Pleurophragmium parvisporum (Ascomycota): One name, seven stories – a case highlighting the need for verification of strains from public culture collections

1 Pleurophragmium parvisporum (Ascomycota): One name, seven stories – a case highlighting the need for verification of strains from public culture collections Martina Réblová1, Jana Nekvindová2, Lucie Bauchová1, Margarita Hernández-Restrepo3 1 CzechAcademyofSciences,InstituteofBotany,DepartmentofTaxonomy,25243Průhonice,CzechRepublic 2 InstituteofClinicalBiochemistryandDiagnostics,UniversityHospitalHradecKrálové,50005HradecKrálové,CzechRepublic 3 WesterdijkFungalBiodiversityInstitute,3584CTUtrecht,Netherlands Correspondingauthor:MartinaRéblová([email protected]) Copyright: © Martina Réblová 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 Public repositories of living fungal strains provide essential reference points and support diverse scientific outcomes. Current best practices for preserving fungal strains emphasise the generation of DNA barcodes and the management of comprehensive metadata. However, challenges arise when type material or authentic reference strains are lacking, as this prevents direct comparison of DNA barcodes and forces identifications to rely solely on morphology. This problem is particularly pronounced for strains deposited during the pre-molecular era, especially those belonging to species with simple or convergent morphologies. In this study, we re-examined seven strains deposited in a public culture collection under the name Pleurophragmium parvisporum, including synonymous designations. Our approach combined cultivation experiments, comparative morphological analyses, multi-locus phylogenetic reconstruction of six nuclear markers, and biogeographic assessments. Our analyses revealed that these strains are scattered across four distinct families or orders in three classes. Two strains belong to Thysanorea (Chaetothyriales, Eurotiomycetes): T. acropleurogena sp. nov. and a sterile strain identified as the already known T. melanica. Two other strains were resolved within Wongia (Papulosaceae incertae sedis, Sordariomycetes) and introduced as W. pallidopolaris sp. nov. and W. rhachidophora sp. nov. Finally, two strains represent novel taxa within the Tubeufiales (Dothideomycetes), described here as Zaanenomyces hilifer sp. nov. and Skoliomycella flava gen. et sp. nov. Of the seven examined strains, only one conformed to the species concept of P. parvisporum and is here regarded as its reference strain. The phylogenetic analyses resolved P. parvisporum within Neomyrmecridium (Myrmecridiales, Sordariomycetes). Consequently, Neomyrmecridium was reduced to synonymy of Pleurophragmium, leading to the proposal of 11 new combinations (P. asiaticum comb. nov., P. asymmetricum comb. nov., P. fusiforme comb. nov., P. gaoligongense comb. nov., P. guizhouense comb. nov., P. luguense comb. nov., P. naviculare comb. nov., P. pteridophytophilum comb. nov., P. septatum comb. nov., P. sichuanense comb. nov., and P. sorbicola comb. nov.), and two new names (P. fluviale nom. nov. and P. jiulongheense nom. nov.). In addition, three species formerly placed in Uncispora are transferred to Thysanorea, with new combinations proposed based on congruent morphology and multi-locus phylogenetic evidence: T. hainanensis comb. nov., T. sinensis comb. nov., and T. wuzhishanensis comb. nov. This study refines the generic limits of Pleurophragmium and morphologically similar genera and reveals several previously unrecognised lineages. It highlights how misinterpretation of subtle morphological features may lead to strains being misidentified and deposited under incorrect names in public collections, where they risk perpetuating taxonomic errors. Academic editor: Sajeewa Maharachchikumbura Received: 24 September 2025 Accepted: 13 November 2025 Published: 25 November 2025 Citation: Réblová M, Nekvindová J, Bauchová L, Hernández-Restrepo M (2025) Pleurophragmium parvisporum (Ascomycota): One name, seven stories – a case highlighting the need for verification of strains from public culture collections. IMA Fungus 16: e173033. https://doi.org/10.3897/ imafungus.16.173033 IMA Fungus 16: e173033 (2025) DOI: 10.3897/imafungus.16.173033 2 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Key words: Ascomycota, Dactylaria, DNA barcode, holoblastic-denticulate conidiogenesis, new taxa, phylogenetics, repository, saprobic, systematics Introduction Public culture collections play a critical role in mycological research by serving as sources of reference material for taxonomy, evolutionary studies, ecology, biotechnology and pathology. They provide essential reference points that offer reliable information that is critical for data-driven research. As Hawksworth (2004) emphasised, such collections are more accurately termed genetic resource collections, reflecting their function as repositories of heritable biological diversity. Fungi are one of the most species-rich yet understudied groups of organisms on Earth. Recent estimates place global fungal diversity between 1.5 to 3.2 million species, with a working mean of approximately 2.53 million (Niskanen et al. 2023). In contrast, only about 204 040 species have been scientifically documented, representing roughly 8% of the estimated mean (Index Fungorum, accessed on 10 September 2025). The discrepancy between known and estimated diversity is even more pronounced in ex situ conservation efforts, such as culture collections, cryobanks, and germplasm storages. Some public collections are skewed toward a limited number of economically or medically important taxa, while vast lineages, especially of non-lichenised, microfungal lineages, remain poorly represented or absent. Lofgren and Stajich (2021) highlighted the need to address the ‘invisible’ fungal biodiversity crisis and positioned culture collections as key tools in fungal conservation and systematics. The World Data Centre for Microorganisms (gcm.wdcm.org, accessed on 10 September 2025) currently lists 588 984 strains of microorganisms, including fungi, yeasts, and bacteria, representing 57 