basal portion sheath-like, erect, straight, unbranched, smooth, thick-walled, comprising 4–13 septa Mononematous, erect, straight, unbranched, smooth, thick-walled, comprising 3–8 septa Macronematous, mononematous, erect, straight or flexuous, unbranched, smooth, thick-walled, comprising 5–11 septa Macronematous, mononematous, erect, straight or flexuous, unbranched, smooth, thick-walled, comprising 10–15 septa Macronematous, mononematous, erect, straight or flexuous, unbranched, smooth, thick-walled, comprising 8–14 septa
Colour Dark brown Brown to dark brown Brown, becoming paler towards the apex Brown Not mentioned Not mentioned Not mentioned
Conidiogenous cells Size (µm) Not mentioned 5–12 µm long, 4–9 µm wide 7.5–13.5 × 4.2–7.4 µm 9–19 × 7.5–10.0 µm Not mentioned Not mentioned Not mentioned
Features Holoblastic, integrated, terminal, sometimes lateral, truncate after conidium secession Holoblastic, enteroblastic, integrated, terminal or intercalary, smooth-walled, ovoid to ampulliform, percurrently proliferating; sometimes new condiogenous cells developing on subtending cells Flask-shaped, integrated, terminal and lateral, truncate after conidial secession, proliferating percurrently up to 4 times Flask-shaped, integrated, terminal, truncate after conidial secession, proliferating percurrently up to 5 times Monotretic, integrated, terminal, sometimes lateral, with up to five successive percurrent proliferations Monotretic, integrated, terminal and lateral Monotretic, integrated, terminal and lateral, with up to three successive percurrent proliferations
Colour Not mentioned Hyaline or brown to dark-brown Brown Brown Not mentioned Not mentioned Not mentioned
Conidia Size (µm) 50–80 µm long, 60–90 µm wide 25–40 µm long, 30–60 µm wide 34–44 µm long, 39–52 µm wide (73–)81–120(–135) µm long, (51–)67–94(–106) µm wide 50–65 µm long, 58–95 µm wide 27–41 µm long, 30–43 µm wide 45–50 µm long, 53–76 µm wide
Features Fan-shaped, lenticular in edge view, 2–3-lobed, dictyoseptate, with rows of cells radiating from a protuberant basal cell 6–7.5 µm wide Dictyospored and cheirospored, fan-shaped to cheiroid, lenticular in edge view, occasionally apically 2–3-lobed, dictyoseptate, with up to 15 columns of cells radiating from a protuberant basal cell 2–3 µm wide Fan-shaped, sometimes 2–5-lobed, lenticular in edge view, dictyoseptate, with up to 15 rows of cells, basal cell protuberant, 3.7–5 µm wide Campanulate or fan-shaped, sometimes apically 2–3-lobed, lenticular in edge view, dictyoseptate, smooth, with up to 18 vertical rows of cells, basal cell protuberant, 7–9 µm wide Fan-shaped, sometimes 2–3-lobed, lenticular in edge view, dictyoseptate, with up to 18 rows of cells radiating from a protuberant basal cell 4.5–6.5 µm wide Fan-shaped, sometimes lobed, lenticular in edge view, dictyoseptate, with eight rows of cells radiating from a protuberant basal cell 2.5–5 µm wide Fan shaped, sometimes 2–3-lobed, lenticular in edge view, dictyoseptate, with 20 rows of cells radiating from a protuberant basal cell 3–5 µm wide
Colour Dark brown Dark brown to olivaceous brown Brown Brown Dark brown Dark brown Dark brown
Apical appendage(s) Size (µm) 75–120 µm long, 4–6 µm wide, tapering to 1.5–2 µm wide (2–)6–16(–20) µm long, 4–5 µm wide at the base and apex 15–30 µm long, 3–4.4 µm wide at the base, tapering to 2.4–2.8 µm wide (33–)52–74(–126) µm long, 6.5–8.0 µm wide at the base 24–99 µm long, 3–5.5 µm wide, tapering to 1–1.5 µm wide 15–35 µm long, 3–4 µm wide, tapering to 2–3 µm wide
Features 2–4 appendages, tapering, 3–5-septate Without or with 1–3 appendages, rounded at the tips, mostly untapered, sometimes tapering 1–5-septate 2 appendages (Up to 4), tapering, 1–2-septate 2–3 appendages (rarely up to 4), tapering, 1–3(–6)-septate 2 appendages (rarely up to 3), smooth, tapering, 0–3-septate Appendages absent 2 appendages, tapering, 1–2-septate
Color Brown, and apically hyaline Pale brown to brown, becoming hyaline at the tips Brown, and apically hyaline Brown, and apically hyaline Brown, and apically hyaline - Brown, and apically hyaline
Host(s) On dead twigs of an unknown host On decaying stems of Broussonetia papyrifera On dried bamboo culms On dead bamboo stem On dead branches of Punica granatum On dead branches of Rhaphidophora decursiva On dead branches of Rosa chinensis
Distribution(s) India Thailand India India China China China
References (Bhat and Kendrick 1993) This study Sureshkumar et al. (2005) Dubey and Jash (2024) Zhang et al. (2009), Zhang et al. (2014) Zhang et al. (2009, 2014) Zhang et al. (2009), Zhang et al. (2014)

A summary of the main findings from the morphological assessment is presented below:

  1. The conidiophores of our species, Xiuguozhangia broussonetiae, are longer than all other Xiuguozhangia taxa (up to 890 μm long), comprising numerous septa (up to 17). Furthermore, conidiophores of other Xiuguozhangia species are unbranched, while those of X. broussonetiae are sometimes branched (Fig. 3E).
  2. The conidia of Xiuguozhangia broussonetiae (Figs 3J–O, Q, R) consist of up to 15 columns of cells radiating from a protuberant basal cell that is smaller (2–3 μm wide) compared to those of other species.
  3. The appendages of X. broussonetiae are mostly untapered (Figs 3N, 3P–R) and sometimes taper towards the tips (Fig. 3B), whereas all other Xiuguozhangia species have only tapering appendages.
  4. Xiuguozhangia broussonetiae differs from all extant species but has a close morphological overlap with X. rosae. However, the primary feature that demarcates X. broussonetiae from X. rosae is the number and features of the apical appendages. Up to three appendages were observed in X. broussonetiae (sometimes seen without appendage), while X. rosae has two appendages. In addition, the appendages of X. broussonetiae are 1–5-septate, while those of X. rosae display one to two septa.
  5. Xiuguozhangia broussonetiae has percurrently proliferating, enteroblastic conidiogenous cells that sometimes produce new conidiogenous cells on subtending cells (Fig. 3D–I). This feature has not been observed in other species of the genus.

