Discoveries of Dothideomycetes (Fungi) associated with pteridophytes in China

1 Discoveries of Dothideomycetes (Fungi) associated with pteridophytes in China Jing-Yi Zhang1,2,3,4,5 , Kevin D. Hyde4,5 , Ming-Fei Yang2, Ya-Ru Sun1,4,5 , Xing-Juan Xiao1,3,4,5 , Ze-Bin Meng6, Dan-Feng Bao7, Yong-Zhong Lu1,3 1 School of Food and Pharmaceutical Engineering, Guizhou Institute of Technology, Guiyang 550025, China 2 School of Public Health, Guiyang Healthcare Vocational University, Guiyang, Guizhou 550081, China 3 Guizhou Key Laboratory of Agricultural Microbiology, Guizhou Academy of Agricultural Sciences, Guiyang 550009, China 4 Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand 5 School of Science, Mae Fah Luang University, Chiang Rai 57100, Thailand 6 Guizhou Tea Seed Resource Utilization Engineering Research Center, Guizhou Education University, Guiyang 550018, China 7 Engineering and Research Center for Southwest Bio-Pharmaceutical Resources of National Education Ministry of China, Guizhou University, Guiyang 550025, China Corresponding authors: Yong-Zhong Lu ([email protected]); Ming-Fei Yang ([email protected]) Copyright: © Jing-Yi Zhang et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Pteridophytes are iconic symbols of the Earth’s biodiversity and harbor diverse fungal communities. In this study, an investigation of saprobic fungi associated with pteridophytes in China identified several fascinating taxonomic groups within Dothideomycetes. A polyphasic approach based on morphology, along with multi-gene phylogenetic analysis using combined LSU, RPB2, SSU, and tef1-α sequence data, revealed nine new collections representing five species in Pleosporales and five new collections representing two species in Muyocopronales. Consequently, six new genera (Cyatheomyces, Microlepicola, Neoberkleasmium, Pseudopalawaniella, Synnematospora, and Xenopleopunctum), six new species (Cyatheomyces synnematosus, Microlepicola guizhouensis, Pseudopalawaniella woodwardiae, Synnematospora pronephrii, Xenopleopunctum guizhouense, and X. sporodochiale), and five new combinations, viz., Neoberkleasmium micronesiacum (≡ Berkleasmium micronesiacum), N. nigroapicale (≡ B. nigroapicale), Xenoberkleasmium crinisium (≡ B. crunisia), X. pandani (≡ B. pandani), and X. typhae (≡ B. typhae), are proposed. Additionally, phylogenetic analysis reveals that four species of Xenoberkleasmium form a distinct lineage within Pleosporales, and they are evolving in a newly proposed family, Xenoberkleasmiaceae. Detailed morphological descriptions and a phylogenetic tree revealing the taxonomic placements of these new taxa are provided. Key words: Ascomycota, 13 new taxa, five new combinations, southwestern China, taxonomy Introduction Fungi associated with pteridophytes (ferns and their allies) play crucial roles in plant colonization and offer unique insights into plant–fungal symbiosis, yet remain understudied (Remy et al. 1994; Guatimosim et al. 2016; Pressel et al. 2016). The most recent study conducted by Zhang et al. (2025) provides a comprehensive Academic editor: Kazuaki Tanaka Received: 4 August 2025 Accepted: 5 December 2025 Published: 17 December 2025 Citation: Zhang J-Y, Hyde KD, Yang M-F, Sun Y-R, Xiao X-J, Meng Z-B, Bao D-F, Lu Y-Z (2025) Discoveries of Dothideomycetes (Fungi) associated with pteridophytes in China. IMA Fungus 16: e167717. https://doi. org/10.3897/imafungus.16.167717 IMA Fungus 16: e167717 (2025) DOI: 10.3897/imafungus.16.167717 2 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes global exploration of fungi associated with pteridophytes and highlights that, based on a conservative estimate, over 92% of these fungi remain undiscovered. Pleosporales was validly introduced by Barr (1987) and is the largest order within Dothideomycetes, Ascomycota, comprising approximately a quarter of all dothideomycetous species (Zhang et al. 2012a; Pem et al. 2024). Pleosporales, comprising approximately 93 families, represents a diverse and ecologically significant group of fungi that are globally distributed in terrestrial, marine, and freshwater habitats (Zhang et al. 2009, 2012a; Hongsanan et al. 2020a; Pem et al. 2024). These species include epiphytes, saprobes, endophytes, parasites, pathogens, lichens, and hyperparasites of fungi, insects, or mammals, with a notable prominence on plant hosts (Zhang et al. 2009, 2012a; Hyde et al. 2013; Li et al. 2023; Pem et al. 2024). Muyocopronales was introduced by Mapook et al. (2016b) to accommodate a single family, Muyocopronaceae. Phylogenetically, Palawaniaceae was introduced in the Dothideomycetes family incertae sedis (Mapook et al. 2016a; Hongsanan et al. 2020b) and was later presumed to belong to the Muyocopronales based on morphological similarity with Muyocopronales and the MCC tree (Mapook et al. 2016a, 2020a). However, subsequent studies did not support this placement (Hongsanan et al. 2020b; Mapook and Hyde 2023). Currently, Muyocopronaceae is the only family included in Muyocopronales (Senwanna et al. 2021; Xu et al. 2024). These species have a widespread distribution, commonly occurring as pathogens or saprobes on various plant substrates (Tibpromma et al. 2016; Crous et al. 2018a; Hernández-Restrepo et al. 2019; Senwanna et al. 2019a), sometimes occurring on soil (Madrid et al. 2012; Hernández-Restrepo et al. 2019), while some are pathogens infecting animals and humans (Hernández-Restrepo et al. 2019). In this study, we investigated the diversity of microfungi on decayed parts of ferns in China, obtaining 14 fresh collections representing seven Dothideomycetes species. Morphological and phylogenetic analyses provide further evidence for the classification of these species. This study is an extension of the research conducted by Zhang et al. (2025), focusing on fungal taxonomy with the aims of 1) investigating the diversity of saprobic fungi associated with ferns, drawing attention to this fungal group, and 2) describing novel taxa based on both morphological characteristics and phylogenetic evidence, enriching the fungal resources related to ferns. Methods Collections, morphology, and isolation Decayed rachides, or petioles, of ferns were collected from terrestrial habitats in China. Specimens were examined to determine the presence of fungi on the host substrate using a stereomicroscope (SMZ168-BL, Motic, China), as well as their macromorphological characters. Micro-morphological features of fungi were observed and photographed using an ECLIPSE Ni-U compound microscope (Nikon, Japan) fitted with an EOS 90D digital camera (Canon, Japan). Measurements were made by Tarosoft (R) Image Frame Work v.0.9.7. Photo-plates were processed and combined with Adobe Photoshop CC 2019 (Adobe Systems, USA). 3 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Single-spore isolations were made onto water agar (WA; 16 g/L distilled water) or potato dextrose agar (PDA; 39 g/L distilled water, Difco potato dextrose), and germinated spores were transferred onto PDA following the method in Senanayake et al. (2020). Pure culture plates were incubated at 26 °C for 3–8 weeks. Dried specimens were deposited in the Herbarium of Kunming Institute of Botany, Academia Sinica (HKAS), Chinese Academy of Sciences, Kunming, China, and the Herbarium of Guizhou Academy of Agricultural Sciences (GZAAS), Guiyang, China. Pure living cultures were deposited in the Kunming Institute of Botany Culture Collection (KUNCC) and Guizhou Culture Collection, China (GZCC). Names of the novel taxa were registered in Index Fungorum (http://www.indexfungorum.org). DNA extraction, amplification, sequencing Fungal genomic DNA was extracted from fresh mycelia grown on PDA, following the manufacturer’s instructions as described in the Biospin Fungus Genomic DNA Extraction Kit (Biospin Fungus Genomic DNA Extraction Kit, BioFlux®, Shanghai, China). Polymerase chain reactions (PCR) were carried out using the following primers: NS1 and NS4 (White et al. 1990), ITS5 and ITS4 (White et al. 1990), LR0R and LR5 (Vilgalys and Hester 1990), fRPB2-5F and fRPB2-7cR (Liu et al. 1999), and EF1-983F and EF1-2218R (Rehner and Buckley 2005). These primers were used to amplify the 18S subunit rDNA (SSU), the internal transcribed spacer (ITS), the large subunit of ribosomal DNA (LSU), the RNA polymerase II subunit 2 (RPB2), and the translation elongation factor 1 (tef1-α) gene regions, respectively. The amplification conditions were based on the protocol described by Zhang et al. (2021). The quality of the PCR products was checked on 1% agarose