219 species preserved across 158 culture collections worldwide. The proportion of fungal species maintained in culture collections is widely recognised to be low. Earlier estimates suggest that fewer than 20 000 fungal species are preserved ex situ (Hawksworth 2004). Among existing repositories, the Westerdijk Fungal Biodiversity Institute in Utrecht, the Netherlands, maintains the most comprehensive and globally recognised collection over 130 000 strains of microorganisms, of which more than 100 000 are fungi, including yeasts, corresponding to approximately 25 000 species. This extensive archive serves as the principal repository for nomenclatural types and authentic reference strains. It provides essential material for taxonomic, ecological, and biotechnological research. The effectiveness and reliability of the culture collections depend heavily on accurate species identification and comprehensive metadata. These collections are commonly used under the assumption that deposited strains are correctly identified, especially when they are assigned to known species. Before molecular tools became available, strain identification relied mainly on morphology. Verification practices varied between institutions, and many names were accepted without re-examination. With the increasing accessibility of DNA sequencing, however, routine generation of DNA barcodes for newly deposited strains has become feasible and is now being implemented in various 3 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum collections (Vu et al. 2016, 2019). In parallel, advances in data management systems (Reimer and Yurkov 2022) are improving the reliability and traceability of reference materials. Moreover, comprehensive monographic studies are being conducted to achieve better species resolution, particularly in genera of medical, industrial, or agricultural importance, such as Acremonium, Alternaria, Aspergillus, Ceratocystis, Fusarium, Penicillium, Phoma, and others (de Gruyter et al. 2013; Woudenberg et al. 2013, 2015; de Beer et al. 2014; Chen et al. 2017; Houbraken et al. 2020; Sandoval-Denis et al. 2025). However, re-identification of historical strains in public collections, especially saprobic fungi, is still not routinely performed. Some strains deposited before the molecular era remain in collections under unverified names, and these may conceal considerable phylogenetic diversity, as demonstrated in the present study. Misidentifications, especially of morphologically defined isolates, can have cascading effects in systematics, phylogenetics, biotechnology, and ecology. In response, the mycological community advocates for standardization, molecular verification, and proper voucher deposition as essential pillars for maintaining the integrity and reproducibility of fungal science (Hawksworth 2004; Aime et al. 2021; Lofgren and Stajich 2021). Nevertheless, even the implementation of routine DNA-based authentication may prove insufficient if reference or ex-type strains of the target species are unavailable, which is often the case. Under these circumstances, the traditional approach of carefully validating the morphology of obtained strains, guided by the principle of ‘trust, but verify’, remains essential for ensuring the reliability and success of any taxonomic or systematic research. In this study, we present specific cases encountered during our own research, which illustrate how unverified identifications of strains in public collections can obscure true phylogenetic relationships and hinder taxonomic clarity. A notable example is provided by the studies of Shenoy et al. (2006, 2010), where reliance on misidentified living strains, without morphological verification by the authors, triggered a cascade of erroneous conclusions, ultimately affecting the outcomes of several subsequent systematic investigations. Using partial nuclear large subunit (LSU) rDNA sequences, Shenoy et al. (2010) proposed that the genus Spadicoides was polyphyletic. The genus, typified by S. bina, was established by Hughes (1958) to accommodate saprobic, primarily lignicolous, dematiaceous hyphomycetes. To support their hypothesis of polyphyly, Shenoy et al. (2010) analysed four available strains: Spadicoides atra CBS 489.77, S. bina CBS 113708, S. verrucosa ex-type CBS 128.86, and S. xylogena CBS 310.31. Although S. atra (GenBank accession: EF204506) and S. bina (GenBank accession: EF204507) are morphologically similar, phylogenetic analysis did not support their congeneric placement. The strain of S. atra formed a sister relationship to three species of Lentomitella (Réblová et al. 2018), whereas S. bina was placed within the Porosphaerella clade (Müller and Samuels 1982). Hernández-Restrepo et al. (2017) accepted the conclusions of Shenoy et al. (2010), recognising Spadicoides as a member of the Cordanaceae, where Porosphaerella and its asexual morph Cordana belong, and segregating S. atra from Spadicoides into a newly established genus, Xenospadicoides. In addition, a new order, Xenospadicoidales, was introduced to accommodate Lentomitella and other spadicoides-like taxa. Using freshly collected material, Réblová et al. (2018) obtained S. bina CBS 137794 in axenic culture by isolating ascospores of 4 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum an unidentified lentomitella-like species. Their multi-locus phylogenetic analyses confirmed that S. bina, S. atra, and other Spadicoides species are congeneric. Based on this evidence, the genus Spadicoides was accepted as a member of the Xenospadicoidales, and Xenospadicoides was reduced to its synonymy. However, the identity of the strain S. bina CBS 113708, previously placed within the Cordanaceae by Shenoy et al. (2010), remained unresolved. Spadicoides bina and Cordana pauciseptata (Preuss 1851) are morphologically strikingly similar, both produce 1-septate, brown, ellipsoidal, acropleurogenous conidia of comparable size. However, they differ fundamentally in their mode of conidiogenesis. In Spadicoides, conidiogenous cells are tretic, whereas in Cordana, conidia are borne on denticles on holoblastic conidiogenous