Based on these morphological differences, we conclude that our taxon is distinct from the existing Xiuguozhangia species.

Discussion

The number of hyphomycetes introduced over the past decade has increased substantially, indicating that their diversity is rather high (Hyde et al. 2020, 2023, 2024a, b; Bhunjun et al. 2022; Manawasinghe et al. 2022; Calabon et al. 2023; Liu et al. 2024b; Tian et al. 2024; Zhang et al. 2024). Hyphomycetes have a ubiquitous distribution in aquatic and terrestrial habitats, occurring on different substrates in tropical and subtropical regions (Seifert et al. 2011; Bao et al. 2021; Wu and Diao 2022, 2023; Dong et al. 2023; Senanayake et al. 2023; Yang et al. 2023; Liu et al. 2024a, b). Several hyphomycetes are rather speciose, for example, Cladosporium, Dictyosporium, Helminthosporium, and Sporidesmium (Shenoy et al. 2006; Hu et al. 2023; Yang et al. 2023). However, the hyphomycete genus, Xiuguozhangia, does not appear to be species-rich, and no studies have reported its occurrence in Thailand. This study introduces a new species, Xiuguozhangia broussonetiae, from Broussonetia papyrifera in northern Thailand.

There are more than 30,000 fungal species that exhibit the asexual morph, belonging to 2265 hyphomycetous genera (Wijayawardene et al. 2022b; Jayawardena et al. 2023; Perera et al. 2023; Ma et al. 2024; Zhang et al. 2024). The Xiuguozhangia species reported so far occur only in their asexual morph as saprobic hyphomycetes. A probable explanation for their asexual morph occurrence might be for survival, as conidia can be produced in large amounts and are easily released for dispersal compared to ascospores (Gilbert and Parker 2023). Their sexual morphs have not been reported yet, and no research has been conducted to link their asexual and sexual morphs, possibly due to the lack of molecular data and their availability in cultures. To date, all Xiuguozhangia taxa have been documented from Asia (China, India and Thailand – this study). We anticipate discovering more species in this genus, as well as the sexual morph, especially in high biodiversity areas like China and Thailand, as these taxa have not yet been extensively collected, isolated, and studied.

In the phylogenetic analysis of dataset 1, which included representative taxa from all families of Pleosporales, Xiuguozhangia forms a sister lineage to Pseudoberkleasmium, with maximum statistical support in both methods of analysis (Fig. 1). Pleosporales taxa were accommodated in over 90 families (with and without molecular data), and have a worldwide distribution in terrestrial and aquatic habitats (Hongsanan et al. 2020; Bhunjun et al. 2021; Jiang et al. 2021; Wijayawardene et al. 2020, 2022a; Yang et al. 2022; Tang et al. 2023; Hyde et al. 2024a, b; Pem et al. 2024; Wei et al. 2024). In this study, we maintain 95 families in this order: 92 with DNA sequence data and two without (Diademaceae and Lizoniaceae). Additionally, we exclude Mycoporaceae (Mycoporum spp.) from the analysis due to its divergent sequences and long branches in the preliminary phylogenetic trees. Further research is needed to determine its placement within Pleosporales. Arthopyreniaceae was classified under Pleosporales in Wijayawardene et al. (2022a) and Pem et al. (2024), but we do not retain it in this order as its taxa cluster basal to Pleosporales (results not shown), similar to the findings by Thiyagaraja et al. (2021). We also retain Sublophiostomataceae within Pleosporales (Hongsanan et al. 2021), but it was not included in Pem et al. (2024). In our phylogenetic analysis, most families in Pleosporales are monophyletic, thereby further supporting their rank. A few families [e.g., Halotthiaceae (87% ML); Pseudomassarinaceae; and Testudinaceae (81% ML)] lack strong statistical support but have consistently been recovered as distinct groups in previous studies using different datasets and/or methods of analysis (Crous et al. 2018; Yang et al. 2022), suggesting that these are separate lineages distinct from other families. However, available genetic markers and DNA sequence data may not be sufficient to resolve their familial relationships with high confidence.

Xiuguozhangia is sister to Pseudoberkleasmium and closely related to Hermatomyces (Figs 1, 2). Given that we assigned Xiuguozhangia in Pseudoberkleasmiaceae based on the analysis of dataset 1, we analysed dataset 2 to further validate its taxonomic placement, which was based on multiple strains representing all species of Hermatomyces and Pseudoberkleasmium. Since ITS and LSU are important genetic markers to delimit fungal genera, we computed the group mean distances across these gene regions between Hermatomyces spp., Pseudoberkleasmium spp. and Xiuguozhangia spp. Considering the inter-generic nucleotide differences between each group (Table 2), the establishment of Xiuguozhangia as a distinct genus from Pseudoberkleasmium is supported.

The characterisation of Xiuguozhangia as a separate genus from Pseudoberkleasmium is further corroborated based on their distinct features. Xiuguozhangia differs from Pseudoberkleasmium in their colonies on natural substrates, and conidiophore, conidiogenous cell and conidial features (Zhang et al. 2014; Tibpromma et al. 2018). Xiuguozhangia species are depicted by dictyoseptate, campanulate, cheiroid, and brown conidia comprising multiple layers of cells, either with or without apical appendages (Zhang et al. 2014). However, Pseudoberkleasmium species are characterised by broadly ellipsoidal to obovoid, flattened, one-cell thick, muriform, and brown olivaceous green conidia without any apical appendage (Tibpromma et al. 2018). Conidial images of Pseudoberkleasmium and Hermatomyces are illustrated in Fig. 2. Since other species of Xiuguozhangia do not have molecular data, we were unable to compare the inter-species nucleotide differences, and we had to rely solely on morphology to establish our new species, X. broussonetiae.