gel stained with ethidium bromide. Successful PCR products were sent to Sangon Biotech (Shanghai, China) for purification and sequencing. The sequences generated in this study have been deposited in NCBI GenBank. Sequence alignments and phylogenetic analysis Forward and reverse sequence reads were assembled using SeqMan v. 7.0.0 (DNASTAR, Madison, WI). Consensus sequences were subjected to a BLASTn search in NCBI GenBank (https://blast.ncbi.nlm.nih.gov/Blast. cgi) to select taxa for subsequent phylogenetic analyses. Consequently, representative sequence data of each family from Pleosporales, Muyocopronales, and several major related lineages used for phylogenetic analyses in Dothideomycetes were selected based on BLASTn searches, as well as recent publications (Table 1). Sequences newly generated in this study were deposited in GenBank. Sequence alignments for different gene loci were performed using the online multiple alignment program MAFFT version 7.2 (https://mafft.cbrc.jp/alignment/server/; Katoh et al. 2019). Trimal v. 1.2 was used to remove ambiguously aligned regions and uninformative positions with the “gt = 0.6” option (Capella-Gutiérrez et al. 2009). The obtained alignment was deposited in Figshare (https://figshare.com/; Suppl. material 1). Sequences of each locus were combined to form a concatenated supermatrix using SequenceMatrix 1.7.8 (Vaidya et al. 2011) and analyzed with maximum likelihood (ML) and Bayesian inference (BI). 4 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Table 1. Taxa used in this study and their GenBank accession numbers. Taxa Strain No. GenBank Accession Numbers References LSU RPB2 SSU tef1-α Acrocalymma medicaginis CPC 24340 KP170713 N/A N/A N/A Trakunyingcharoen et al. (2014) Acrocalymma pterocarpi MFLUCC 17-0926 MK347949 MK434897 MK347840 MK360040 Jayasiri et al. (2019) Aigialus parvus BCC 18403 GU479778 GU479817 GU479744 GU479842 Suetrong et al. (2009) Aigialus rhizophorae BCC 33572 GU479780 GU479819 GU479745 GU479844 Suetrong et al. (2009) Alternaria alternata CBS 916.96 DQ678082 DQ677980 DQ678031 DQ677927 Schoch et al. (2006a) Amniculicola aquatica MFLUCC 16-1123 MK106096 N/A N/A MK109800 Hyde et al. (2019) Amniculicola asexualis GZCC 20-0482 OP377926 OP473094 OP378011 OP473006 Yang et al. (2023) Amorocoelophoma cassiae MFLUCC 17-2283 MK347956 MK434894 NG_065775 MK360041 Jayasiri et al. (2019) Angustimassarina lonicerae MFLUCC 15-0087 KY496724 N/A N/A N/A Tibpromma et al. (2017) Anteaglonium parvulum SMH5223 GQ221909 N/A N/A GQ221918 Mugambi and Huhndorf (2009a) Aposphaeria corallinolutea MFLU 15-2752 KY554197 KY554207 KY554200 KY554205 Tibpromma et al. (2017) Aquasubmersa japonica HHUF 30469 NG_057138 LC194421 NG_062426 LC194384 Ariyawansa et al. (2015a); Hashimoto et al. (2017a) Aquasubmersa mircensis MFLUCC 11-0401 NG_042699 N/A NG_061141 N/A Zhang et al. (2012b) Arxiella longispora SGSF 303 MW519910 MW717995 N/A MW883564 Xu et al. (2023a) Ascocylindrica marina MD6011 KT252905 N/A KT252907 N/A Ariyawansa et al. (2015a) Ascocylindrica marina MF416 MK007123 N/A MK007124 N/A Ariyawansa et al. (2015a) Astragalicola vasilyevae MFLUCC 17-0832 MG828986 MG829248 MG829098 MG829193 Wanasinghe et al. (2018) Astrosphaeriella fusispora MFLUCC 10-0555 KT955462 KT955413 KT955443 KT955425 Phookamsak et al. (2015) Austropleospora ochracea GZCC 19-0430 MW133817 N/A MW134597 OP473010 Yang et al. (2023) Bahusandhika indica GUFCC 18001 KF460274 N/A N/A N/A Pratibha et al. (2014) Bambusicola bambusae MFLUCC 11-0614 JX442035 KP761718 JX442039 KP761722 Dai et al. (2012, 2015) Bambusicola massarinia MFLUCC 11-0389 NG_058658 KP761716 NG_061198 KP761725 Dai et al. (2012) Berkleasmium ariense NFCCI 4026 KY039165 N/A N/A N/A Tibpromma et al. (2017) Berkleasmium crunisia BCC 17023 DQ280271 N/A N/A N/A Pinnoi et al. (2007) Berkleasmium micronesiacum BCC 8141 DQ280272 N/A DQ280268 N/A Pinnoi et al. (2007) Berkleasmium nigroapicale BCC 8220 DQ280273 N/A DQ280269 N/A Pinnoi et al. (2007) Berkleasmium typhae BCC 12536 DQ280275 N/A N/A N/A Pinnoi et al. (2007) Botryosphaeria dothidea CMW 8000 KF766319 DQ677944 KF766233 DQ767637 Schoch et al. (2006a) Brevicollum versicolor HHUF 30591 NG_058716 LC271250 NG_065124 LC271246 Tanaka et al. (2017) Camarosporidiella caraganicola MFLUCCC 14-0605 KP711381 N/A KP711382 N/A Liu et al. (2015) Camarosporium quaternatum CPC 31081 NG_064442 N/A KY929123 KY929201 Crous and Groenewald (2017) Camarosporomyces flavigenus CBS 314.80 GU238076 N/A NG_061093 N/A Aveskamp et al. (2010) Capnodium coffeae CBS 147.52 DQ247800 KT216519 DQ247808 DQ471089 Schoch et al. (2006b); Spatafora et al. (2006); Ismail et al. (2016); Capnodium salicinum CBS 131.34 DQ678050 KT216553 DQ677997 DQ677889 Schoch et al. (2006a) Clematidis italica MFLUCC 15-0084 KU842381 N/A NG_061236 N/A Li et al. (2016) Clypeoloculus akitaensis KT 788 AB807543 N/A AB797253 AB808519 Tanaka et al. (2015) Coniothyrium palmarum CBS 400.71 JX681084 KT389592 EU754054 DQ677903 Schoch et al. (2006a) Corynespora cassiicola CBS 100822 GU301808 GU371742 GU296144 GU349052 Schoch et al. (2009) Corynespora torulosa CPC 15989 KF777207 N/A N/A N/A Crous et al. (2013) Crassiperidium octosporum MAFF 242971 NG_066389 LC373132 NG_065689 LC373120 Matsumura et al. (2018) Cryptocoryneum japonicum HHUF 30482 NG_059035 LC194438 NG_065118 LC096144 Hashimoto et al. (2017a) Cryptocoryneum pseudorilstonei CBS 113641 NG_059036 LC194446 LC194322 LC096152 Hashimoto et al. (2017a) Cucurbitaria berberidis MFLUCC 11-0387 KC506796 N/A KC506800 N/A Doilom et al. (2013) Cyatheomyces synnematosus KUNCC 23-13865 PV862390 PV948862 N/A PV948875 This study Cyatheomyces synnematosus GZCC 23-0670 PV862391 PV948863 N/A PV948876 This study Cyatheomyces synnematosus KUNCC 23-14156 PV862392 PV948864 N/A PV948877 This study 5 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Taxa Strain No. GenBank Accession Numbers References LSU RPB2 SSU tef1-α Cyatheomyces synnematosus KUNCC 23-14159 PV862393 PV948865 N/A PV948878 This study Cyclothyriella rubronotata CBS 141486 KX650544 KX650574 NG_061252 KX650519 Jaklitsch et al. (2016) Cyclothyriella rubronotata CPC 27604 MH107933 N/A N/A N/A Crous et al. (2018b) Dacampia engeliana Hafellner 72868 KT383791 N/A N/A N/A Ertz et al. (2015) Dacampia hookeri Hafellner 73897 KT383792 N/A N/A N/A Ertz et al. (2015) Delitschia chaetomioides SMH 3253.2 GU390656 N/A N/A GU327753 Mugambi and Huhndorf (2009b) Delitschia winteri AFTOL-ID 1599 DQ678077 DQ677975 DQ678026 DQ677922 Schoch et al. (2006a) Dendrographa decolorans Ertz 5003 (BR) AY548815 DQ883715 AY548809 DQ883725 Spatafora et al. (2006) Dictyocheirospora bannica KH 332 AB807513 N/A AB797223 AB808489 Tanaka et al. (2015) Didymella exigua CBS 183.55 MH868977 GU371764 N/A N/A Schoch et al. (2009); Vu et al. (2019) Didymella rumicicola CBS 683.79 MH873007 KT389622 N/A N/A Vu et al. (2019) Didymosphaeria rubi-ulmifolii MFLUCC 14-0023 KJ436586 N/A KJ436588 N/A Ariyawansa et al. (2014) Digitodesmium chishuiense GZCC 20-0510 OP377907 OP473082 OP377993 OP472990 Yang et al. (2023) Dothidotthia aceris MFLUCC 16-1183 MK751816 N/A MK751761 N/A Senwanna et al. (2019b) Equiseticola fusispora MFLUCC 14-0522 NG_059249 N/A NG_061238 MG520895 Abd-Elsalam et al. (2016) Fuscostagonospora cytisi MFLUCC 16-0622 KY770978 N/A KY770977 KY770979 Hyde et al. (2017) Fuscostagonospora sasae HHUF 29106 AB807548 N/A AB797258 AB808524 Tanaka et al. (2015) Fusculina eucalypti CBS 145083 MK047499 N/A N/A N/A Crous et al. (2018a) Gordonomyces mucovaginatus CBS 127273 NG_057941 N/A N/A N/A Crous et al. (2011a) Halobyssothecium aquifusiforme GZCC 20-0481 OP377925 OP473093 OP378010 OP473005 Yang et al. (2023) Halojulella avicenniae BCC 20173 GU371822 GU371786 GU371830 GU371815 Schoch et al. (2009) Halojulella avicenniae JK 5326A GU479790 N/A GU479756 N/A Suetrong et al. (2009) Halotthia posidoniae BBH 22481 GU479786 N/A GU479752 N/A Suetrong et al. (2009) Helicosporium liuzhouense GZCC 22-2014 OQ981402 OQ980474 N/A OQ980476 Xiao et al. (2023) Helminthosporium velutinum L131 KY984352 KY984413 KY984432 KY984463 Voglmayr and Jaklitsch (2017) Hermatomyces iriomotensis HHUF 30518 LC194367 LC194449 LC194325 LC194394 Hashimoto et al. (2017a) Hermatomyces jinghaensis HKAS 112167 MW989519 N/A N/A MZ042642 Ren et al. (2021) Hermatomyces turbinatus MFLUCC 21-0038 MW989518 MZ042638 N/A MZ042641 Ren et al. (2021) Hyphodiscosia jaipurensis MFLU 23-0472 PP112042 N/A PP101307 N/A Xu et al. (2024) Hyphodiscosia jaipurensis MFLUCC 23-0302 PP112041 N/A PP101306 N/A Xu et al. (2024) Hypsostroma caimitalense GKM1165 GU385180 