cells from terminal and intercalary (nodal) swellings. Detailed examination and comparison of the original voucher specimen housed in the Uppsala herbarium, the original dried culture and the derived living strain CBS 113708, both deposited in the CBS culture collection, confirmed that the fungus in question is, in fact, C. pauciseptata (Réblová et al. 2018). It is evident that the strain was initially misidentified, yet the deposition of all related material in public collections ultimately enabled accurate identification. Another example concerns the misidentification of a strain of Bahusutrabeeja dwaya (MTCC 9680), the type species of the genus, by Shenoy et al. (2010). It was placed within the Botryosphaeriales (Dothideomycetes). The authors were unable to account for this placement morphologically and discussed the apparent disparity between the phylogenetic and morphological data. Based on the ex-type strain, the species is currently assigned to Codinaea within the Chaetosphaeriales (Réblová et al. 2021). A similar case involves the misidentification of a non-type strain of Linkosia multiseptatum (HKUCC 10825), reported by Shenoy et al. (2006) as a member of the Rhamphoriales. More recent molecular data from the ex-type strain of L. multiseptatum (CGMCC 3.20786; Wu and Diao 2022) have placed Linkosia within the Chaetosphaeriales, together with other morphologically similar sporidesmium-like fungi. This study is focused on the case of Pleurophragmium parvisporum and re-evaluates the strains deposited under this name to clarify the species’ identity and assess the broader implications of strain misidentification in public collections. Pleurophragmium (Costantin 1888), typified by P. bicolor (= P. parvisporum, Holubová-Jechová 1972), was introduced for dematiaceous hyphomycetes. It is characterised by transversely septate, hyaline conidia that become subhyaline or pale brown at maturity, borne terminally and laterally on denticles on holoblastic conidiogenous cells, arising from simple, macronematous conidiophores. De Hoog and von Arx (1974) transferred Pleurophragmium to Dactylaria (Saccardo 1880) based on overlapping conidial morphology, and de Hoog (1985) later accommodated it as Dactylaria sect. Pleurophragmium. The concept of this species was primarily derived from observations of two strains, CBS 531.73 and CBS 770.83, which were used to describe both species-level traits and colony morphology (de Hoog 1985). Pleurophragmium parvisporum is represented by seven strains in the CBS culture collection. The only available molecular data for P. parvisporum consist of a partial LSU sequence from the non-type strain CBS 531.73 (GenBank accession: EU107296; Bhilabutra et al., unpublished), which was generated without prior morphological verification. In a phylogenetic analysis 5 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum by Réblová (2009), the placement of Rhodoveronaea and morphologically similar genera with holoblastic-denticulate conidiogenesis was examined using homologous LSU sequences representing species from 21 orders and families across three fungal classes. Pleurophragmium parvisporum CBS 531.73 grouped with Papulosa amerospora AFTOL-ID 748 (Spatafora et al. 2006) in the Sordariomycetes. Based on these results, Réblová (2009) excluded Pleurophragmium from Dactylaria and tentatively re-evaluated its placement within the Papulosaceae. However, the author noted that this decision was made in the absence of verified original material. In addition to Dactylaria, Pleurophragmium also shows striking morphological similarity to Neomyrmecridium (Crous et al. 2018a). Neomyrmecridium, based on N. septatum, currently encompassing 13 species, is characterised by dematiaceous, macronematous, erect, unbranched conidiophores, holoblastic-denticulate conidiogenous cells, and solitary, fusoid-ellipsoidal, obovoid or naviculate, transversely septate conidia that are initially hyaline and become pale brown with age, and have a mucoid sheath. Nevertheless, the close morphological similarity between the two genera has largely been overlooked in studies proposing new Neomyrmecridium species (e.g. Crous et al. 2018a, b; Serano et al. 2020; Xu et al. 2023; Zhang et al. 2024, 2025; Cao et al. 2025). During a revision of the genus Pleurophragmium, we aimed to clarify the systematic position of P. parvisporum, a morphologically distinctive yet poorly understood taxon. Although it has been reported from temperate regions of Asia (Matsushima 1975), North America (Wang 2010), and Europe (Ellis 1968, 1971; Holubová-Jechová 1972), we were unable to recollect it in nature. Moreover, our recent examination of the strain CBS 531.73 raised suspicion that it may be misidentified. In response, we acquired seven strains of this species available in the CBS culture collection. These isolates were identified based on morphological characteristics at the time of their deposition. Although ITS barcodes are now available for these strains, the absence of ex-type or verified strain of P. parvisporum makes accurate identification impossible without manual validation. In this study, we present a comprehensive investigation of P. parvisporum, with particular emphasis on clarifying the identity of seven strains obtained from the CBS culture collection. Our approach combined cultivation experiments, comparative morphological analyses, multi-locus phylogenetic reconstruction using six nuclear markers and biogeographic assessments. The results revealed unexpected phylogenetic diversity among these strains, which were shown to represent novel and previously known species, distributed across several genera, spanning three fungal classes. Materials and methods Isolates and morphological studies Living strains were sourced from the Westerdijk Fungal Biodiversity Institute (CBS) at Utrecht, the Netherlands. Strains CBS 122759, CBS 440.70, and CBS 862.68 were originally deposited as Pleurophragmium simplex (a synonym of P. parvisporum) and later reassigned to Dactylaria parvispora or Dactylaria sp. Strain CBS 531.73 was deposited as P. parvisporum and subsequently