With respect to the inter-species morphological differences in the genus, the new species, X. broussonetiae, differs from all extant species (Table 3), but exhibits overlapping features with X. rosae. Nonetheless, X. broussonetiae differs from X. rosae in terms of the number and characters of the apical appendages, as outlined in the note section above. Other observed differences are given in Table 3. Xiuguozhangia was classified as Ascomycota genus incertae sedis in the latest outline by Wijayawardene et al. (2022a), as it was difficult to propose its precise familial placement without the availability of DNA sequence data. Since our study is the first to provide molecular data, we refer Xiuguozhangia in Pseudoberkleasmiaceae. Hyde et al. (2019) initially introduced this family as monotypic to accommodate Pseudoberkleasmium species based on morphology and molecular data. Pseudoberkleasmium was described by Tibpromma et al. (2018) based on morphology and multigene phylogenetic analyses. Pseudoberkleasmium taxa resemble those of Berkleasmium, but Berkleasmium belongs to Tubeufiales (Tibpromma et al. 2018; Wijayawardene et al. 2022a; this study). Besides Pseudoberkleasmium, we treat Xiuguozhangia as a member of Pseudoberkleasmiaceae based on our inferred phylogenies. Upon the addition of X. broussonetiae to the genus, there are seven species of Xiuguozhangia, sharing the same holoblastic conidial ontogeny as a generic feature.

Due to the limited taxon sampling and analysis, and the lack of comprehensive DNA sequence data, there remains uncertainty in using conidial ontogeny as a basis for distinguishing Xiuguozhangia from Piricaudiopsis. As such, we refrain from drawing further conclusions regarding their taxonomic separation at this stage. Instead, a more effective approach would involve extensive sampling across diverse localities and habitats, particularly those that support the growth of hyphomycetes. Coupled with a systematic acquisition of DNA sequence data, these efforts would provide a more robust support for classifying Xiuguozhangia species and other poorly studied Pleosporalean taxa (Liu et al. 2024b; Pem et al. 2024). Moreover, a thorough re-examination of the type species of each genus is essential to ascertain whether these two genera are indeed distinct or if they might represent a single taxonomic entity.

Acknowledgments

We thank Dr Shaun Pennycook for suggesting and validating the nomenclature of the new species. Deecksha Gomdola thanks the Mae Fah Luang University for providing the MFU student scholarship 2020 (reference: GR-ST-FS-63-06) to pursue to PhD program, and for allocating the thesis writing grant [reference: 7702(6)/201 (No.0275)]. Deecksha Gomdola thanks the Mushroom Research Foundation (MRF) for partly funding her project, and the staffs of the MFU botanical garden for their help in the sample collection and identification of the host. Deecksha Gomdola also thanks Prof. Dr. Yong Wang of the Guizhou University (Department of Plant Pathology) for providing funds to conduct molecular work under his NFSC grant No. 31972222. We would also like to acknowledge the grant GDPX[2019]13 for partly funding this project. We thank Miss Tang Xia and Mr. Xingguo Tian for providing conidial images of Hermatomyces and Pseudoberkleasmium to complement Fig. 2. The authors extend their appreciation to the Researchers Supporting Project number (RSP2024R114), King Saud University, Riyadh, Saudi Arabia.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by Guizhou University (NFSC grant No. 31972222 and GDPX[2019]13), Mae Fah Luang University, Mushroom Research Foundation and MFU student scholarship 2020 (reference: GR-ST-FS-63-06); thesis writing grant [reference: 7702(6)/201 (No.0275)]; Researchers Supporting Project number (RSP2024R114).

Author contributions

Conceptualization: DG, EG, RJ. Data curation: DG. Formal analysis: DG. Funding acquisition: YW. Investigation: DG. Methodology: DG. Resources: YW, FA, KDH. Software: DG. Supervision: KDH, YW, RSJ. Validation: RSJ, YW, RJ, FA, KDH, EG. Visualization: DG. Writing - original draft: DG. Writing - review and editing: RSJ, YW, RJ, KDH, EG.

Author ORCIDs

Deecksha Gomdola https://orcid.org/0000-0002-0817-1555

Rajesh Jeewon https://orcid.org/0000-0002-8563-957X

Eleni Gentekaki https://orcid.org/0000-0002-3306-6714

Ruvishika S. Jayawardena https://orcid.org/0000-0001-7702-4885

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

Fatimah Alotibi https://orcid.org/0000-0003-3629-5755

Data availability

All data generated or analysed during this study are included in this published article.