N/A N/A N/A Mugambi and Huhndorf (2009b) Hypsostroma saxicola SMH 5005 GU385181 N/A N/A N/A Mugambi and Huhndorf (2009b) Hysterium angustatum CBS 123334 FJ161207 FJ161129 FJ161167 FJ161111 Boehm et al. (2009) Hysterobrevium smilacis CBS 114601 FJ161174 FJ161114 FJ161135 FJ161091 Boehm et al. (2009) Latorua caligans CBS 576.65 NG_058180 N/A N/A N/A Crous et al. (2015) Latorua grootfonteinensis CBS 369.72 NG_058181 N/A N/A N/A Crous et al. (2015) Lecanactis abietina Ertz 5068 (BR) AY548812 AH013900 AY548805 N/A Lutzoni et al. (2004) Lentimurispora urniformis MFLUCC 18-0497 MH179144 N/A MH179160 MH188055 Liu et al. (2018a) Lentithecium pseudoclioninum HHUF 29055 NG_059392 N/A NG_064847 AB808521 Tanaka et al. (2015) Leptodiscella africana CBS 400.65 MH870275 MK492711 N/A MK495955 Hernández-Restrepo et al. (2019) Leptodiscella sexualis MFLU 19-2783 MW293930 N/A N/A N/A Tennakoon et al. (2021) Leptosphaeria cichorium MFLUCC 14-1063 KT454712 N/A KT454728 N/A Ariyawansa et al. (2015b) Leptoxyphium fumago CBS 123.26 GU301831 GU371741 GU296161 GU349051 Schoch et al. (2009) Libertasomyces myopori CPC 27354 NG_058241 N/A N/A N/A Crous et al. (2016) Ligninsphaeria jonesii GZCC 15-0080 KU221038 N/A N/A N/A Zhang et al. (2016a) Ligninsphaeria jonesii MFLUCC 15-0641 NG_059642 N/A N/A N/A Zhang et al. (2016a) Lignosphaeria thailandica MFLUCC 11-0376 KP888645 N/A N/A N/A Thambugala et al. (2015) Lindgomyces cigarosporus G619 KX655804 N/A KX655805 N/A Raja et al. (2017) Lindgomyces ingoldianus ATCC 200398 AB521736 N/A NG_016531 N/A Hirayama et al. (2010) 6 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Taxa Strain No. GenBank Accession Numbers References LSU RPB2 SSU tef1-α Longipedicellata aptrootii MFLU 10-0297 KU238894 KU238891 KU238895 KU238892 Zhang et al. (2016b) Longipedicellata aquatica MFLUCC 15-0630 OP377961 OP473115 OP378038 OP473053 Yang et al. (2023) Lophiostoma macrostomum KT508 AB619010 JN993491 AB618691 LC001751 Hirayama and Tanaka (2011); Schoch et al. (2012); Thambugala et al. (2015) Lophiotrema fallopiae MAFF 245612 LC149915 LC194459 LC149911 LC194404 Hyde et al. (2016) Lophiotrema nucula CBS 627.86 GU301837 GU371792 GU296167 GU349073 Schoch et al. (2009) Macrodiplodiopsis desmazieri CBS 140062 NG_058182 N/A N/A N/A Crous et al. (2015) Macrodiplodiopsis desmazieri CBS 222.37 KR909316 KR909322 KR909318 KR909319 Ahmed et al. (2015) Magnicamarosporium iriomotense HHUF 30125 NG_059389 N/A AB797219 AB808485 Tanaka et al. (2015) Massaria anomia CBS 591.78 GU301839 GU371769 GU296169 N/A Schoch et al. (2009) Massaria inquinans WU 30527 HQ599402 HQ599460 HQ599444 HQ599342 Voglmayr et al. (2011) Mauritiana rhizophorae BCC 28866 GU371824 N/A GU371832 GU371817 Schoch et al. (2009) Melanomma japonicum MAFF 239634 NG_060360 LC203395 NG_065122 LC203367 Hashimoto et al. (2017b) Melanomma pulvis-pyrius CBS 124080 MH874873 GU456350 GU456302 GU456265 Zhang et al. (2009) Microlepicola guizhouensis KUNCC 23-14007 PV862399 PV948870 PV862385 PV948884 This study Morosphaeria muthupetensis PUFD87 MF614796 N/A MF614797 MF614798 Devadatha et al. (2018) Morosphaeria velatispora KH 221 AB807556 N/A AB797266 AB808532 Tanaka et al. (2015) Muyocopron chromolaenae MFLUCC 17-1513 NG_068700 MT136761 NG_070150 MT136756 Mapook et al. (2020a) Muyocopron dipterocarpi MFLUCC 14-1103 KU726966 KY225779 KU726969 MT136754 Mapook et al. (2016b) Mycoleptodiscus endophyticus MFLUCC 17-0545 MG646946 N/A NG_065724 MG646985 Tibpromma et al. (2018b) Mycoleptodiscus suttonii CBS 276.72 MK487728 MK492732 N/A MK495974 Hernández-Restrepo et al. (2019) Mycoleptodiscus terrestris IMI 159038 MK487731 MK492735 N/A MK495977 Hernández-Restrepo et al. (2019) Neoastrosphaeriella krabiensis MFLUCC 11-0025 JN846729 N/A JN846739 N/A Liu et al. (2011) Neocamarographium carpini CBS 128781 JQ044450 N/A N/A N/A Crous et al. (2011b) Neocamarosporium goegapense CBS 138008 KJ869220 N/A N/A N/A Crous et al. (2014) Neocamarosporium phragmitis MFLUCC 17-0756 NG_070431 N/A NG_065736 MG844351 Hyde et al. (2018) Neocochlearomyces chromolaenae BCC 68250 NG_066431 N/A NG_065766 MK047573 Crous et al. (2018a) Neocochlearomyces chromolaenae BCC 68252 MK047516 N/A MK047554 MK047575 Crous et al. (2018a) Neodeightonia palmicola MFLUCC 10-0822 HQ199222 N/A HQ199223 N/A Liu et al. (2010) Neohendersonia kickxii CBS 112403 NG_058264 N/A N/A N/A Giraldo et al. (2017) Neokalmusia aquibrunnea GZCC 17-0045 OP377920 N/A OP378005 OP473000 Yang et al. (2023) Neomassaria fabacearum MFLUCC 16-1875 KX524145 N/A NG_061245 KX524149 Hyde et al. (2016) Neomassaria formosana NTUCC 17-007 MH714756 MH714765 MH714759 MH714762 Ariyawansa et al. (2018) Neomassarina chromolaenae MFLUCC 17-1480 NG_068715 MT235822 NG_070168 MT235785 Mapook et al. (2020a) Neomassarina thailandica MFLUCC 17-1432 MT214467 MT235823 MT214420 MT235786 Mapook et al. (2020a) Neomassariosphaeria aquimucosa GZCC 19-0500 MW133803 N/A MW134591 OP473014 Yang et al. (2023) Neomycoleptodiscus alishanense MFLUCC 19-0390 ON024150 N/A N/A N/A Liu et al. (2024b) Neomycoleptodiscus venezuelense CBS 100519 NG_066340 MK492736 N/A MK495978 Hernández-Restrepo et al. (2019) Neooccultibambusa thailandensis MFLUCC 16-0274 NG_068827 MH412758 MH260348 MH412780 Tibpromma et al. (2018a) Neopaucispora rosaecae MFLUCC 17-0807 MG829033 N/A NG_061293 MG829217 Wanasinghe et al. (2018) Neophaeosphaeria agaves CPC 21264 KF777227 N/A N/A N/A Crous et al. (2013) Neophaeosphaeria filamentosa CBS 102202 GQ387577 GU371773 GQ387516 GU349084 de Gruyter et al. (2013) Neophaeosphaeria phragmiticola KUMCC 16-0216 MG837009 N/A NG_065735 MG838020 Hyde et al. (2018) Neopyrenochaeta acicola CBS 812.95 GQ387602 LT623271 NG_065567 N/A de Gruyter et al. (2010); ValenzuelaLopez et al. (2018) Neopyrenochaeta cercidis MFLU 18-2089 MK347932 MK434908 MK347823 N/A Jayasiri et al. (2019) Neoroussoella bambusae MFLUCC 11-0124 KJ474839 KJ474856 N/A KJ474848 Liu et al. (2014) Nigrograna fuscidula CBS 141556 KX650550 N/A N/A KX650525 Jaklitsch and Voglmayr (2016) Nigrograna mackinnonii CBS 674.75 GQ387613 KF015703 NG_061081 KF407986 de Gruyter et al. (2010); Ahmed et al. (2014) 7 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Taxa Strain No. GenBank Accession Numbers References LSU RPB2 SSU tef1-α Nigrograna obliqua CBS 141475 KX650558 KX650579 KX650512 KX650530 Jaklitsch and Voglmayr (2016) Occultibambusa bambusae MFLUCC 13-0855 KU863112 KU940170 KU872116 KU940193 Dai et al. (2016) Occultibambusa jonesii GZCC 16-0117 KY628322 KY814758 KY628324 KY814756 Zhang et al. (2017) Ohleria modesta OM KX650563 KX650583 KX650513 KX650534 Jaklitsch et al. (2016) Ohleria modesta MGC KX650562 KX650582 N/A KX650533 Jaklitsch et al. (2016) Palawania thailandensis MFLU 16-1871 KY086494 N/A N/A N/A Mapook et al. (2016a) Palawania thailandensis MFLUCC 14-1121 NG_241882 KY086496 NG_242378 N/A Mapook et al. (2016a) Parabambusicola bambusina KH 139 AB807537 N/A AB797247 AB808512 Tanaka et al. (2015) Paradictyoarthrinium diffractum MFLUCC 13-0466 KP744498 KX437764 KP753960 N/A Liu et al. (2015) Paradictyoarthrinium hydei MFLUCC 17-2512 NG_067558 MG780232 NG_065757 N/A Liu et al. (2018b) Paralophiostoma hysterioides PUFNI 17617 MT912850 MT926117 MT914175 N/A Hongsanan et al. (2020a) Paramycoleptodiscus albizziae CBS 141320 KX228330 MK492737 N/A MK495979 Hernández-Restrepo et al. (2019) Parapyrenochaeta protearum CBS 131315 JQ044453 LT717683 N/A N/A Valenzuela-Lopez et al. (2018) Paratrimmatostroma kunmingensis KUN-HKAS 102224A MK098196 N/A MK098204 MK098208 Jayawardena et al. (2022) Periconia delonicis MFLUCC 17-2584 NG_068611 MK434901 NG_065770 MK360071 Jayasiri et al. (2019) Periconia pseudodigitata KT 1395 AB807564 N/A AB797274 AB808540 Tanaka et al. (2015) Phaeoseptum terricola MFLUCC 10-0102 MH105779 MH105782 NG_065749 MH105781 Hyde et al. (2018) Phaeosphaeria oryzae CBS 110110 KF251689 KF252193 GQ387530 ON419509 de Gruyter et al. (2010); Quaedvlieg et al. (2013); Ashrafi et al. (2023) Phaeosphaeriopsis triseptata MFLUCC 13-0271 KJ522479 KJ522485 KJ522484 MG520919 Thambugala et al. (2014) Phaeotrichum benjaminii CBS 541.72 AY004340 DQ677946 AY016348 DQ677892 Lumbsch et al. (2005); Schoch et al. (2006a) Pleomonodictys capensis CBS 968.97 KY853521 N/A N/A N/A Hernández-Restrepo et al. (2017) Pleomonodictys descalsii FMR 12716 KY853522 N/A N/A N/A Hernández-Restrepo et al. (2017) Preussia funiculata CBS 659.74 GU301864 GU371799 GU296187 GU349032 de Gruyter et al. (2009) Prosthemium betulinum CBS 279.74 MH872591 KT216532 DQ678027 DQ677923 Schoch et al. (2006a); Vu et al. (2019) Prosthemium stellare CBS 126964 MH875800 