placed under D. parvispora, while strains CBS 215.96, CBS 770.83, and CBS 113561 were 6 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum deposited directly as D. parvispora. Dried cultures were deposited in the CBS Fungarium, while additional herbarium specimens of P. parvisporum were deposited in the Herbarium of the Institute of Botany (PRA), Czech Academy of Sciences at Průhonice, Czech Republic. Biogeographic assessments were conducted using environmental ITS datasets retrieved from the GlobalFungi database (Větrovský et al. 2020). Newly described taxa were registered in MycoBank (Crous et al. 2004). Details of the strains examined, including their origin and GenBank accession numbers of the generated sequences, are summarised in Table 1. Colony macromorphology was examined using an Olympus SZX12 dissecting microscope (Olympus America, Inc., Melville, NY, USA). Microscopic preparations were mounted in 90% lactic acid, water, or Melzer’s reagent, with measurements taken from specimens in Melzer’s reagent. Conidial dimensions are presented as mean ± standard deviation (SD) based on 20–25 measurements. Micromorphological traits were studied with an Olympus BX51 light microscope, and micrographs were captured using an Olympus DP75 camera operated via Olympus cellSens Dimension software v. 4.3. Colony images were taken with a Canon EOS 77D digital camera equipped with a Canon EF 100 mm f/2.8L Macro IS USM lens (Canon Europe Ltd., Middlesex, UK), illuminated with 5500K 16W LED lights. All images were processed in Adobe Photoshop CS6 (Adobe Systems, San Jose, CA, USA). To examine colony morphology, pigment production, and growth rates, strains were cultivated on four media: cornmeal dextrose agar (CMD) (cornmeal agar, Oxoid Limited, Basingstoke, UK, supplemented with 2% w/v dextrose), malt extract agar (MEA) (Oxoid), Modified Leonian’s agar (MLA) (Malloch 1981), oatmeal agar (OA), and potato-carrot agar (PCA) (Crous et al. 2019). To promote sporulation, strains were also grown on cornmeal agar (CMA) (Crous et al. 2019) supplemented with dried stems of Urtica dioica and incubated under alternating 12-h cycles of UV light and darkness. Colony features were recorded from 4-wk-old cultures incubated at 23 °C in darkness. Scientific names of fungal genera reported in this study are abbreviated as follows: Botryosphaeria (B.), Camporesiomyces (Ca.), Cancellidium (Cn.) Cordana (C.), Cyphellophora (Cy.), Helicoma (H.), Minimelanolocus (M.), Neomyrmecridium (N.), Pleurophragmium (P. ), Pseudospiropes (Ps.), Skoliomycella (S.), Thysanorea (T.), Tubeufia (Tu.), Uncispora (U.), Wongia (W.), and Zaanenomyces (Z.). Molecular methods Phylogenetic relationships were evaluated using six gene markers: internal transcribed spacer ITS1–5.8S–ITS2 (ITS) of the nuclear rDNA cistron (ITS barcode) (Schoch et al. 2012), nuclear large subunit (LSU) rDNA gene, the nuclear small subunit (SSU) rDNA gene, and three coding markers, i.e. the second largest subunit of RNA polymerase II (DNA-directed RNA polymerase) (rpb2), the intermediate section of the translation elongation factor 1-α (tef1), and β-tubulin (tub2) marked by exons 3−6. The LSU and SSU markers provide reliable phylogenetic resolution at the generic and higher taxonomic levels in fungi (e.g. Zhang et al. 2007; Schoch et al. 2009). The coding gene markers are recognised for their effectiveness in resolving interspecific relationships (Stielow et al. 2015; Meyer et al. 2019). 7 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Protocols for DNA extraction from 2–4-d old cultures and PCR amplification of cited gene markers were conducted following the methods described by Réblová et al. (2022) and Réblová and Nekvindová (2023). Automated sequencing was carried out by Eurofins Genomics Europe Sequencing Service (Cologne, Germany). Analyses of raw sequence data and assembly of sequence contigs were performed using Sequencher v. 5.4.6 (Gene Codes Corp., Ann Arbor, MI, USA). Phylogenetic analyses Preliminary similarity searches were performed using the BLASTn and megaBLAST algorithms with ITS and LSU sequences of all studied strains to identify their closest relatives. Corresponding ITS, LSU, SSU, rpb2, tef1, and tub2 sequences of related taxa were then retrieved from GenBank sequence database at NCBI (Sayers et al. 2022) and used in subsequent analyses. Sequences of these species, together with their GenBank accession numbers and references, are listed in Table 2. Sequence alignments were generated with MAFFT v. 7.487 (Katoh and Standley 2013) via the CIPRES Science Gateway v. 3.3 (Miller et al. 2010) and adjusted manually in BioEdit v. 7.1.8 (Hall 1999) when necessary. The best nucleotide substitution models for each partition (ITS, LSU, SSU, rpb2, tef1, tub2) were selected under the Akaike Information Criterion using MrModeltest v. 2.4 (Nylander 2004). Phylogenetic reconstructions were performed with Maximum Likelihood (ML) and Bayesian Inference (BI) methods implemented in the CIPRES Science Gateway. Table 1. Species, isolate information and new sequences determined for this study (in bold) and additional sequences retrieved from GenBank. Organism Strain Status* Host Substrate Country GenBank accessions Reference ITS LSU SSU rpb2 tef1 tub2 Pleurophragmium simplex CBS 770.83 unidentified dead twigs Japan PX283736 PX283746 PX283743 PX310214 PX310206 –This study Skoliomycella flava CBS 122759 Tunidentified plant debris Portugal PX283737 PX283747 –PX310215 PX310207 –This study Thysanorea acropleurogena CBS 215.96 Tunidentified wood Papua New Guinea PX283738 PX283748 – – – PX310220 This study Thysanorea melanica CBS 862.68 n/a wheat field soil Netherlands PX283739 PX283749 – – – PX310221 This study Wongia pallidopolaris CBS 440.70 Tn/a sandy soil Netherlands PX283740 PX283750 PX283744 PX310216 PX310208 –This study Wongia rhachidophora CBS 531.73 TBambusa sp. dead leaf India PX283741 PX283751 PX283745 PX310217 PX310209 –This study Zaanenomyces hilifer CBS 113561 TCarex sp. litter Iran PX283742 PX283752 –PX310218 PX310210 –This study Zaanenomyces moderatricisacademiae CBS 148315 TJuncus inflexus dead culm Netherlands OK664723 OK663762 –OK651167 PX310211 –Crous et al. 2021 Zaanenomyces quadripartis CBS 148310 TJuncus effusus dead culm Netherlands OK664721 OK663760 –PX310219 PX310212 –Crous et al. 2021 Zaanenomyces versatilis CBS 148312 TJuncus inflexus dead culm Netherlands OK664730 OK663769 – – PX310213 –Crous et al. 2021 *T denotes ex-type culture. 