References

  • Bao DF, Hyde KD, McKenzie EHC, Jeewon R, Su HY, Nalumpang S, Luo ZL (2021) Biodiversity of lignicolous freshwater hyphomycetes from China and Thailand and description of sixteen species. Journal of Fungi 7(8): 669. https://doi.org/10.3390/jof7080669
  • Bhat DJ, Kendrick B (1993) Twenty-five new conidial fungi from the Western Ghats and the Andaman Islands (India). Mycotaxon 49: 19–90.
  • Bhunjun CS, Phukhamsakda C, Jeewon R, Promputtha I, Hyde KD (2021) Integrating different lines of evidence to establish a novel ascomycete genus and family (Anastomitrabeculia, anastomitrabeculiaceae) in pleosporales. Journal of Fungi 7(2): 1–18. https://doi.org/10.3390/jof7020094
  • Bhunjun CS, Niskanen T, Suwannarach N, Wannathes N, Chen YJ, McKenzie EHC, Maharachchikumbura SSN, Buyck B, Zhao CL, Fan YG, Zhang JY, Dissanayake AJ, Marasinghe DS, Jayawardena RS, Kumla J, Padamsee M, Chen YY, Liimatainen K, Ammirati JF, Phukhamsakda C, Liu JK, Phonrob W, Randrianjohany É, Hongsanan S, Cheewangkoon R, Bundhun D, Khuna S, Yu WJ, Deng LS, Lu YZ, Hyde KD, Lumyong S (2022) The numbers of fungi: Are the most speciose genera truly diverse? Fungal Diversity 114(1): 387–462. https://doi.org/10.1007/s13225-022-00501-4
  • Calabon MS, Hyde KD, Jones EBG, Bao DF, Bhunjun CS, Phukhamsakda C, Shen HW, Gentekaki E, Al Sharie AH, Barros J, Chandrasiri KSU, Hu DM, Hurdeal VG, Rossi W, Valle LG, Zhang H, Figueroa M, Raja HA, Seena S, Song HY, Dong W, El-Elimat T, Leonardi M, Li Y, Li YJ, Luo ZL, Ritter CD, Strongman DB, Wei MJ, Balasuriya A (2023) Freshwater fungal biology. Mycosphere 14(1): 195–413. https://doi.org/10.5943/mycosphere/14/1/4
  • Capella-Gutiérrez S, Silla-Martínez JM, Gabaldón T (2009) trimAl: A tool for automated alignment trimming in large-scale phylogenetic analyses. Bioinformatics 25(15): 1972–1973. https://doi.org/10.1093/bioinformatics/btp348
  • Chaiwan N, Gomdola D, Wang S, Monkai J, Tibpromma S, Doilom M, Wanasinghe DN, Mortimer PE, Lumyong S, Hyde KD (2021) an online database providing updated information of microfungi in the Greater Mekong Subregion. Mycosphere 12(1): 1513–1526. https://doi.org/10.5943/mycosphere/12/1/19
  • Crous PW, Schumacher RK, Wingfield MJ, Akulov A, Denman S, Roux J, Braun U, Burgess TI, Carnegie AJ, Váczy KZ, Guatimosim E, Schwartsburd PB, Barreto RW, Hernández-Restrepo M, Lombard L, Groenewald JZ (2018) New and interesting Fungi. 1. Fungal Systematics and Evolution 1(1): 169–215. https://doi.org/10.3114/fuse.2018.01.08
  • Dong W, Hyde KD, Jeewon R, Liao CF, Zhao HJ, Kularathnage ND, Li H, Yang YH, Pem D, Shu YX, Gafforov Y, Manawasinghe IS, Doilom M (2023) Mycosphere notes 449–468: Saprobic and endophytic fungi in China, Thailand, and Uzbekistan. Mycosphere 14(1): 2208–2262. https://doi.org/10.5943/mycosphere/14/1/26
  • Hongsanan S, Hyde KD, Phookamsak R, Wanasinghe DN, McKenzie EHC, Sarma VV, Lücking R, Boonmee S, Bhat JD, Liu NG, Tennakoon DS, Pem D, Karunarathna A, Jiang SH, Jones GEB, Phillips AJL, Manawasinghe IS, Tibpromma S, Jayasiri SC, Sandamali D, Jayawardena RS, Wijayawardene NN, Ekanayaka AH, Jeewon R, Lu YZ, Phukhamsakda C, Dissanayake AJ, Zeng XY, Luo ZL, Tian Q, Thambugala KM, Dai D, Samarakoon MC, Chethana KWT, Ertz D, Doilom M, Liu JK, Pérez-Ortega S, Suija A, Senwanna C, Wijesinghe SN, Niranjan M, Zhang S-N, Ariyawansa HA, Jiang H-B, Zhang J-F, Norphanphoun C, de Silva NI, Thiyagaraja V, Zhang H, Bezerra JDP, Miranda-González R, Aptroot A, Kashiwadani H, Harishchandra D, Sérusiaux E, Abeywickrama PD, Bao D-F, Devadatha B, Wu H-X, Moon KH, Gueidan C, Schumm F, Bundhun D, Mapook A, Monkai J, Bhunjun CS, Chomnunti P, Suetrong S, Chaiwan N, Dayarathne MC, Yang J, Rathnayaka AR, Xu J-C, Zheng J, Liu G, Feng Y, Xie N (2020) Refined families of Dothideomycetes: Orders and families incertae sedis in Dothideomycetes. Fungal Diversity 105(1): 17–318. https://doi.org/10.1007/s13225-020-00462-6
  • Hongsanan S, Phookamsak R, Goonasekara ID, Thambugala KM, Hyde KD, Bhat JD, Suwannarach N, Cheewangkoon R (2021) Introducing a new pleosporalean family Sublophiostomataceae fam. nov. to accommodate Sublophiostoma gen. nov. Scientific Reports 11(1): 9496. https://doi.org/10.1038/s41598-021-88772-w
  • Hu Y, Liu J, Xu Z, Castañeda-Ruíz RF, Zhang K, Ma J (2023) Morphology and multigene phylogeny revealed three new species of Helminthosporium (Massarinaceae, Pleosporales) from China. Journal of Fungi 9(2): 280. https://doi.org/10.3390/jof9020280
  • Huelsenbeck JP, Ronquist F, Nielsen R, Bollback JP (2001) Bayesian inference of phylogeny and its impact on evolutionary biology. Science 294(5550): 2310–2314. https://doi.org/10.1126/science.1065889