N/A AB553650 N/A Tanaka et al. (2010); Vu et al. (2019) Protofenestella ulmi FP5 = CBS 143000 MF795791 MF795833 N/A MF795879 Jaklitsch et al. (2018) Psedotubeufia laxispora GZCC 22-2011 OR030831 OR046682 N/A OR046675 Ma et al. (2023a) Pseudoastrosphaeriella bambusae MFLUCC 11-0205 KT955475 KT955414 KT955455 KT955437 Phookamsak et al. (2015) Pseudoastrosphaeriella longicolla MFLUCC 11-0171 KT955476 KT955420 N/A KT955438 Phookamsak et al. (2015) Pseudoastrosphaeriella thailandensis MFLUCC 11-0144 KT955478 KT955416 KT955457 KT955440 Phookamsak et al. (2015) Pseudoberkleasmium chiangmaiense MFLUCC 17-1809 MK131260 N/A N/A MK131261 Hyde et al. (2019) Pseudoberkleasmium pandanicola KUMCC 17-0178 MH260304 N/A MH260344 N/A Tibpromma et al. (2018a) Pseudocoleodictyospora tectonae MFLUCC 12-0385 KU764709 KU712491 NG_061232 N/A Doilom et al. (2017) Pseudocoleodictyospora thailandica MFLUCC 12-0565 KU764701 KU712494 NG_062417 N/A Doilom et al. (2017) Pseudolophiotrema elymicola KT 1450 LC194381 LC194473 LC194339 LC194418 Hashimoto et al. (2017a) Pseudomassarina clematidis MFLU 16-0493 NG_073850 MT394700 NG_070663 MT394644 Thambugala et al. (2015) Pseudopalawania siamensis MFLUCC 17-1476a N/A N/A MT137789 MT136752 Mapook et al. (2020b) Pseudopalawania siamensis MFLUCC 17-1476b NA N/A MT137790 N/A Mapook et al. (2020b) Pseudopalawaniella woodwardiae KUNCC 23-13877 PV862394 PV948866 PV862381 PV948879 This study Pseudopyrenochaeta lycopersici CBS 306.65 EU754205 LT717680 NG_062728 N/A Valenzuela-Lopez et al. (2018) Pyrenochaetopsis leptospora CBS 101635 GQ387627 LT623282 NG_063097 MF795881 de Gruyter et al. (2010) Pyrenochaetopsis tabarestanensis IBRC M 30051 KF803343 N/A NG_065034 N/A Papizadeh et al. (2017) Pyrenophora phaeocomes AFTOL-ID 283 NG_027575 DQ497614 JN940960 DQ497607 Goonasekara et al. (2020） Quadrisporella heveae MFLUCC 18-0308 OL782057 OL828755 N/A OL875101 Senwanna et al. (2021) Quercicola fusiformis MFLUCC 18-0479 MK348009 MK434864 MK347898 MK360085 Jayasiri et al. (2019) Quercicola guttulospora MFLUCC 18-0481 MK348010 N/A MK347899 MK360086 Jayasiri et al. (2019) 8 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Taxa Strain No. GenBank Accession Numbers References LSU RPB2 SSU tef1-α Quixadomyces cearensis HUEFS 238438 MG970695 N/A N/A N/A Crous et al. (2018c) Ramusculicola thailandica MFLUCC 13-0284 KP888647 N/A KP899131 KR075167 Thambugala et al. (2015) Roccella fuciformis Tehler 8171 FJ638979 N/A N/A N/A Zhang et al. (2019) Rostriconidium pandanicola KUMCC 17-0176 NG_068830 MH412759 MH260358 MH412781 Tibpromma et al. (2018a) Roussoella nitidula MFLUCC 11-0634 KJ474842 KJ474858 N/A KJ474851 Liu et al. (2014) Salsuginea phoenicis MFLU 19-0015 MK405280 N/A N/A MK404650 Jones er al. (2019) Salsuginea ramicola KT 2597.2 GU479801 GU479834 GU479768 GU479862 Suetrong et al. (2009) Setoapiospora thailandica AND3 OL457707 N/A OL700220 OL998895 de Silva et al. (2022) Setoapiospora thailandica MFLUCC 17-1426 NG_068914 N/A NG_068420 MN648731 Hyde et al. (2020) Shiraia bambusicola GZAAS2.629 KC460980 N/A N/A N/A Liu et al. (2013) Sirodesmium olivaceum CBS 395.59 GU250894 GU250947 GU250915 N/A Ruibal et al. (2009) Sporormia fimetaria Gr.81.194 GQ203729 N/A N/A N/A Kruys and Wedin (2009) Stemphylium vesicarium CBS 191.86 GU238160 DQ247794 GU238232 DQ471090 Schoch et al. (2006b); Spatafora et al. (2006); Aveskamp et al. (2010) Striatiguttula nypae MFLUCC 18-0265 MK035992 MK034440 MK035977 MK034432 Zhang et al. (2019) Striatiguttula phoenicis MFLUCC 18-0266 MK035995 MK034442 MK035980 MK034435 Zhang et al. (2019) Subglobosporium tectonae MFLUCC 12-0393 KU764703 KU712485 NG_061233 N/A Doilom et al. (2017) Subplenodomus violicola CBS 306.68 MH870849 N/A GU238231 N/A Aveskamp et al. (2010); Vu et al. (2019) Sulcatispora acerina KUMCC 21-0821 ON009112 ON009294 ON009096 ON009271 Wanasinghe et al. (2022) Sulcatispora acerina KT 2982 LC014610 N/A LC014605 LC014615 Tanaka et al. (2015) Sulcosporium thailandica MFLUCC 12-0004 KT426563 N/A KT426564 N/A Ariyawansa et al. (2015a) Synnematospora pronephrii KUNCC 23-13965 PV862395 N/A PV862382 PV948880 This study Teichospora trabicola C134 KU601591 KU601600 N/A KU601601 Jaklitsch et al. (2016) Tetraploa bambusae KUMCC 21-0844 ON077067 N/A ON077073 ON075061 Phookamsak et al. (2022) Tetraploa cylindrica KUMCC 20-0205 MT893204 N/A MT893203 N/A Liao et al. (2022) Tetraploa dashaoensis KUMCC 21-0010 OL473555 N/A OL473556 N/A Jayawardena et al. (2022) Thyridaria broussonetiae TB1 = CBS 141481 KX650568 KX650586 KX650515 KX650539 Jaklitsch and Voglmayr (2016) Torula aquatica MFLUCC 16-1115 MG208146 MG207977 N/A N/A Su et al. (2018) Torula pluriseptata MFLUCC 14-0437 KY197855 KY197869 KY197862 KY197875 Li et al. (2017) Trematosphaeria grisea CBS 332.50 NG_057979 KF015720 NG_062930 KF015698 Ahmed et al. (2014) Trematosphaeria pertusa CBS 122368 NG_057809 FJ795476 FJ201991 KF015701 Ahmed et al. (2014) Trichodelitschia munkii Kruys 201 (UPS) DQ384096 N/A DQ384070 N/A Kruys et al. (2006) Tubeufia muriformis GZCC 22-2039 OR030836 OR046686 N/A OR046680 Ma et al. (2023b) Tzeanania taiwanensis NTUCC 17-006 MH461121 MH461129 MH461127 MH461131 Ariyawansa et al. (2018) Verruculina enalia BCC 18402 GU479803 GU479836 GU479771 GU479864 Suetrong et al. (2009) Westerdykella angulata CBS 610.74 NG_057754 N/A NG_062146 GU371821 Kruys et al. (2006) Wicklowia aquatica CBS 125634 MH875044 N/A NG_061099 N/A Schoch et al. (2009); Vu et al. (2019) Wicklowia submersa MFLUCC 18-0373 MK637644 N/A MK637643 N/A Boonmee et al. (2019) Xenoberkleasmium chiangraiense GZAAS 24-0051 PP657340 PP887797 N/A N/A Liu et al. (2024a) Xenoberkleasmium pandani KUNCC 23-13876 PV862396 PV948867 N/A PV948881 This study Xenoberkleasmium pandani KUNCC 23-13878 PV862397 PV948868 PV862383 PV948882 This study Xenoberkleasmium pandani KUNCC 23-14012 PV862398 PV948869 PV862384 PV948883 This study Xenopleopunctum guizhouense KUNCC 23-13881 PV862400 PV948871 PV862386 PV948885 This study Xenopleopunctum guizhouense KUNCC 23-13880 PV862401 PV948872 PV862387 PV948886 This study Xenopleopunctum guizhouense KUNCC 23-13882 PV862402 PV948873 PV862388 PV948887 This study Xenopleopunctum sporodochiale GZCC 23-0742 PV862403 PV948874 PV862389 PV948888 This study Xenopyrenochaetopsis pratorum CBS 445.81 GU238136 KT389671 NG_062792 N/A Aveskamp et al. (2010) 9 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes ML and BI analyses were performed through the CIPRES Science Gateway (Miller et al. 2010). ML analysis was conducted with RAxML-HPC v. 8.2.12 (Stamatakis 2014) using a GTRGAMMA approximation with rapid bootstrap analysis followed by 1000 bootstrap replicates. BI analysis was performed in a likelihood framework implemented on XSEDE 3.2.7a (Ronquist et al. 2012). Bayesian posterior probabilities (PPs) (Rannala and Yang 1996; Huelsenbeck 2001) were evaluated based on Markov Chain Monte Carlo (MCMC) sampling. Four simultaneous Markov chains were run for 100,000,000 generations, or the searches were stopped when the average standard deviation of split frequencies was below 0.01 (stopval = 0.01). Trees were sampled every 1000 generations, yielding 100,000 trees in total. The first 25% of trees were set as burn-in and discarded. The remaining trees were used to calculate PPs (Larget and Simon 1999). The phylogenetic tree was visualized using FigTree v. 1.4.4 (Rambaut et al. 2014), and the layouts were reorganized online following the methods described in Xie et al. (2023) and finalized with Adobe Illustrator CS6 (Adobe Systems, USA). Results Phylogenetic result A combined dataset of LSU, RPB2, SSU, and tef1-α sequence data was used to evaluate the phylogenetic placement of the new taxa in Pleosporales and Muyocopronales, Dothideomycetes. The concatenated sequence matrix comprised 234 taxa with representative taxa of Pleosporales and Muyocopronales and several major groups in Dothideomycetes, with three species of Arthoniomycetes as outgroup taxa. After alignment, the dataset contained 3,785 characters (LSU: 1–853; RPB2: 854–1,870 bp; SSU: 1,871–2,879, tef1-α: 2,880–3,785), including 2,589 distinct alignment patterns, with 32.15% comprising undetermined characters or gaps, 1,717 parsimony-informative sites, 426 singleton sites, and 1,642 constant sites. The total tree length is 27.938381. The best-scoring RAxML tree is shown in Fig. 1, with a final likelihood value of −112695.550615. Our 14 fresh collections representing seven species are placed in two orders (Muyocopronales and Pleosporales). Among these, five isolates represent two distinct, independent lineages identified as Cyatheomyces synnematosus and Pseudopalawaniella woodwardiae in Muyocopronaceae, Muyocopronales. Notably, these two lineages share a sister relationship with high support (100% ML/1.00 PP, Fig. 1). The analysis confirms that Lentimurisporaceae is a monophyletic family within Pleosporales. Synnematospora pronephrii, representing a newly proposed genus, is sister to the genus Bahusandhika with high support. Another new genus, Neoberkleasmium, introduced in Lentimurisporaceae, forms a distinct clade that includes two new combinations transferred from Berkleasmium. The phylogenetic tree reveals that our three new collections representing Xenoberkleasmium pandani (≡ Berkleasmium pandani) and two additional Berkleasmium species cluster with X. chiangraiense, the type species of Xenoberkleasmium. These four species form an independent, monophyletic clade with strong support (100% ML/1.00 PP, Fig. 1), representing a new family, Xenoberkleasmiaceae, which is closely related to Hypsostromataceae. 