8 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Table 2. Species, isolate information and sequences retrieved from GenBank. Taxon Strain Status GenBank Accession numbers Reference ITS LSU SSU rpb2 tef1 tub2 Acanthohelicospora pinicola MFLUCC 10-0116 T KF301526 KF301534 – – KF301555 – Boonmee et al. 2014 Acanthostigma chiangmaiense MFLUCC 10-0125 T JN865209 JN865197 – – KF301560 – Boonmee et al. 2011, 2014 Acanthostigma perpusillum UAMH 7237 AY916492 AY856892 – – – – Tsui et al. 2006 Aciculomyces restrictus FMR 18994 T ON009870 ON009950 – – – ON667802 Torres-Garcia et al. 2023 Aculeata aquatica MFLUCC 11-0529 T MG922571 MG922575 – – – – Dong et al. 2018 Atrokylindriopsis setulosa HMAS 245592 T KP337330 KP337329 – – – – Ma et al. 2015 Berkleasmium aquaticum MFLUCC 17-0049 T KY790444 KY790432 –MF535268 KY792608 – Lu et al. 2017c, 2018b Berkleasmium fusiforme MFLUCC 17-1978 T MH558693 MH558820 –MH551007 MH550884 – Lu et al. 2018a Berlesiella nigerrima MUCL 39954 AF050251 AF050251 – – – – Untereiner and Naveau 1999 Boerlagiomyces macrosporus MFLUCC 12-0388 KU144927 KU764712 – – KU872750 – Doilom et al. 2017 Botryosphaeria agaves MFLUCC 10-0051 JX646790 JX646807 – – JX646855 – Liu et al. 2012 Botryosphaeria dothidea CBS 115476 EKF766151 DQ678051 –DQ677944 DQ767637 – Schoch et al. 2006; Slippers et al. 2013 Botryosphaeria wangensis HGUP 190007 MZ541933 MZ540051 –OP321271 – – Zhang et al. 2021, 2022 Brunneosporella aquatica HKUCC 3708 AF177154 AF132326 – – – Ranghoo et al. 1999 Camporesiomyces bhatii GMBCC 1120 T PQ763360 PQ842543 –PV388888 PV388894 – Han et al. 2025 Ca. bhatii GMBCC 1125 PQ763361 PQ842544 –PV388889 PV388895 – Han et al. 2025 Camporesiomyces coffeae GMBCC 1130 T PQ763358 PQ842545 –PV388890 PV388896 – Han et al. 2025 Ca. coffeae GMBCC 1131 PQ763359 PQ842546 –PV388891 PV388897 – Han et al. 2025 Camporesiomyces mali KUMCC 19-0216 T MN792813 MN792811 – – MN794018 – Hyde et al. 2020 Camporesiomyces patagoniensis BBB MVB 573 T JN127358 JN127359 – – – – Sánchez et al. 2012 Camporesiomyces puerensis GMBCC 1113 T PQ763356 PQ842541 –PV388886 PV388892 – Han et al. 2025 Ca. puerensis GMBCC 1114 PQ763357 PQ842542 –PV388887 PV388893 – Han et al. 2025 Camporesiomyces vaccinii CBS 216.90 T AY916486 AY856879 – – – – Tsui et al. 2006 Cancellidium atrobrunneum MFLUCC 20-0100 T MT422724 MT422740 MT422726 –MT436438 – Hyde et al. 2021 Cancellidium cinereum MFLUCC 18-0424 T MT370353 MT370363 MT370351 MT370486 MT370488 – Hyde et al. 2021 Cancellidium griseonigrum MFLUCC 17-2117 T MT370354 MT370364 MT370352 MT370487 – – Hyde et al. 2021 Capronia camelliaeyunnanensis CGMCC 3.19061 T MH807377 MH807378 – – – – Phookamsak et al. 2019 Capronia kleinmondensis CBS 122671 T MH863226 MH874753 – – – – Vu et al. 2019 Capronia leucadendri CBS 122672 T MH863227 MH874754 – – – – Vu et al. 2019 Capronia lijiangensis CGMCC 3.20501 T OK487581 OK487580 – – – – Phukhamsakda et al. 2022 Capronia pilosella AFTOL-ID 657 DQ826737 DQ823099 – – – – James et al. 2006 Cladophialophora boppii CBS 126.86 T EU103997 NG_058762 – – – – Gueidan et al. 2008 Cladophialophora carrionii CBS 114393 KF928452 KF928516 – – – EU137151 de Hoog et al. 2007; Attili-Angelis et al. 2014 Cl. carrionii CBS 160.54 T AB109177 LC192080 – – – EU137201 Abliz et al. 2004; de Hoog et al. 2007; Kiyuna et al. 2018 Cladophialophora floridana NRRL 66282 T AB986343 AB986343 – – – – Obase et al. 2016 Cladophialophora chaetospira CBS 114747 EU035403 KF928514 – – – KF928578 Crous et al. 2007; AttiliAngelis et al. 2014 Cladophialophora matsushimae MFC-1P384 T FN549916 FN400758 – – – – Koukol 2010 Cladophialophora mycetomatis CBS 122637 T FJ385276 LC192076 – – – – Badali et al. 2008; Kiyuna et al. 2018 Cladophialophora yegresii CBS 114405 T EU137322 KX822323 – – – EU137209 de Hoog et al. 2007; Vasse et al. 2017 Cyphellophora aestiva aCBS 228.86 T KC455244 KC455257 – – – KC455227 Réblová et al. 2013 Cyphellophora laciniata CBS 190.61 T EU035416 FJ358239 – – – JQ766329 Crous et al. 2007; Gueidan et al. 2008; Feng et al. 2014 9 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Taxon Strain Status GenBank Accession numbers Reference ITS LSU SSU rpb2 tef1 tub2 Cyphellophora suttonii CBS 449.91 T KC455243 KC455256 – – – KC455226 Réblová et al. 2013 Dematiohelicomyces helicosporus MFLUCC 16-0213 T KX454169 KX454170 –MF535258 KY117035 – Hyde et al. 2016; Lu et al. 2017b, 2018b Dematiohelicosporum guttulatum MFLUCC 17-2011 T MH558705 MH558833 –MH551021 MH550896 – Lu et al. 2018a Dematiotubeufia chiangraiensis MFLUCC 10-0115 T JN865200 JN865188 – – KF301551 – Boonmee et al. 2011, 2014 Dictyospora thailandica MFLUCC 16-0001 T KY873627 KY873622 –MH551023 KY873286 – Brahamanage et al. 2017; Lu et al. 2018a Exophiala angulospora CBS 482.92 T JF747046 MH874033 – – – JN112426 de Hoog et al. 2011; Vu et al. 2019 Exophiala castellani CBS 158.58 T KF928458 MH869272 – – – KF928586 Attili-Angelis et al. 2014; Vu et al. 2019 Exophiala dermatitidis CBS 207.35 T AF050269 KF928508 – – – KF928572 Untereiner and Naveau 1999; Attili-Angelis et al. 2014 Exophiala equina FMR 18335 ON009852 ON009932 – – – ON491590 Torres-Garcia et al. 2023 Exophiala heteromorpha CBS 232.33 T AY857524 MH866871 – – – – Prenafeta-Boldú et al. 2006; Vu et al. 2019 Exophiala jeanselmei CBS 507.90 T MH862234 MH873915 – – – – Vu et al. 2019 Exophiala mesophila CBS 402.95 T JF747111 KX712349 – – – JN112476 de Hoog et al. 2011; Teixeira et al. 2017 Exophiala nigra CBS 535.94 T MH86248 KX712353 – – – – Teixeira et al. 2017; Vu et al. 2019 Exophiala nishimurae CBS 101538 T AY163560 KX712351 – – – JX482552 de Hoog et al. 2003; Woo et al. 2013; Teixeira et al. 2017 Exophiala pisciphila CBS 537.73 T NR_121269 MH872483 – – – JN112493 de Hoog et al. 2011; Schoch et al. 2014; Vu et al. 2019 Exophiala prototropha CBS 534.94 