  • Hyde KD, Tennakoon DS, Jeewon R, Bhat DJ, Maharachchikumbura SSN, Rossi W, Leonardi M, Lee HB, Mun HY, Houbraken J, Nguyen TTT, Jeon SJ, Frisvad JC, Wanasinghe DN, Lücking R, Aptroot A, Cáceres MES, Karunarathna SC, Hongsanan S, Phookamsak R, de Silva NI, Thambugala KM, Jayawardena RS, Senanayake IC, Boonmee S, Chen J, Luo ZL, Phukhamsakda C, Pereira OL, Abreu VP, Rosado AWC, Bart B, Randrianjohany E, Hofstetter V, Gibertoni TB, Soares AM da S, Plautz Jr HL, Sotão HMP, Xavier WKS, Bezerra JDP, de Oliveira TGL, de Souza-Motta CM, Magalhães OMC, Bundhun D, Harishchandra D, Manawasinghe IS, Dong W, Zhang SN, Bao DF, Samarakoon MC, Pem D, Karunarathna A, Lin CG, Yang J, Perera RH, Kumar V, Huang SK, Dayarathne MC, Ekanayaka AH, Jayasiri SC, Xiao Y, Konta S, Niskanen T, Liimatainen K, Dai YC, Ji XH, Tian XM, Mešić A, Singh SK, Phutthacharoen K, Cai L, Sorvongxay T, Thiyagaraja V, Norphanphoun C, Chaiwan N, Lu YZ, Jiang HB, Zhang JF, Abeywickrama PD, Aluthmuhandiram JVS, Brahmanage RS, Zeng M, Chethana T, Wei D, Réblová M, Fournier J, Nekvindová J, do Nascimento Barbosa R, dos Santos JEF, de Oliveira NT, Li G-J, Ertz D, Shang Q-J, Phillips AJL, Kuo C-H, Camporesi E, Bulgakov TS, Lumyong S, Jones EBG, Chomnunti P, Gentekaki E, Bungartz F, Zeng X-Y, Fryar S, Tkalčec Z, Liang J, Li G, Wen T-C, Singh PN, Gafforov Y, Promputtha I, Yasanthika E, Goonasekara ID, Zhao R-L, Zhao Q, Kirk PM, Liu J-K, Yan JY, Mortimer PE, Xu J, Doilom M (2019) Fungal diversity notes 1036–1150: Taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 96(1): 1–242. https://doi.org/10.1007/s13225-019-00429-2
  • Hyde KD, Jeewon R, Chen YJ, Bhunjun CS, Calabon MS, Jiang HB, Lin CG, Norphanphoun C, Sysouphanthong P, Pem D, Tibpromma S, Zhang Q, Doilom M, Jayawardena RS, Liu JK, Maharachchikumbura SSN, Phukhamsakda C, Phookamsak R, Al-Sadi AM, Thongklang N, Wang Y, Gafforov Y, Gareth Jones EB, Lumyong S (2020) The numbers of fungi: Is the descriptive curve flattening? Fungal Diversity 103(1): 219–271. https://doi.org/10.1007/s13225-020-00458-2
  • Hyde KD, Norphanphoun C, Ma J, Yang HD, Zhang JY, Du TY, Gao Y, Gomes de Farias AR, He SC, He YK, Li CJY, Li JY, Liu XF, Lu L, Su HL, Tang X, Tian XG, Wang SY, Wei DP, Xu RF, Xu RJ, Yang YY, Zhang F, Zhang Q, Bahkali AH, Boonmee S, Chethana KWT, Jayawardena RS, Lu YZ, Karunarathna SC, Tibpromma S, Wang Y, Zhao Q (2023) Mycosphere notes 387–412 – novel species of fungal taxa from around the world. Mycosphere 14(1): 663–744. https://doi.org/10.5943/mycosphere/14/1/8
  • Hyde KD, Wijesinghe SN, Afshari N, Aumentado HD, Bhunjun CS, Boonmee S, Camporesi E, Chethana KWT, Doilom M, Dong W, Du TY, Farias ARG, Gao Y, Jayawardena RS, Karimi O, Karunarathna SC, Kularathnage ND, Lestari AS, Li CJY, Li YX, Liao CF, Liu XF, Lu L, Lu YZ, Luo ZL, Ma J, Mamarabadi M, Manawasinghe IS, Mapook A, Mi LX, Niranjan M, Senanayake IC, Shen HW, Su HL, Tibpromma S, Xu RJ, Yan JY, Yang YH, Yang YY, Yu FQ, Kang JC, Zhang JY (2024a) Mycosphere Notes 469–520. Mycosphere 15(1): 1294–1454. https://doi.org/10.5943/mycosphere/15/1/11
  • Hyde KD, Wijesinghe SN, Amuhenage TB, Bera I, Bhunjun CS, Chethana KWT, De Farias ARG, Dong W, Du TY, Gao Y, Gui H, Gunarathne A, Jayawardena RS, Li H, Liu JK, Madagammana AD, Manawasinghe IS, Mapook A, Mukhopadhyay S, Niranjan M, Phukhamsakda C, Ren GC, Senanayake IC, Shu YX, Thakshila SAD, Thongklang N, Tibpromma S, Tun ZL, Wu N, Xiong YR, Xu RF, Yang YH, Yu FQ, Zhao HJ (2024b) Current Research in Applied and Environmental Mycology Fungal Profiles 1–30. Current Research in Environmental & Applied Mycology 14: 167–266. [Journal of Fungal Biology]
  • Hyde KD, Noorabadi MT, Thiyagaraja V, He MQ, Johnston PR et al. (2024c) The 2024 Outline of Fungi and fungus-like taxa. Mycosphere 15(1): 5146–6239.
  • Jayasiri SC, Hyde KD, Ariyawansa HA, Bhat J, Buyck B, Cai L, Dai YC, Abd-Elsalam KA, Ertz D, Hidayat I, Jeewon R, Jones EBG, Bahkali AH, Karunarathna SC, Liu JK, Luangsa-ard JJ, Lumbsch HT, Maharachchikumbura SSN, McKenzie EHC, Moncalvo JM, Ghobad-Nejhad M, Nilsson H, Pang KL, Pereira OL, Phillips AJL, Raspé O, Rollins AW, Romero AI, Etayo J, Selçuk F, Stephenson SL, Suetrong S, Taylor JE, Tsui CKM, Vizzini A, Abdel-Wahab MA, Wen TC, Boonmee S, Dai DQ, Daranagama DA, Dissanayake AJ, Ekanayaka AH, Fryar SC, Hongsanan S, Jayawardena RS, Li WJ, Perera RH, Phookamsak R, de Silva NI, Thambugala KM, Tian Q, Wijayawardene NN, Zhao RL, Zhao Q, Kang JC, Promputtha I (2015) The Faces of Fungi database: Fungal names linked with morphology, phylogeny and human impacts. Fungal Diversity 74(1): 3–18. https://doi.org/10.1007/s13225-015-0351-8