16 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Figure 3. Cyatheomyces synnematosus (HKAS 129695, paratype). A Colonies on host substrate; B, C Conidiophores with conidia; D–F Conidiogenous cells with conidia; G, H Conidia. Scale bars: 50 µm (B, C); 10 µm (D–H). 17 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes with entire margin, flat with a protuberance in the center, veined, from which several indentations extend outwards, and cut into fan shapes at the surface, white form above; brown form below, and not producing pigmentation in cultures. Material examined. CHINA • Guizhou Province, Zunyi City, Chishui County, Hushi Town, Chishui Alsophila Natural Reserve, on dead frond stalks of Cyathea sp. (Cyatheaceae), 27 July 2022, J.Y. Zhang, CX4 (HKAS 129699, holotype; GZAAS 23-0669, isotype), ex-type living culture, KUNCC 23-13865; • ibid., 22 September 2019, J.Y. Zhang, C4 (HKAS 129695 = GZAAS 23-0776, paratype), living culture, GZCC 23-0670; • ibid., 14 April 2023, J.Y. Zhang, ZY16 (HKAS 129859, paratype), living culture, KUNCC 23-14156; • ibid., ZY18 (HKAS 147019, paratype), living culture, KUNCC 23-14159. Additional sequence. KUNCC 23-13865: ITS (PV862363); GZCC 23-0670: ITS (PV862364); KUNCC 23-14156: ITS (PV862365); KUNCC 23-14159: ITS (PV862366). Notes. Four new strains formed a phylogenetically distinct lineage within Muyocopronaceae and are described as a new genus, Cyatheomyces. Cyatheomyces synnematosus has a unique morphology, characterized by macronematous, synnematous conidiophores, which notably distinguishes it from other genera in this family, which typically have solitary, microor macronematous, mononematous conidiophores (Crous et al. 2018a; Hernández-Restrepo et al. 2019; Xu et al. 2024). The morphological characteristics are similar to species in Phaeoisaria and Rhamphoriopsis (Hyde et al. 2018, 2019; Yang et al. 2023). However, phylogenetic analysis showed that Phaeoisaria was placed in Pleurotheciaceae, Pleurotheciales, Sordariomycetes, and Rhamphoriopsis in Rhamphoriaceae, Rhamphoriales, Sordariomycetes (Hyde et al. 2024). Comparatively, Cyatheomyces is assigned to Muyocopronaceae (Muyocopronales, Dothideomycetes) based on evidence from morphology and phylogeny. Pseudopalawaniella J.Y. Zhang, K.D. Hyde & Y.Z. Lu, gen. nov. Index Fungorum: IF904133 Etymology. The genus name refers to the similarity to Pseudopalawania. Type species. Pseudopalawaniella woodwardiae J.Y. Zhang, K.D. Hyde & Y.Z. Lu. Description. Sexual morph: Ascomata superficial, solitary or scattered, sub-carbonaceous to carbonaceous, flattened or raised, with a poorly developed basal layer and an irregular margin, dark brown to black. Peridium composed of dark brown or black cells of textura angularis. Hamathecium cylindrical to filiform, branched, septate, pseudoparaphyses, hyaline. Asci 8-spored, bitunicate, fissitunicate, cylindric-clavate, apically rounded, straight or slightly curved, sessile or short pedicellate. Ascospores overlapping, irregularly arranged, ellipsoid to broadly fusiform, with rounded ends, straight or slightly curved, 1-septate, constricted at the septum, guttulate, hyaline. Asexual morph: Undetermined. Pseudopalawaniella woodwardiae J.Y. Zhang, Y.Z. Lu & K.D. Hyde, sp. nov. Index Fungorum: IF904134 Fig. 4 Etymology. The species epithet refers to this fungal host, “Woodwardia japonica.” 18 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Diagnosis. Differs from Pseudopalawania species in its shapes of ascomata (Raised vs. Flattened), asci (asci without distinct ocular chambers vs. asci with distinct ocular chambers) and ascospores (ellipsoid to broadly fusiform ascospores with obtuse ends vs. fusiform to inequilateral ascospores with pointed ends). Holotype. HKAS 129705 Description. Sexual morph: Ascomata superficial, solitary or scattered, raised, rarely flattened, sub-carbonaceous to carbonaceous, appearing as circular, flattened, covering the host, with a poorly developed basal layer and an irregular margin, dark brown to black spots, 139–189 µm × (73–)85–111.5 µm high (x – = 163 × 97 µm, n = 15). Ostioles central. Peridium comprising dark brown or black cells of textura angularis, 13–22.5 µm wide. Hamathecium cylindrical to filiform, septate, pseudoparaphyses, hyaline, 1–2(–2.6) µm wide. Asci 8-spored, bitunicate, fissitunicate, cylindric-clavate, straight or slightly curved, sessile or inconspicuous pedicellate, apically rounded, with an inconspicuous ocular chamber, hyaline, 46–53 × 9.5–12 µm (x – = 47.5 × 10.5 µm, n = 20). Ascospores overlapping, 2–3-seriate, ellipsoid to broadly fusiform, inequilateral, with obtuse ends, straight or slightly curved, 1-septate, 3.5–6.5 wide at septum, constricted at the septum, with a slightly small lower cell, hyaline or subhyaline, 14.6–19.7 × 4.7–6.5 µm (x – = 17 × 5 µm, n = 20). Asexual morph: Undetermined. Culture characteristics. Ascospores germinating on WA within 15 h at 26 °C. Colonies on PDA, circular with slight wavy margin, flat with protuberance and wrinkle in the center, veined, yellowish white with ashen in the center from above; beige with dark brown in the middle from below, and not producing pigmentation in cultures. Material examined. CHINA • Guizhou Province, Tongren City, Jiangkou County, (27°46'38"N, 108°45'22"E), on dead leaf axis of Woodwardia japonica (Blechnaceae) in a forest near the roadside, 21 May 2022, J.Y. Zhang, F31-3 (HKAS 129705, holotype; GZAAS 23-0673, isotype), ex-type living culture KUNCC 23-13877. Additional sequence. ITS (PV862367). Notes. Pseudopalawaniella resembles Pseudopalawania in superficial, sub-carbonaceous to carbonaceous ascomata covering the host, cylindric-clavate asci with an ocular chamber, and hyaline, 1-septate ascospores (Mapook et al. 2020b). However, they are phylogenetically distinct and also differ in the shapes of their ascomata (raised vs. flattened), asci (asci without distinct ocular chambers vs. asci with distinct ocular chambers), and ascospores (ellipsoid to broadly fusiform ascospores with obtuse ends vs. fusiform to inequilateral ascospores with pointed ends). Multigene phylogenetic analysis shows that Pseudopalawaniella woodwardiae forms an independent clade within Muyocopronaceae, sister to Cyatheomyces synnematosus, with good bootstrap support (100% ML/1.00 PP, Fig. 1). However, we were unable to compare the morphological characteristics of these two species, as Pseudopalawaniella woodwardiae presents a sexual morph in nature, whereas Cyatheomyces synnematosus exhibits an asexual morph. Attempts were made to culture the asexual morph in Pseudopalawaniella woodwardiae and the sexual morph in Cyatheomyces synnematosus, but these attempts failed. Therefore, there is no morphological evidence to prove that the two species belong to the same genus. Furthermore, a comparison of nucleotide base pairs of LSU, ITS, RPB2, and tef1-α between Pseudopalawaniella woodwardiae (HKAS 129705) and Cyatheomyces synnematosus (HKAS 129699) showed 27/833 bp (3.2%, including 3 gaps), 97/839 bp (11.6%, including 42 gaps), 95/1079 bp (8.8%, without gaps), and 45/991 bp 19 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Figure 4. Pseudopalawaniella woodwardiae (HKAS 129705, holotype). A The host; B, C Colonies; D, E Sections through ascomata; F Peridium mixed with host substrate at the base of an ascoma; G, H Asci and pseudoparaphyses; I–L Ascospores; M Germinated ascospores; N Pure culture from above and below. Scale bars: 50 μm (D, E); 20 μm (F–H); 10 μm (I–M). (4.5%, without gaps) differences. Hence, to avoid taxonomic confusion, we introduce Pseudopalawaniella as a new genus in Muyocopronaceae (Muyocopronales). Further morphological investigations, together with more collections and molecular data, are needed to clarify the status of these two genera. 