OR371992 – – – – – Unpublished Exophiala radicis FMR 18645 ON009857 ON009937 – – – ON667789 Torres-Garcia et al. 2023 Exophiala salmonis CBS 157.67 T MH858932 MH870616 – – – JN112499 de Hoog et al. 2011; Vu et al. 2019 Fluminicola aquatica MFLUCC 15-0962 T MF374357 MF374366 MF374374 –MF370960 – Zhang et al. 2017 Fluminicola saprophytica MFLUCC 15-0976 T NR_153493 MF374367 MF374375 MF370954 MF370956 – Zhang et al. 2017 Fluminicola striata MFLUCC 18-0990 T MW286496 MW287770 – – – – Dong et al. 2021 Fonsecaea monophora CBS 289.93 AY366925 – – – – EU938554 de Hoog et al. 2004; Najafzadeh et al. 2009 Fonsecaea nubica CBS 269.64 T EU938592 – – – – EU938574 Najafzadeh et al. 2009 Fonsecaea pedrosoi CBS 271.37 NAB114127 KJ930166 – – – EU938559 de Azevedo et al. 2015; Najafzadeh et al. 2009 Fonsecaea pugnacius CBS 139214 T KR706553 KR706549 – – – KR706547 de Azevedo et al. 2015 Helicangiospora lignicola MFLUCC 11-0378 T KF301523 KF301531 – – KF301552 – Boonmee et al. 2014 Helicoarctatus aquaticus MFLUCC 17-1996 T MH558707 MH558835 –MH551024 MH550898 – Lu et al. 2018a Helicodochium aquaticum MFLUCC 17-2016 T MH558709 MH558837 –MH551026 MH550900 – Lu et al. 2018a Helicohyalinum aquaticum MFLUCC 16-0014 MH558711 MH558839 –MH551028 MH550902 – Lu et al. 2018a Helicohyalinum infundibulum MFLUCC 16-1133 T MH558712 MH558840 –MH551029 MH550903 – Lu et al. 2018a Helicoma brunneisporum MFLUCC 17-1983 T MH558714 MH558842 –MH551031 MH550905 – Lu et al. 2018a Helicoma longisporum MFLUCC 17-1997 T MH558720 MH558846 –MH551037 MH550911 – Lu et al. 2018a Helicomyces hyalosporus MFLUCC 17-0051 T MH558731 MH558857 –MH551047 MH550922 – Lu et al. 2018a Helicomyces torquatus MFLUCC 16-0217 MH558732 MH558858 –MH551048 MH550923 – Lu et al. 2018a Helicosporium aquaticum MFLUCC 17-2008 T MH558733 MH558859 –MH551049 MH550924 – Lu et al. 2018a Helicosporium flavisporum MFLUCC 17-2020 T MH558734 MH558860 –MH551050 MH550925 – Lu et al. 2018a Helicosporium luteosporum MFLUCC 16-0226 T KY321324 KY321327 –MH551056 KY792601 – Lu et al. 2017a; 2018a Helicosporium setiferum MFLUCC 17-1994 T MH558735 MH558861 –MH551051 MH550926 – Lu et al. 2018a Helicosporium vegetum CBS 254.75 – DQ470982 –DQ470934 DQ471105 – Spatafora et al. 2006 Helicosporium vesicarium MFLUCC 17-1795 T MH558739 MH558864 –MH551055 MH550930 – Lu et al. 2018a Helicotubeufia guangxiensis MFLUCC 17-0040 T MH290018 MH290023 –MH290033 MH290028 – Liu et al. 2018b Helicotubeufia hydei MFLUCC 17-1980 T MH290021 MH290026 –MH290036 MH290031 – Liu et al. 2018b Chlamydotubeufia cylindrica MFLUCC 16-1130 T MH558702 MH558830 –MH551018 MH550893 – Lu et al. 2018a 16 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum CBS 449.91 T Cyphellophora suttonii CBS 190.61 T Cyphellophora laciniata CBS 228.86 T aCyphellophora aestiva 99/1 BSS 376 T171 Tm12 CBS 269.64 T Fonsecaeanubica CBS 139214 T Fonsecaeapugnacius CBS 289.93 Fonsecaea monophora CBS 271.37 T Fonsecaeapedrosoi 81/– MFC 1P384 T Cladophialophora matsushimae CBS 122637 T Cladophialophora mycetomatis CBS 126.86 T Cladophialophora boppii NRRL 66282 T Cladophialophora floridana CBS 114747 Cladophialophora chaetospira CBS 114405 T Cladophialophora yegresii CBS 160.54 T CBS 862.96 UAMH 10875 T Phialophora americana CBS 140325 E Phialophora verrucosa BMU 01890 T Phialophora chinensis 91/0.99 87/1 92/1 JAUCC M0840-1 T Neoveronaea sinensis FMR 18994 T Aciculomyces restrictus HMAS 245592 T Atrokylindriopsis setulosa 90/1 CBS 482.92 T Exophiala angulospora MFLUCC 120389 T Melanoctona tectonae MUCL 39954 Berlesiella nigerrima CGMCC 3.19061 T Capronia camelliae yunnanensis CGMCC 3.20501 T Capronia lijiangensis AFTOL ID-657 Capronia pilosella MFLUCC 16-1449 T Marinophialophora garethjonesii CBS 534.94 Exophiala prototropha CBS 232.33 T Exophiala heteromorpha CBS 207.35 T Exophiala dermatitidis 87/– 99/1 CBS 181.65 IN Rhinocladiellaanceps CBS 496.78 T Rhinocladiellaphaeophora CBS 313.73 T Rhinocladiellaaquaspersa MFLUCC 11-0529 T Aculeata aquatica CBS 101538 T Exophiala nishimurae CBS 507.90 T Exophiala jeanselmei CBS 535.94 T Exophiala nigra CBS 101597 Phaeoannellomyces elegans CBS 264.49 T Melanchlenus eumetabolus CBS 317.33 AS Rhinocladiella atrovirens CBS 122672 Capronia leucadendri CBS 122671 Capronia kleinmondensis 93/1 CBS 147585 T Neoherpotrichiella juglandicola CBS 537.73 T Exophiala pisciphila CBS 157.67 T Exophiala salmonis FMR 18335 Exophiala equina FMR 18645 Exophiala radicis 93/1CBS 572.90 T e CBS 254.57 T CBS 268.75 T Veronaea compacta CBS 776.83 T Veronaea japonica 88/1 99/1 76/0.95 CBS 158.58 Exophiala castellanii CBS 402.95 T Exophiala mesophila CBS 145909 T Thysanorea cantrelliae CBS 145910 T Thysanorea seifertii CBS 215.96 T Thysanorea acropleurogena sp. nov. YMF 1.03683 T Thysanorea sinensis comb. nov. YMF 1.04038 T Thysanorea hainanensis comb. nov. YMF 1.04080 T Thysanorea wuzhishanensis comb. nov. 99/1 MFLUCC 15-0416 Thysanorea obscura MFLUCC 17-2378 T Thysanorea nonramosa CBS 212.96 T MFLUCC 15-0966 T d MFLUCC 15-0971 T Thysanorea thailandensis CBS 862.68 MFLUCC 15-0415 T MFLUCC 15-0237 T Thysanorea asiatica MFLUCC 15-0259 T Thysanorea curvata CBS 126086 Thysanorea rousseliana KUMCC 15-0206 T Thysanorea submersa MFLUCC 15-0414 T Thysanorea yunnanensis 86/1 82/– 86/0.99 77/0.99 91/1 88/1 –/0.99 –/0.98 –/0.98 –/0.97 –/0.95 –/1 –/0.97 100/– –/1 –/0.95 0.1 Herpotrichiellaceae (Chaetothyriales) Thysanorea melanica Thysanorea papuana outgroup Cyphellophoraceae Thysanorea Veronaeabotryosa Cladophialophora carrionii Valentiella maceioensis Figure 2. Maximum likelihood phylogenetic tree of the Herpotrichiellaceae based on ITS–LSU–tub2 DNA sequences. Names in bold indicate taxonomic novelties; strains with their accession numbers in bold and highlighted in violet colour were sequenced in this study; a ex-type of Cyphellophora vermispora, d ex-type of Thysanorea aquatica, e ex-type of Veronaea constricta. T, E, IN and AS denote ex-type, ex-epitype, ex-isoneotype, and authentic strains, respectively. Thickened branches indicate support ML BS = 100% and PP values = 1.0. Branch support of nodes ≥ 70% ML and ≥ 0.95 PP is indicated above or below branches. A hyphen (–) indicates values lower than 75% ML BS or 0.95 PP. 17 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum (YMF1.03683), and U. wuzhishanensis (YMF1.04080), were nested within the Thysanorea clade, corroborating their transfer to Thysanorea. These species together formed a sister lineage (71/95) to T. acropleurogena. The systematic position of U. harroldiae, the type species of Uncispora, remains unknown. The third phylogenetic analysis (Fig. 3), based on the ITS–LSU–SSU–rpb2– tef1 dataset, included 27 ingroup strains representing 21 species in four genera of the Papulosaceae and related genera (Sordariomycetes incertae sedis). A total of 258 nucleotides were excluded from the 5′ and/or 3′ ends of LSU, SSU, rpb2, and tef1 datasets due to incomplete sequences in most strains. Of the 4 472 characters (including gaps), 1 335 were unique sites identified by RAxML: 328 in ITS, 190 in LSU, 87 in SSU, 481 in rpb2, and 249 in tef1. Three species of Cancellidium (Cancellidiales), i.e. Cn. atrobrunneum MFLUCC 20-0100, Cn. cinereum MFLUCC 18-0424, and Cn. griseo-nigrum MFLUCC 17-2117, were used as outgroup taxa. The best-fit models of nucleotide substitution selected were: SYM+I+G (ITS), GTR+I+G (LSU, rpb2, tef1), and GTR+I (SSU). Papulosaceae were recovered as a strongly supported lineage (100/1.0) compri sing Brunneosporella aquatica, Fluminicola (96/90), Papulosa amerospora and Wongia (100/1.0), together with the closely related genera Pseudostanjehughesia (100/1.0) and Platytrachelon. Both strains, CBS 440.70 and CBS 531.73, clustered within the well-supported Wongia subclade. The strain CBS 440.70 grouped with W. aquatica in a basal position and is here described as a new species, W. pallidopolaris. Strain CBS 531.73 was resolved as a new lineage, W. rhachidophora, which was nested within a strongly supported subclade (100/1.0). It includes morphologically similar species such as W. bambusae, W. bandungensis, W. fusiformis, and W. suae that likely represent a species complex. To assess the relationships of W. rhachidophora with these species, three loci commonly employed for species-level delimitation, ITS, tef1, and rpb2, were compared. Pairwise sequence identities indicated that W. rhachidophora is most closely related to W. bandungensis (ITS: 99.4%, tef1: 99.0%, rpb2: 99.4%) and W. suae (ITS: 98.4%, tef1: 98.7%, rpb2: 99.6%), while showing lower similarity to W. bambusae (ITS: 97.1–97.8%, tef1: 98.5%, rpb2: 98.7%), and W. fusiformis (ITS: 95.3%, tef1: 98.5%, rpb2: 96.9%). The ITS region shows moderate divergence (0.6–1.6%). The ITS sequences show that W. rhachidophora is closest to W. bandungensis (99.4%). However, it still shows small differences that may or may not exceed intraspecific variation thresholds. There is also high similarity among W. rhachidophora and W. suae/W. bambusae (97–98%), but below typical species threshold (≥98.5– 99%) (Vu et al. 2019; Lücking et al. 2020). In the tef1 dataset, all values hover around 98.5–99%, showing strong relatedness across species; W. rhachidophora is again most similar to W. bandungensis. The rpb2 sequence of W. rhachidophora is shorter 755 bp vs 900–1000 bp in others, which can inflate identity values slightly because less variable regions are often retained. Apart from W. fusiformis, all rpb2 identities are very high (≥98.7%). The highest sequence identity is to W. suae and to W. bandungensis. Together with morphological distinctiveness (see notes to W. rhachidophora), the molecular data support W. rhachidophora as a distinct species within the species complex. In addition, the sequence comparison of W. bandungensis and W. suae indicate that ITS identity (98.6%) is right at the species boundary in fungi. Their tef1 and rpb2, however, show nearly complete identity (≥99.9%), strongly indicating they are the same species. 18 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum MFLUCC 18-0424 T Cancellidium cinereum MFLUCC 20-0100 T Cancellidium atrobrunneum MFLUCC 17-2117 T Cancellidium griseonigrum CBS 125235 Platytrachelonabietis MFLUCC 16-0569 T Pseudostanjehughesia aquitropica MFLUCC 15-0352 T Pseudostanjehughesia lignicola AFTOL ID-748 Papulosaamerospora HKUCC 3708 Brunneosporellaaquatica MFLUCC 18-0990 T Fluminicola striata MFLUCC 15-0962 Fluminicola aquatica MFLUCC 15-0976 T Fluminicola saprophytica 99/1 96/– MFLUCC 18-1607 T Wongia aquatica CBS 440.70 T Wongia pallidopolarissp. nov. KUNCC 24-17692 T Wongia guttulata KUNCC 23-13715 T Wongia flava BCRC FU32062 T Wongia miscanthi BRIP 69019 T Wongia ficherai DAR 79637 T Wongia garrettii DAR 80512 T BRIP 60377 DLUCC 1767 KUNCC 23-16632 MFLUCC 21-0032 T MFLUCC 21-0028 82/–KUNCC 24-17699 CGMCC 3.24360 T CBS 531.73 T Wongia rhachidophora sp. nov. CGMCC 3.24295 T Wongia suae TBRC-BCC 95171 T TBRC-BCC 95343 I 91/1 98/0.98 99/1 98/1 94/1 81/0.97 96/1 97/1 76/– 0.05 100/1 outgroup Cancellidiales Papulosaceae Wongia Sordariomycetidaeincertaesedis Wongia bandungensis Wongia bambusae Wongia fusiformis Wongia griffinii Figure 3. Maximum likelihood phylogenetic tree of the Papulosaceae based on ITS–LSU–SSU–rpb2–tef1 DNA sequences. Names in bold indicate taxonomic novelties; strains with their accession numbers in bold and highlighted in violet colour were sequenced in this study. T and I denote ex-type and ex-isotype strains, respectively. Thickened branches indicate support ML BS = 100% and PP values = 1.0. Branch support of nodes ≥ 70% ML and ≥ 0.95 PP is indicated above or below branches. A hyphen (–) indicates values lower than 75% ML BS or 0.95 PP. The final phylogenetic analysis (Fig. 4), conducted on the ITS–LSU–rpb2– tef1 dataset, included 71 ingroup strains representing 65 species of the Tubeufiaceae (Tubeufiales, Dothideomycetes). Ninety-five nucleotides were excluded from the 5′ end of the LSU dataset due to incomplete sequences in most strains. Of the total of 3 512 characters (including gaps), 1 631 were unique sites identified by RAxML: 442 in ITS, 271 in LSU, 538 in rpb2, and 380 in tef1. Three members of Botryosphaeria (Botryosphaeriales), i.e. B. agaves MFLUCC 10-0051, B. wangensis HGUP190007, and B. dothidea CBS 115476, were