  • Jayawardena RS, Hyde KD, Wang S, Sun YR, Suwannarach N, Sysouphanthong P, Abdel-Wahab MA, Abdel-Aziz FA, Abeywickrama PD, Abreu VP, Armand A, Aptroot A, Bao DF, Begerow D, Bellanger JM, Bezerra JDP, Bundhun D, Calabon MS, Cao T, Cantillo T, Carvalho JLVR, Chaiwan N, Chen CC, Courtecuisse R, Cui BK, Damm U, Denchev CM, Denchev TT, Deng CY, Devadatha B, de Silva NI, dos Santos LA, Dubey NK, Dumez S, Ferdinandez HS, Firmino AL, Gafforov Y, Gajanayake AJ, Gomdola D, Gunaseelan S, Shucheng-He Htet ZH, Kaliyaperumal M, Kemler M, Kezo K, Kularathnage ND, Leonardi M, Li J-P, Liao C, Liu S, Loizides M, Luangharn T, Ma J, Madrid H, Mahadevakumar S, Maharachchikumbura SSN, Manamgoda DS, Martín MP, Mekala N, Moreau P-A, Mu Y-H, Pahoua P, Pem D, Pereira OL, Phonrob W, Phukhamsakda C, Raza M, Ren G-C, Rinaldi AC, Rossi W, Samarakoon BC, Samarakoon MC, Sarma VV, Senanayake IC, Singh A, Souza MF, Souza-Motta CM, Spielmann AA, Su W, Tang X, Tian XG, Thambugala KM, Thongklang N, Tennakoon DS, Wannathes N, Wei DP, Welti S, Wijesinghe SN, Yang H, Yang Y, Yuan H-S, Zhang H, Zhang J, Balasuriya A, Bhunjun CS, Bulgakov TS, Cai L, Camporesi E, Chomnunti P, Deepika YS, Doilom M, Duan W-J, Han S-L, Huanraluek N, Jones EBG, Lakshmidevi N, Li Y, Lumyong S, Luo Z-L, Khuna S, Kumla J, Manawasinghe IS, Mapook A, Punyaboon W, Tibpromma S, Lu Y-Z, Yan JY, Wang Y (2023) Fungal diversity notes 1512–1610: Taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 117(1): 1–272. https://doi.org/10.1007/s13225-022-00513-0
  • Jiang HB, Jeewon R, Karunarathna SC, Phukhamsakda C, Doilom M, Kakumyan P, Suwannarach N, Phookamsak R, Lumyong S (2021) Reappraisal of Immotthia in Dictyosporiaceae, Pleosporales: Introducing Immotthia bambusae sp. nov. and Pseudocyclothyriella clematidis comb. et gen. nov. based on morphology and phylogeny. Frontiers in Microbiology 12: 656235. https://doi.org/10.3389/fmicb.2021.656235
  • Katoh K, Rozewicki J, Yamada KD (2019) MAFFT online service: Multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics 20(4): 1160–1166. https://doi.org/10.1093/bib/bbx108
  • Liu SL, Wang XW, Li GJ, Deng CY, Rossi W, Leonardi M, Liimatainen K, Kekki T, Niskanen T, Smith ME, Ammirati J, Bojantchev D, Abdel-Wahab MA, Zhang M, Tian E, Lu YZ, Zhang JY, Ma J, Dutta AK, Acharya K, Du TY, Xu J, Kim JS, Lim YW, Gerlach A, Zeng NK, Han YX, Razaghi P, Raza M, Cai L, Calabon MS, Jones EBG, Saha R, Kumar TKA, Krishnapriya K, Thomas A, Kaliyaperumal M, Kezo K, Gunaseelan S, Singh SK, Singh PN, Lagashetti AC, Pawar KS, Jiang S, Zhang C, Zhang H, Qing Y, Bau T, Peng XC, Wen TC, Ramirez NA, Niveiro N, Li MX, Yang ZL, Wu G, Tarafder E, Tennakoon DS, Kuo CH, da Silva TM, Souza-Motta CM, Bezerra JDP, He G, Ji XH, Suwannarach N, Kumla J, Lumyong S, Wannathes N, Rana S, Hyde KD, Zhou LW (2024a) Fungal diversity notes 1717–1817: Taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 124(1): 1–216. https://doi.org/10.1007/s13225-023-00529-0
  • Liu NG, Hyde KD, Sun YR, Bhat DJ, Jones EBG, Jumpathong J, Lin CG, Lu YZ, Yang J, Liu LL, Liu ZY, Liu JK (2024b) Notes, outline, taxonomy and phylogeny of brown-spored hyphomycetes. Fungal Diversity 128: 1–282. https://doi.org/10.1007/s13225-023-00529-0
  • Ma J, Gomdola D, Boonmee S, Shen HW, Tang X, Zhang LJ, Lu YZ, Hyde KD (2024) Three new species of Neohelicomyces (Tubeufiales, Tubeufiaceae) from freshwater and terrestrial habitats in China. MycoKeys 105: 317–336. https://doi.org/10.3897/mycokeys.105.124129
  • Maharachchikumbura SSN, Chen Y, Ariyawansa HA, Hyde KD, Haelewaters D, Perera RH, Samarakoon MC, Wanasinghe DN, Bustamante DE, Liu JK, Lawrence DP, Cheewangkoon R, Stadler M (2021) Integrative approaches for species delimitation in Ascomycota. Fungal Diversity 109(1): 155–179. https://doi.org/10.1007/s13225-021-00486-6
  • Manawasinghe IS, Calabon MS, Jones EBG, Zhang YX, Liao CF, Xiong Y, Chaiwan N, Kularathnage ND, Liu NG, Tang SM, Sysouphanthong P, Du TY, Luo M, Pasouvang P, Pem D, Phonemany M, Ishaq M, Chen JW, Karunarathna SC, Mai ZL, Rathnayaka AR, Samarakoon MC, Tennakoon DS, Wijesinghe SN, Yang YH, Zhao HJ, Fiaz M, Doilom M, Dutta AK, Khalid AN, Liu JW, Thongklang N, Senanayake IC, Tibpromma S, You LQ, Camporesi E, Gafforov YS, Hyde KD (2022) Mycosphere notes 345–386. Mycosphere 13(1): 454–557. https://doi.org/10.5943/mycosphere/13/1/3
  • Mena PJ, Mercado SA (1987) Piricaudiopsis (Hyphomycetes, Deuteromycotina), nuevo género enteroblástico de Cuba. Acta Botanica Cubana 51: 1–5.
  • Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the 2011 TeraGrid Conference: extreme digital discovery, 8 pp. https://doi.org/10.1145/2016741.2016785
  • Nguyen LT, Schmidt HA, Von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32(1): 268–274. https://doi.org/10.1093/molbev/msu300
  • O’Donnell K, Kistlerr HC, Cigelnik E, Ploetz RC (1998) Multiple evolutionary origins of the fungus causing panama disease of banana: Concordant evidence from nuclear and mitochondrial gene genealogies. Proceedings of the National Academy of Sciences of the United States of America 95(5): 2044–2049. https://doi.org/10.1073/pnas.95.5.2044
  • Pem D, Jeewon R, Chethana KWT, Hongsanan S, Doilom M, Suwannarach N, Hyde KD (2021) Species concepts of Dothideomycetes: Classification, phylogenetic inconsistencies and taxonomic standardization. Fungal Diversity 109(1): 283–319. https://doi.org/10.1007/s13225-021-00485-7
  • Pem D, Hyde KD, McKenzie EHC, Hongsanan S, Wanasinghe DN, Boonmee S, Darmostuk V, Bhat JD, Tian Q, Htet ZH, Senanayake IC, Niranjan M, Sarma VV, Doilom M, Dong W (2024) A comprehensive overview of genera in Dothideomycetes. Mycosphere 15(1): 2175–4568. https://doi.org/10.5943/mycosphere/15/1/18