20 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Pleosporales Luttr. ex M.E. Barr, Prodr. Cl. Loculoasc. (Amherst): 67 (1987). Lentimurisporaceae N.G. Liu, Jian K. Liu & K.D. Hyde, Cryptog. Mycol. 39(2): 270 (2018). Notes. Lentimurisporaceae was introduced as a new pleosporalean family by Liu et al. (2018) to accommodate Bahusandhika, Lentimurispora, and two Berkleasmium species based on morphology, phylogeny, and divergence time estimates. Members of Lentimurisporaceae are dematiaceous hyphomycetes, which are characterized by punctiform colonies, sporodochial conidiophores, blastic conidiogenous cells, muriform or fusiform, cylindrical or rhomboidal conidia (Liu et al. 2018a; Hongsanan et al. 2020a). In this study, we introduce two new genera (Neoberkleasmium and Synnematospora) and emend the concept of Lentimurisporaceae to include synnematous conidiophores. Neoberkleasmium J.Y. Zhang, Y.Z. Lu & K.D. Hyde, gen. nov. Index Fungorum: IF904137 Etymology. The genus name refers to the similar genus Berkleasmium. Type species. Neoberkleasmium nigroapicale (Bussaban, Lumyong, P. Lumyong, McKenzie & K.D. Hyde) J.Y. Zhang, K.D. Hyde & Y.Z. Lu. Description. Sexual morph: Undetermined. Asexual morph: hyphomycetous. Colonies on natural substrate superficial, effuse, scattered or aggregated, sporodochial, pulvinata, punctiform, black. Mycelium partly immersed, partly superficial, composed of branched, septate, subhyaline to brown hyphae. Conidiophores macronematous, septate, hyaline to pale brown. Conidiogenous cells blastic, integrated, terminal, hyaline. Conidia acrogenous, solitary, cylindrical to broadly clavate, thick-walled, muriform, constricted at septa, brown. Notes. Neoberkleasmium is introduced as a segregated genus from Berkleasmium to accommodate Berkleasmium micronesiacum and B. nigroapicale. Berkleasmium species have been associated with helicosporous fungi and were placed in Tubeufiaceae, Tubeufiales (Lu et al. 2017; Tibpromma et al. 2017). In comparison, Neoberkleasmium is a member of Lentimurisporaceae, Pleosporales. Morphologically, Neoberkleasmium fits well with the concept of Lentimurisporaceae and resembles Lentimurispora in having sporodochial, brown to black conidiomata, monoblastic conidiogenous cells, and muriform, dematiaceous conidia (Table 2). However, Neoberkleasmium is distinguished Table 2. Morphological comparison of four accepted genera in Lentimurisporaceae. Genera Conidiophores Conidiogenous cells Conidia Reference(s) Bahusandhika Sporodochial, micronematous Spherical, ovoid, or ampulliform Fusiform, cylindrical, or rhomboidal; 1–3-septate; catenate; brown Pratibha et al. (2014); Crous et al. (2015); Crane and Miller (2016) Lentimurispora Sporodochial, micronematous Inverted vase-like, clavate Lenticular; muriform, dark brown central cells and subhyaline to pale brown peripheral cells Liu et al. (2018a) Neoberkleasmium Sporodochial, macronematous Cylindrical Cylindrical to broadly clavate; muriform; brown Matsushima (1981); Bussaban et al. (2001); Liu et al. (2018a) Synnematospora Synnematous, macronematous Ampulliform Cylindrical to oblong; (1–)3-septate; catenate; brown This study 21 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes from Lentimurispora by its cylindrical to broadly clavate conidia, often darkened at the upper part, whereas Lentimurispora produces conidia that are lenticular with dark brown central cells and pale-colored peripheral cells. Neoberkleasmium micronesiacum (Matsush.) J.Y. Zhang, K.D. Hyde & Y.Z. Lu, comb. nov. Index Fungorum: IF904138 ≡ Berkleasmium micronesiacum Matsush., Mycol. Mem. 2: 2 (1981) Holotype. USA, Guam, Mangilao, on the dead petiole of Cocoris nuciferae, 19 September 1980, Dried culture CMA, MFC-10321. Description. see Matsushima (1981). Notes. Berkleasmium micronesiacum was introduced by Matsushima (1981) based on morphological characteristics alone. However, phylogenetic analysis showed that B. micronesiacum and B. nigroapicale are separate from the monophyletic Berkleasmium lineage (Pinnoi et al. 2007; Liu et al. 2018a; Lu et al. 2018). Therefore, we transfer Berkleasmium micronesiacum and B. nigroapicale to Neoberkleasmium. Neoberkleasmium nigroapicale (Bussaban, Lumyong, P. Lumyong, McKenzie & K.D. Hyde) J.Y. Zhang, K.D. Hyde & Y.Z. Lu, comb. nov. Index Fungorum: IF904139 ≡ Berkleasmium nigroapicale Bussaban, Lumyong, P. Lumyong, McKenzie & K.D. Hyde, Fungal Diversity 8: 80 (2001) Holotype. THAILAND • Chiang Mai, Doi Suthep-Pui National Park, on dead pseudostems of Amomum siamense (Zingiberaceae), 15 October 2000, B. Bussaban CMUZS2 (POD 74415, holotype), ex-types living culture, BCC 8220 and HKUCC 7909. Description. see Bussaban et al. (2001). Notes. Berkleasmium nigroapicale was introduced by Bussaban et al. (2001), and this species was transferred to Neoberkleasmium based on phylogenetic evidence (Fig. 1). Synnematospora J.Y. Zhang, Y.Z. Lu & K.D. Hyde, gen. nov. Index Fungorum: IF904135 Etymology. The genus name refers to the synnematous conidiophores. Type species. Synnematospora pronephrii J.Y. Zhang, Y.Z. Lu & K.D. Hyde. Description. Sexual morph: undetermined. Asexual morph: hyphomycetous. Colonies on natural substrate effuse, scattered, dark brown, with conidial masses at the upper part of conidiophores. Mycelium mostly immersed, composed of septate, hyaline to brown hyphae. Synnemata composed of compactly adpressed conidiophores, brown to dark brown. Conidiophores macronematous, 22 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes synnematous, cylindrical, thick-walled, septate, brown. Conidiogenous cells blastic, discrete, terminal and lateral, ampulliform, flask-shaped, light brown. Conidia phragmosporous, solitary or catenate, cylindrical to oblong, straight or slightly curved, thick-walled, septate, constricted at septa, guttulate, brown. Conidial secession rhexolytic. Synnematospora pronephrii J.Y. Zhang, Y.Z. Lu & K.D. Hyde, sp. nov. Index Fungorum: IF904136 Fig. 5 Etymology. The species epithet refers to the fungal host, “Pronephrium penangianum.” Diagnosis. Differs from torula-like species in having macronematous, synnematous conidiophores. Holotype. HAKS 129756 Description. Sexual morph: undetermined. Asexual morph: hyphomycetous. Colonies on natural substrate effuse, scattered, dark brown, with conidial masses at the upper half of conidiophores. Mycelium mostly immersed, composed of septate, smooth, hyaline to brown hyphae. Synnemata composed of compact, parallel, adpressed conidiophores, brown to dark brown, 328–564 μm long and up to 35 μm wide at the base. Conidiophores macronematous, synnematous, cylindrical, unbranched, thick-walled, septate, pale brown to brown, 1.8–2.7 µm wide. Conidiogenous cells, blastic, discrete, ampulliform, flask-shaped, light brown, 2.5–5.3 × 2.8–4.2 µm (x – = 4 × 3.4 µm, n = 20). Conidia phragmosporous, solitary or catenate, cylindrical to oblong with rounded or truncated ends, straight, thick-walled, (1–)3-septate when mature, constricted at septa, always with a single guttule in each cell, brown, 9–15 × 3.7–5 µm (x – = 12.5 × 4.4 µm, n = 25). Conidial secession rhexolytic. Culture characteristics. Conidia germinating on WA within 15 h and germ tube produced from the ends of conidia. Colonies growing on PDA, reaching ca. 36 mm diameter in 20 days at 26 °C, circular, with entire margin, flat, with raise in the central part, dry, gray in the central part, brownness to pale brown towards the margin in front; dark brown in the center, paler to light brown towards the edge from below, and not producing pigmentation in cultures. Material examined. CHINA• Guizhou Province, Qianxinan Buyi and Miao Autonomous Prefecture, Anlong County, Xianheping National Forest Park, on dead stems of Pronephrium penangianum (Thelypteridaceae), 16 March 2022, J.Y. Zhang, J248 (HAKS 129756, holotype; GZAAS 23-0697, isotype), ex-type living culture KUNCC 23-13965. Additional sequence. ITS (PV862368). Notes. In the phylogenetic analysis, Synnematospora pronephrii formed a distinct and strongly supported lineage (81% ML/0.97 PP, Fig. 1), which is sister to Bahusandhika indica (GUFCC 18001) within Lentimurisporaceae. Members of Bahusandhika are torula-like (Pratibha et al. 2014; Crane and Miller 2016; Pem et al. 2024). Although the conidial characteristics of Synnematospora pronephrii resemble those of Bahusandhika species (Table 2), the species features macronematous, synnematous conidiophores. These characteristics also clearly distinguish it from other genera 23 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Figure 5. Synnematospora pronephrii (HAKS 129756, holotype). A–C Colonies on the host substrate; D–H Conidiophores; I–L Conidiogenous cells; M–P Conidia; Q Geminated conidium; R, S Pure culture from above and below. Scale bars: 100 μm (D–F); 20 μm (G, L); 50 μm (H); 5 μm (I–K); 10 μm (M–Q). 