  • Perera RH, Hyde KD, Jones EBG, Maharachchikumbura SSN, Bundhun D, Camporesi E, Akulov A, Liu JK, Liu ZY (2023) Profile of Bionectriaceae, Calcarisporiaceae, Hypocreaceae, Nectriaceae, Tilachlidiaceae, Ijuhyaceae fam. nov., Stromatonectriaceae fam. nov. and Xanthonectriaceae fam. nov. Fungal Diversity 118(1): 95–271. https://doi.org/10.1007/s13225-022-00512-1
  • Rambaut A, Drummond A (2014) FigTree v1. 3.1 Institute of Evolutionary Biology. University of Edinburgh.
  • Rathnayaka AR, Tennakoon DS, Jones GE, Wanasinghe DN, Bhat DJ, Priyashantha AH, Stephenson SL, Tibpromma S, Karunarathna SC (2024) Significance of precise documentation of hosts and geospatial data of fungal collections, with an emphasis on plant-associated fungi. New Zealand Journal of Botany 31: 1–28. https://doi.org/10.1080/0028825X.2024.2381734
  • Seifert K, Morgan-Jones G, Gams W, Kendrick B (2011) The genera of hyphomycetes. CBS Biodiversity Series 9: 1–997.
  • Senanayake IC, Rathnayaka AR, Marasinghe DS, Calabon MS, Gentekaki E, Lee HB, Hurdeal VG, Pem D, Dissanayake LS, Wijesinghe SN, Bundhun D, Nguyen TT, Goonasekara ID, Abeywickrama PD, Bhunjun CS, Jayawardena RS, Wanasinghe DN, Jeewon R, Bhat DJ, Xiang MM (2020) Morphological approaches in studying fungi: Collection, examination, isolation, sporulation and preservation. Mycosphere 11(1): 2678–2754. https://doi.org/10.5943/mycosphere/11/1/20
  • Senanayake IC, Rossi W, Leonardi M, Weir A, McHugh M, Rajeshkumar KC, Verma RK, Karunarathna SC, Tibpromma S, Ashtekar N, Ashtamoorthy SK, Raveendran S, Kour G, Singh A, De la Peña-Lastra S, Mateos A, Kolařík M, Antonín V, Ševčíková H, Esteve-Raventós F, Larsson E, Pancorbo F, Moreno G, Altés A, Turégano Y, Du TY, Lu L, Li QR, Kang JC, Gunaseelan S, Kezo K, Kaliyaperumal M, Fu J, Samarakoon MC, Gafforov Y, Teshaboeva S, Kunjan PC, Chamaparambath A, Flakus A, Etayo J, Rodriguez-Flakus P, Zhurbenko MP, de Silva NI, Tennakoon DS, Latha KPD, Manimohan P, Raj KNA, Calabon MS, Ahmadpour A, Heidarian Z, Alavi Z, Alavi F, Ghosta Y, Azizi R, Luo M, Zhao MP, Kularathnage ND, Hua L, Yang YH, Liao CF, Zhao HJ, Lestari AS, Jayasiri SC, Yu FM, Lei L, Liu JW, Karimi O, Tang SM, Sun YR, Wang Y, Zeng M, Htet ZH, Linaldeddu BT, Alves A, Phillips AJL, Bregant C, Montecchio L, De Kesel A, Hustad VP, Miller AN, Fedosova AG, Kučera V, Raza M, Hussain M, Chen YP, Thiyagaraja V, Gomdola D, Rathnayaka AR, Dissanayake AJ, Suwannarach N, Hongsanan S, Maharachchikumbura SSN, Dissanayake LS, Wijayawardene NN, Phookamsak R, Lumyong S, Jones EBG, Yapa N, Wanasinghe DN, Xie N, Doilom M, Manawasinghe IS, Liu JK, Zhao Q, Xu B, Hyde KD, Song J (2023) Fungal diversity notes 1611–1716: Taxonomic and phylogenetic contributions on fungal genera and species emphasis in south China. Fungal Diversity 122(1): 161–403. https://doi.org/10.1007/s13225-023-00523-6
  • Shenoy BD, Jeewon R, Wu WP, Bhat DJ, Hyde KD (2006) Ribosomal and RPB2 DNA sequence analyses suggest that Sporidesmium and morphologically similar genera are polyphyletic. Mycological Research 110(8): 916–928. https://doi.org/10.1016/j.mycres.2006.06.004
  • Sureshkumar G, Sharath Babu K, Kunwar IK, Manoharachary C (2005) Two new hyphomycetous fungal species from India. Mycotaxon 92: 279–283.
  • Tamura K, Peterson D, Stecher G, Filipski A, Kumar S (2013) MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution 30(12): 2725–2729. https://doi.org/10.1093/molbev/mst197
  • Tang X, Jeewon R, Lu YZ, Alrefaei AF, Jayawardena RS, Xu RJ, Ma J, Chen XM, Kang JC (2023) Morphophylogenetic evidence reveals four new fungal species within Tetraplosphaeriaceae (Pleosporales, Ascomycota) from tropical and subtropical forest in China. MycoKeys 100: 153–170. https://doi.org/10.3897/mycokeys.100.113141
  • Thiyagaraja V, Lücking R, Ertz D, Coppins BJ, Wanasinghe DN, Karunarathna SC, Suwannarach N, To-Anun C, Cheewangkoon R, Hyde KD (2021) Sequencing of the type species of Arthopyrenia places Arthopyreniaceae as a synonym of Trypetheliaceae. Mycosphere 12(1): 993–1011. https://doi.org/10.5943/mycosphere/12/1/10
  • Tian XG, Bao DF, Karunarathna SC, Jayawardena RS, Hyde KD, Bhat DJ, Luo ZL, Elgorban AM, Hongsanan S, Rajeshkumar KC, Maharachchikumbura SSN, Suwannarach N, Dawoud TM, Lu YZ, Han JJ, Xiao YP, Du TY, Lu L, Xu RF, Dai DQ, Liu XF, Liu C, Tibpromma S (2024) Taxonomy and phylogeny of Ascomycetes associated with selected economically important monocotyledons in China and Thailand. Mycosphere 15(1): 1–274. https://doi.org/10.5943/mycosphere/15/1/1
  • Tibpromma S, Hyde KD, McKenzie EHC, Bhat DJ, Phillips AJL, Wanasinghe DN, Samarakoon MC, Jayawardena RS, Dissanayake AJ, Tennakoon DS, Doilom M, Phookamsak R, Tang AMC, Xu J, Mortimer PE, Promputtha I, Maharachchikumbura SSN, Khan S, Karunarathna SC (2018) Fungal diversity notes 840–928: Micro-fungi associated with Pandanaceae. Fungal Diversity 93(1): 1–160. https://doi.org/10.1007/s13225-018-0408-6
  • Vaidya G, Lohman DJ, Meier R (2011) SequenceMatrix: Concatenation software for the fast assembly of multi-gene datasets with character set and codon information. Cladistics 27(2): 171–180. https://doi.org/10.1111/j.1096-0031.2010.00329.x
  • Wei GY, Mo WD, Hu Y, Zou MT, Wen JT, Chang LF, Chen YS, Norphanphoun C, Yang QI, Wang Y (2024) – Discovery of a Novel Occultibambusa Species (Occultibambusaceae, Dothideomycetes) from Poaceae Host from Guizhou Province, China. Current Research in Environmental & Applied Mycology 14: 146–156. [Journal of Fungal Biology]
  • Wijayawardene NN, Hyde KD, Al-Ani LKT, Tedersoo L, Haelewaters D, Rajeshkumar KC, Zhao RL, Aptroot A, Leontyev DV, Saxena RK, Tokarev YS, Dai DQ, Letcher PM, Stephenson SL, Ertz D, Lumbsch HT, Kukwa M, Issi IV, Madrid H, Phillips AJL, Selbmann L, Pfliegler WP, Horváth E, Bensch K, Kirk PM, Kolaříková K, Raja HA, Radek R, Papp V, Dima V, Ma J, Malosso E, Takamatsu S, Rambold G, Gannibal PB, Triebel D, Gautam AK, Avasthi S, Suetrong S, Timdal E, Fryar SC, Delgado G, Réblová M, Doilom M, Dolatabadi S, Pawłowska J, Humber RA, Kodsueb R, Sánchez-Castro I, Goto BT, Silva DKA, de Souza FA, Oehl F, da Silva GA, Silva IR, Błaszkowski J, Jobim K, Maia LC, Barbosa FR, Fiuza PO, Divakar PK, Shenoy BD, Castañeda-Ruiz RF, Somrithipol S, Lateef AA, Karunarathna SC, Tibpromma S, Mortimer PE, Wanasinghe DN, Phookamsak R, Xu J, Wang Y, Tian F, Alvarado P, Li DW, Kušan I, Matočec N, Maharachchikumbura SSN, Papizadeh M, Heredia G, Wartchow F, Bakhshi M, Boehm E, Youssef N, Hustad VP, Lawrey JD, Santiago ALCMA, Bezerra JDP, Souza-Motta CM, Firmino AL, Tian Q, Houbraken J, Hongsanan S, Tanaka K, Dissanayake AJ, Monteiro JS, Grossart HP, Suija A, Weerakoon G, Etayo J, Tsurykau A, Vázquez V, Mungai P, Damm U, Li QR, Zhang H, Boonmee S, Lu YZ, Becerra AG, Kendrick B, Brearley FQ, Motiejūnaitė J, Sharma B, Khare R, Gaikwad S, Wijesundara DSA, Tang LZ, He MQ, Flakus A, Rodriguez-Flakus P, Zhurbenko MP, McKenzie EHC, Stadler M, Bhat DJ, Liu JK, Raza M, Jeewon R, Nassonova ES, Prieto M, Jayalal RGU, Erdoğdu M, Yurkov A, Schnittler M, Shchepin ON, Novozhilov YK, Silva-Filho AGS, Liu P, Cavender JC, Kang Y, Mohammad S, Zhang LF, Xu RF, Li YM, Dayarathne MC, Ekanayaka AH, Wen TC, Deng CY, Pereira OL, Navathe S, Hawksworth DL, Fan XL, Dissanayake LS, Kuhnert E, Grossart HP, Thines M (2020) Outline of Fungi and fungus-like taxa. Mycosphere 11(1): 1060–1456. https://doi.org/10.5943/mycosphere/11/1/8
  • Wijayawardene NN, Hyde KD, Dai DQ, Sánchez-García M, Goto BT, Saxena RK, Erdoğdu M, Selçuk F, Rajeshkumar KC, Aptroot A, Błaszkowski J, Boonyuen N, da Silva GA, de Souza FA, Dong W, Ertz D, Haelewaters D, Jones EBG, Karunarathna SC, Kirk PM, Kukwa M, Kumla J, Leontyev DV, Lumbsch HT, Maharachchikumbura SSN, Marguno F, Martínez-Rodríguez P, Mešić A, Monteiro JS, Oehl F, Pawłowska J, Pem D, Pfliegler WP, Phillips AJL, Pošta A, He MQ, Li JX, Raza M, Sruthi OP, Suetrong S, Suwannarach N, Tedersoo L, Thiyagaraja V, Tibpromma S, Tkalčec Z, Tokarev YS, Wanasinghe DN, Wijesundara DSA, Wimalaseana SDMK, Madrid H, Zhang GQ, Gao Y, Sánchez-Castro I, Tang LZ, Stadler M, Yurkov A, Thines M (2022a) – Outline of Fungi and fungus-like taxa – 2021. Mycosphere 13(1): 53–453. https://doi.org/10.5943/mycosphere/13/1/2
  • Wijayawardene NN, Phillips AJL, Pereira DS, Dai DQ, Aptroot A, Monteiro JS, Druzhinina IS, Cai F, Fan X, Selbmann L, Coleine C, Castañeda-Ruiz RF, Kukwa M, Flakus A, Fiuza PO, Kirk PM, Kumar KCR, leperuma Arachchi IS, Suwannarach N, Tang L-Z, Boekhout T, Tan CS, Jayasinghe RPPK, Thines M (2022b) Forecasting the number of species of asexually reproducing fungi (Ascomycota and Basidiomycota). Fungal Diversity 114(1): 463–490. https://doi.org/10.1007/s13225-022-00500-5
  • Yang EF, Tibpromma S, Karunarathna SC, Phookamsak R, Xu JC, Zhao ZX, Karunanayake C, Promputtha I (2022) Taxonomy and phylogeny of novel and extant taxa in Pleosporales associated with Mangifera indica from Yunnan, China (Series I). Journal of Fungi 8(2): 152. https://doi.org/10.3390/jof8020152
  • Yang J, Liu LL, Jones EBG, Hyde KD, Liu ZY, Bao DF, Liu NG, Li WL, Shen HW, Yu XD, Liu JK (2023) Freshwater fungi from karst landscapes in China and Thailand. Fungal Diversity 119(1): 1–212. https://doi.org/10.1007/s13225-023-00514-7
  • Zhang K, Ma J, Wang Y, Zhang XG (2009) Three new species of Piricaudiopsis from southern China. Mycologia 101(3): 417–422. https://doi.org/10.3852/08-147
  • Zhang K, Ma L, Ma J, Castañeda-Ruiz RF (2014) Xiuguozhangia, a new genus of microfungi to accommodate five Piricaudiopsis species. Mycotaxon 128(1): 131–135. https://doi.org/10.5248/128.131
  • Zhang SN, Hyde KD, Jones EBG, Yu XD, Cheewangkoon R, Liu JK (2024) Current insights into palm fungi with emphasis on taxonomy and phylogeny. Fungal Diversity 127(1): 55–301. https://doi.org/10.1007/s13225-024-00536-9
login to comment