24 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes in Lentimurisporaceae, which possess sporodochial conidiomata (Liu et al. 2018a). Hence, we establish a new genus to accommodate Synnematospora pronephrii based on the morpho-phylogenetic evidence. Xenoberkleasmiaceae J.Y. Zhang, Y.Z. Lu & K.D. Hyde, fam. nov. Index Fungorum: IF904140 Etymology. The family name refers to the type genus. Type genus. Xenoberkleasmium N.G. Liu, Jian K. Liu & K.D. Hyde. Description. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies on natural substratum effuse, scattered, punctiform or powdery, dark-brown to black, glistening. Mycelium partly immersed, partly superficial, composed of branched, septate, hyaline to pale brown hyphae. Conidiomata sporodochial. Conidiophores micronematous to macronematous, mononematous, sometime reduced to conidiogenous cells. Conidiogenous cells blastic, integrated, terminal, determinate. Conidia acrogenous, broadly ellipsoidal to obovoid, thick-walled, muriform, brown to olivaceous green, with or without guttules, usually with basal cell attached. Notes. Xenoberkleasmiaceae is introduced to accommodate the genus Xenoberkleasmium, which is characterized by sporodochial conidiomata and muriform, brown conidia. This group forms a distinct and well-supported clade within Pleosporales and shares a close relationship with the morphologically unrelated family Hypsostromataceae. Xenoberkleasmium N.G. Liu, Jian K. Liu & K.D. Hyde, Fungal Diversity 129:1– 281 (2024) Type species. Xenoberkleasmium chiangraiense N.G. Liu, Jian K. Liu & K.D. Hyde 2024. Notes. Xenoberkleasmium was introduced as a monotypic genus to accommodate X. chiangraiense in Pleosporales (Liu et al. 2024a). In this study, X. chiangraiense clusters with Berkleasmium crunisia, B. pandani, and B. typhae in Pleosporales, distinct from the monophyletic lineage of Berkleasmium. Consequently, these three Berkleasmium species are transferred to Xenoberkleasmium based on shared morphological characteristics and congeneric phylogenetic placement. Xenoberkleasmium crinisium (Pinnoi) J.Y. Zhang, Y.Z. Lu & K.D. Hyde, comb. nov. Index Fungorum: IF904141 ≡ Berkleasmium crunisia Pinnoi, in Pinnoi, Jeewon, Sakayaroj, Hyde & Jones, Mycologia 99(3): 379 (2007) Holotype. THAILAND• Satun: Khuan Ka Long, on decaying rachis Calamus sp., 10 December 2004, A. Pinnoi in BIOTEC Bangkok Herbarium (BBH13084, holotype), ex-type culture, BCC 17023, 17024. 25 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Description. see Pinnoi et al. (2007). Notes. Berkleasmium crunisia was introduced by Pinnoi et al. (2007). In our phylogenetic analysis, Berkleasmium crunisia (BCC 17023) formed a distinct clade nested within Xenoberkleasmium. Morphologically, this species is characterized by sporodochial, punctiform colonies; macronematous, mononematous, clavate, aseptate, hyaline conidiophores; holoblastic conidiogenous cells; and muriform, oval to ellipsoidal, pale brown conidia (Pinnoi et al. 2007). These characteristics align with the generic concept of Xenoberkleasmium. Based on the molecular data and morphological characters, we therefore recognize Berkleasmium crunisia as a member of Xenoberkleasmium, proposing a new combination, X. crinisium. Xenoberkleasmium pandani (McKenzie) J.Y. Zhang, Y.Z. Lu & K.D. Hyde, comb. nov. Index Fungorum: IF904142 Fig. 6 ≡ Berkleasmium pandani McKenzie, Mycotaxon 104: 24 (2008) Holotype. MALAYSIA • Genting Highlands, Ganung Buah, in the dead leaves of Pandanus species (Pandanaceae), 18 August 1992, E.H.C. McKenzie (PDD 60532). Description. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies on natural substrate effuse, superficial, scattered to gregarious, sporodochial, velvety, punctiform, brown to black. Mycelium mostly superficial, composed of branched, septate, hyaline to brown hyphae. Conidiophores macronematous, mononematous, cylindrical, hyaline to middle brown, up to 32 µm. Conidiogenous cells monoblastic, holoblastic, terminal, inverted lageniform, clavate, narrowed towards the base, truncate at apex after conidial secession, with dense cytoplasm, hyaline, 13–33 × (4–)8.5–12.8 µm (x – = 23 × 10.5 µm, n = 20). Conidia 25–35 × 15–21 µm (x – = 30 × 18 µm, n = 30), acrogenous, solitary, oval to ellipsoidal, usually broadly obtuse at apex and slightly truncate at base, thick-walled muriform, not constricted or slightly constricted at the septa, sometimes with a dark median septum, brown. Culture characteristics. Conidia germinating on WA within 15 h and germ tube produced from the ends of conidia. Colonies growing on PDA, reaching about. 42 mm diameter in one month at 26 °C, circular, with an indentation at the entire margin, flat, with a protuberance in the center, dry, celadon to gray from central part to margin from above; pale brown to celadon to brown from center towards the margin from below, and not producing pigmentation in cultures. Material examined. CHINA • Guizhou Province, Tongren City, Jiangkou County, (27°46'38"N, 108°45'22"E), on dead stems of Woodwardia japonica (Blechnaceae) in a forest near the roadside, 21 May 2022, J.Y. Zhang, F31-1 (HKAS 147017 = GZAAS 23-0824), living culture KUNCC 23-13876; • ibid., F32-1 (HKAS 129706 = GZAAS 23-0674), living culture, KUNCC 23-13878; CHINA • Guizhou Province, Qiandongnan Miao and Dong Autonomous Prefecture, Liping County, Yongcong Town, on dead fronds of Blechnopsis orientalis (Blechnaceae) in a forest, 27 March 2022, J.Y. Zhang, J352 (HKAS 147020), living culture KUNCC 23-14012. 32 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes to dark brown, sometime slightly darked at the upper part, 46.5–66.5 × 22– 32.5 µm (x – = 53.5 × 26.3 µm, n = 30), often with a cylindrical to subglobose, hyaline or subhyaline to pale brown basal cell, 8.5–14 × 6–15 µm. Culture characteristics. Conidia germinating on WA within 15 h and germ tube produced from the base of conidia. Colonies growing on PDA under natural light, reaching ca. 36 mm diameter after one month at 26 °C, circular with entire margin, flat with a protuberance in the center, dry, velvety, white to taupe brown in concentric circles from center towards margin in the front; pale brown in the center, followed by brown to pale brown in concentric circles towards the edge, not producing pigmentation in culture. Material examined. CHINA • Guizhou Province, Tongren City, Jiangkou County, 27°46'38"N, 108°45'22"E 522M, on dead petiole of Woodwardia japonica, in a distributed forest, 21 May 2022, J.Y. Zhang, F37 (HKAS 129707, holotype; GZAAS 23-0675, isotype), ex-type living culture KUNCC 23-13881; • ibid., F34 (HKAS 147018 = GZAAS 23-0825, paratype), living culture, KUNCC 23-13880; • ibid., F38-1 (HKAS 129708, paratype), living culture, KUNCC 23-13882. Additional sequence. KUNCC 23-13881: ITS (PV862373); KUNCC 23-13880: ITS (PV862374); KUNCC 23-13882: ITS (PV862375). Notes. Three collections representing a new species, Xenopleopunctum guizhouense, form an independent lineage that is sister to X. sporodochiale with strong support value (100% ML/1.00 PP, Fig. 1). Xenopleopunctum guizhouense and X. sporodochiale share a similar morphology in having sporodochial, punctiform colonies, integrated, monoblastic, brown conidiogenous cells, and muriform, brown conidia with a hyaline or subhyaline to pale brown basal cell. However, Xenopleopunctum guizhouense differs from X. sporodochiale in having larger conidia (46.5–66.5 × 22–32.5 µm vs. 35–45(–47.8) × 18.3–25.5 µm) with diverse shapes (cylindrical to oval to irregular ellipsoidal vs. oval to ellipsoidal). Additionally, a comparison of nucleotide base pairs of ITS, LSU, RPB2, SSU, and tef1-α between X. guizhouense (F37-KUNCC 23-13881) and X. sporodochiale (C22-HKAS 129694) shows 27/469 bp (6%, including 4 gaps), 12/857 bp (1%, with 2 gaps), 60/1062 bp (6%, 0 gaps), 2/1022 bp (-%, 0 gaps), and 50/908 bp (6%, with 1 gap) differences, confirming they are distinct species. Xenopleopunctum sporodochiale J.Y. Zhang, Y.Z. Lu & K.D. Hyde, sp. nov. Index Fungorum: IF904149 Fig. 9 Etymology. The species epithet refers to its sporodochial conidiomata. Diagnosis. Differs from Xenopleopunctum guizhouense in having small conidia (35–45(–47.8) × 18.3–25.5 µm vs. 46.5–66.5 × 22–32.5 µm). Holotype. HKAS 129694 Description. Sexual morph: Undetermined. Asexual morph: Hyphomycetous. Colonies on natural substrate effuse, superficial, scattered to gregarious, sporodochial, punctiform, brown to black. Mycelium partly superficial, partly immersed, composed of branched, septate, hyaline to brown hyphae. Conidiophores micronematous, mononematous, reduced to conidiogenous cells. Conidiogenous cells monoblastic, integrated, terminal, cylindrical, thick-walled, brown, 3–6.2 × 2.5–3.1 µm (x – = 4.6 × 2.8 µm, n = 15). Conidia acrogenous, oval to ellipsoidal, 33 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Figure 9. Xenopleopunctum sporodochiale (HKAS 129694, holotype). A, B Colonies on the host substrate; C, D Mycelium; E–I Conidia; J, K Pure culture from above and below. Scale bars: 20 μm (C–I). muriform, slightly constricted at septa, hyaline or subhyaline when immature, hyaline to brown from base to upper part during maturation, brown when mature, darked at the septum, 35–45(–47.8) × 18.3–25.5 µm (x – = 40.7 × 21.5 µm, n = 30), often with a cylindrical to subglobose, hyaline or subhyaline to pale brown basal cell, 7.5–12 × 7.3–12 µm (x – = 9.8 × 10.9 µm, n = 20). 34 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Culture characteristics. Conidia germinating on WA within 13 h and germ tube produced from the base of conidia. Colonies growing on PDA under natural light, reaching ca. 26 mm diameter after one month at 26 °C, irregular or subrotund in shape with lobate edge, raised with convex in the center, dry, fluff, white to pale brown towards the edge from the front; middle brown to dark brown to white from the center to margin from below, and not producing pigmentation in culture. Material examined. CHINA • Guizhou Province, Zunyi City, Chishui County, Hushi Town, Chishui Alsophila Natural Reserve, on dead frond stalks of Cyatheaceae sp., 22 September 2019, J.Y. Zhang, C22 (HKAS 129694, holotype, GZAAS 23-0774, isotype), living culture, GZCC 23-0742. Additional sequence. ITS (PV862376). Notes. Xenopleopunctum sporodochiale formed a distinct clade and shared a sister relationship with X. guizhouense, which confirmed they are separate species. The difference between these two species was mentioned above. Discussion China is presently the second richest source of fungi associated with ferns and their allies (Bao et al. 2025; Zhang et al. 2025). In this study conducted in Guizhou Province, China, we isolated 14 strains from ferns identified in Dothideomycetes. Our phylogenetic analysis based on LSU, RPB2, SSU, and tef1-α sequence data supports a natural classification, leading to scientific adjustments in some lineages (Phukhamsakda et al. 2022). We propose one new family, six new genera, six new species, and five new combinations. This discovery further supports the previous study by Zhang et al. (2025) regarding the diversity and prevalence of Dothideomycetes within fungal communities associated with fern hosts. These findings enhance our understanding of fern-related fungal diversity in China and lay the groundwork for future studies on the ecological roles of fungi linked to ferns and their allies. Berkleasmium (Zobel 1854) has traditionally been considered polyphyletic and phylogenetically unresolved, as its broadly delimited concept accommodates sporodochial species that cluster in different families (Pinnoi et al. 2007; Hyde et al. 2016; Lu et al. 2017; Tibpromma et al. 2017). Tanney and Miller (2017) provided the asexual–sexual morph connection for B. concinnum and placed this generic type of B. concinnum in Tubeufiaceae, Tubeufiales, based on phylogenetic analysis. Subsequently, Lu et al. (2018) accepted only seven species in Berkleasmium sensu stricto based on strong phylogenetic evidence and similar sexual morph characteristics (Tanney and Miller 2017; Lu et al. 2018). Berkleasmium now has two asexual morph types of helicosporous and dictyosporous hyphomycetes, represented by B. concinnum. In this study, three Berkleasmium species (B. crunisia, B. pandani, and B. typhae) are transferred to Xenoberkleasmium, which shares similar morphological characteristics with the type species X. chiangraiense in having punctiform colonies; macronematous, mononematous conidiophores; monoblastic, cylindrical to inverted lageniform, hyaline or pale brown conidiogenous cells; and muriform, oval to ellipsoidal, black conidia with oblique and irregular septa (Liu et al. 2024a). The sporodochial dictyoconidia of Berkleasmium are characterized by cylindrical, brown conidiogenous cells and conidia with regular horizontal septa, which can 35 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes be differentiated from Xenoberkleasmium (Hughes 1958; Bussaban et al. 2001; Tanney and Miller 2017; Verma et al. 2019). Similarly, two species of Berkleasmium (B. micronesiacum and B. nigroapicale) form a phylogenetically monophyletic clade within the morphologically related family Lentimurisporaceae, with sporodochial conidiomata (Liu et al. 2018a). Thus, a new genus, Neoberkleasmium, is proposed to accommodate two new combinations, N. micronesiacum and N. nigroapicale. Furthermore, as many Berkleasmium species lack molecular data, they were not accepted by Lu et al. (2018), rendering their generic position dubious and necessitating clarification with new collections and isolates (Bussaban et al. 2001; Somrithipol and Jones 2003; Zhao and Zhang 2004; Qu et al. 2014; Tibpromma et al. 2017). The new genus Xenopleopunctum morphologically resembles Pleopunctum (Phaeoseptaceae), exhibiting an asexual morphology characterized by punctiform, brown colonies; monoblastic conidiogenous cells; and muriform, oval to ellipsoidal, dematiaceous conidia with a hyaline basal cell (Liu et al. 2019; Boonmee et al. 2021; Xu et al. 2023b; Yang et al. 2023). Phylogenetically, Xenopleopunctum shares a sister relationship with Pseudomassarina, which was placed within Pseudomassarinaceae. Pseudomassarina clematidis is the only accepted species in this family and exhibits similar sexual morphological traits to Lignosphaeria, a genus belonging to Phaeoseptaceae (Thambugala et al. 2015; Pem et al. 2024). Both genera possess immersed to erumpent, coriaceous ascomata with a central ostiole; cylindrical to clavate asci with an apically rounded and ocular chamber; and hyaline, septate, fusiform conidia (Thambugala et al. 2015; Phukhamsakda et al. 2020; Pem et al. 2024). Given these similarities, it is possible that our new genus, Xenopleopunctum, may represent an asexual morph within Pseudomassarinaceae, potentially related to Pseudomassarina. While we were unable to culture the sexual morph of Xenopleopunctum, this prevents us from verifying our hypothesis. Therefore, more collections and analyses are necessary to confirm any taxonomic assumptions and establish a connection to the sexual morph of Xenopleopunctum. Acknowledgments The authors would like to thank Shaun Pennycook (Manaaki Whenua Landcare Research, New Zealand) for advising on the fungal names. Jing-Yi Zhang thanks Ning-Guo Liu for his guidance. Kevin D. Hyde extends his appreciation to the Ongoing Research Funding Program (ORF-2025-114), 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. Use of AI No use of AI was reported. 36 IMA Fungus 16: e167717 (2025), DOI: 10.3897/imafungus.16.167717 Jing-Yi Zhang et al.: Discoveries of Dothideomycetes (Fungi) associated with pteridophytes Adherence to national and international regulations All the fungal strains used in this study have been legally obtained, respecting the Convention on Biological Diversity (Rio Convention). Funding This work was funded by the National Natural Science Foundation of China (NSFC 32060013), the Guizhou Institute of Technology High-Level Talent Research Start-up Project (2023GCC069), and the Science and Technology Foundation of Guizhou Province (Qian Ke He Pingtai ZSYS[2025]029). Author contributions Writing original draft: Jing-Yi Zhang. Review and editing: Kevin D. Hyde, Ming-Fei Yang, Ya-Ru Sun, Xing-Juan Xiao, Ze-Bin Meng, Dan-Feng Bao, Yong-Zhong Lu. Formal analysis: Jing-Yi Zhang, Ya-Ru Sun. Supervision: Yong-Zhong Lu. Author ORCIDs Jing-Yi Zhang https://orcid.org/0000-0003-0606-6169 Kevin D. 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