selected as outgroup taxa. The best-fit models of nucleotide 19 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum CBS 115476 E Botryosphaeriadothidea HGUP 190007 Botryosphaeria wangensis MFLUCC 10-0051 Botryosphaeriaagaves MFLUCC 10-0124 E MFLUCC 16-0024 MFLUCC 10-0125 T Acanthostigma chiangmaiense UAMH 7237 Acanthostigma perpusillum CBS 122759 T Skoliomycella flavagen. et sp.nov. CBS 148315 T Zaanenomyces moderatricis-academiae CBS 149453 CBS 148312 T CBS 113561 T Zaanenomyces hilifer sp. nov. CBS 148310 T CBS 148272 GZAAS 19-1751 T Neodictyospora karsti MFLUCC 16-0226 T Helicosporium luteosporum MFLUCC 17-1994 T Helicosporium setiferum MFLUCC 17-2008 T Helicosporiumaquaticum CBS 254.75 Helicosporium vegetum MFLUCC 17-2020 T Helicosporium flavisporum MFLUCC 17-1795 T Helicosporium vesicarium 88/1 KUMCC 19-0216 T Camporesiomyces mali BBB MVB 573 Camporesiomyces patagoniensis CBS 216.90 T Camporesiomyces vaccinii GMBCC 1125 GMBCC 1120 T 90/1GMBCC 1114 GMBCC 1113 T GMBCC 1130 T GMBCC 1131 92/1 85/0.99 99/1 83/1 MFLUCC 16-1125 T Neotubeufia krabiensis MFLUCC 11-0378 T Helicangiospora lignicola MFLUCC 16-0001 T Dictyospora thailandica MFLUCC 17-2011 T Dematiohelicosporumguttulatum MFLUCC 10-0116 T Acanthohelicospora pinicola MFLUCC 11-0510 T Neoacanthostigma fusiforme MFLUCC 17-0049 T Berkleasmium aquaticum MFLUCC 17-1978 T Berkleasmium fusiforme MFLUCC 10-0115 T Dematiotubeufia chiangraiensis MFLUCC 11-0379 T Neohelicoma fagacearum MFLUCC 17-1997 T Helicoma longisporum MFLUCC 17-1983 T Helicoma brunneisporum CGMCC 3.23539 T Neomanoharachariellaaquatica MFLUCC 17-1996 T Helicoarctatus aquaticus MFLU 11-0228 T Kamalomyces bambusicola MFLUCC 13-0233 T Kamalomyces thailandicus 86/1 93/1 77/1 MFLUCC 17-1982 T Pleurohelicosporium parvisporum GZCC 20-0489 T Pleurohelicosporium hyalinum MFLUCC 16-1134 T Chlamydotubeufia krabiensis MFLUCC 16-1130 T Chlamydotubeufia cylindrica MFLUCC 16-0010 T Aquaphila albicans MFLUCC 17-1980 T Helicotubeufia hydei MFLUCC 17-0040 T Helicotubeufia guangxiensis MFLUCC 16-0016 T Neochlamydotubeufia fusiformis MFLUCC 10-0118 T Neochlamydotubeufia khunkornensis MFLUCC 16-0213 T Dematiohelicomyces helicosporus MFLUCC 17-2016 T Helicodochiumaquaticum MFLUCC 16-1133 T Helicohyalinuminfundibulum MFLUCC 16-0014 Helicohyalinumaquaticum MFLUCC 17-1523 T Neohelicosporiumparvisporum MFLUCC 17-1522 T Neohelicosporiumguangxiense MFLUCC 12-0170 T Manoharachariella tectonae MFLUCC 12-0388 Boerlagiomyces macrosporus KUMCC 15-0276 Muripulchraaquatica MFLUCC 16-0993 T Neohelicomyces aquaticus KUMCC 15-0470 T Neohelicomyces grandisporus MFLUCC 17-0051 T Helicomyces hyalosporus MFLUCC 16-0217 Helicomyces torquatus MFLUCC 17-2021 Parahelicomyces talbotii CGMCC 32-3535 Parahelicomyces hyalosporus MFLUCC 17-0053 T Tubeufia inaequalis MFLUCC 17-1524 T Tubeufia eccentrica GZCC 22-2010 T Pseudotubeufia hyalospora GZCC 22-2011 T Pseudotubeufia laxispora 94/1 86/1 83/1 99/1 81/– 92/1 0.1 –/0.98 –/0.98 –/0.99–/1 –/0.98 1x outgroup Botryosphaeriales Tubeufiaceae (Tubeufiales) Camporesiomyces coffeae Camporesiomyces puerensis Camporesiomyces bhatii Zaanenomyces quadripartis Zaanenomyces versatilis Thaxteriellopsis lignicola Figure 4. Maximum likelihood phylogenetic tree of the Tubeufiaceae based on ITS–LSU–rpb2–tef1 DNA sequences. Names in bold indicate taxonomic novelties; strains with their accession numbers in bold and highlighted in violet colour were sequenced in this study. T and E denote ex-type and ex-epitype strains, respectively. Thickened branches indicate support ML BS = 100% and PP values = 1.0. Branch support of nodes ≥ 70% ML and ≥ 0.95 PP is indicated above or below branches. A hyphen (–) indicates values lower than 75% ML BS or 0.95 PP. 20 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum substitution selected were: GTR+I+G (ITS, LSU, and tef1) and SYM+I+G (rpb2). Strains CBS 113561 and CBS 122759 were resolved as members of the Tubeufiaceae. The resulting topology recovered 37 well-supported generic lineages within the family. Strain CBS 113561 clustered within Zaanenomyces clade (83/1.0) closely related to Z. quadripartis, supporting its recognition as a new species, Z. hilifer. They formed a strongly supported sister lineage (100/1.0) to a subclade (100/1.0) comprising Z. moderatricisacademiae and Z. versatilis. In contrast, strain CBS 122759 formed a distinct lineage, justifying the establishment of a new monotypic genus, Skoliomycella, with S. flava as its type species. Both Skoliomycella and Zaanenomyces were placed within a robust subclade (92/1.0) together with Acanthostigma (100/1.0), Camporesiomyces (99/1.0), Helicosporium (88/1.0), and Neodictyospora karstii. Morphological studies Except for CBS 113561 (see below) and CBS 862.68 (which remained sterile), all other strains could be readily distinguished from P. parvisporum under culture conditions. Two strains represent two distinct species of Thysanorea (Arzanlou et al. 2007) in the Herpotrichiellaceae. Thysanorea acropleurogena (CBS 215.96) sporulated abundantly, producing simple or apically loosely branched conidiophores with terminal and intercalary conidiogenous cells forming an elongated denticulate rachis, and pale brown to pale olivaceous brown, predominantly 3-septate conidia. In contrast, T. melanica (CBS 862.68) (Liu et al. 2015) remained sterile on all tested media, and its identification relied exclusively on molecular data. Two other strains belong to the genus Wongia in the Papulosaceae, where they represent distinct species lineages, here described as W. pallidopolaris (CBS 440.70) and W. rhachidophora (CBS 531.73). Both species produced brown to dark brown conidia, borne on terminal conidiogenous cells with one to several denticles on relatively short, robust conidiophores. The two last strains, CBS 113561 and CBS 122759, are members of the Tubeufiaceae. Strain CBS 113561 produced hyaline, mostly 3-septate conidia, and simple conidiophores with a short denticulate rachis and sometimes bearing nodulose swellings. In culture, its morphology closely resembled that of P. parvisporum, making its distinction challenging. This strain is described here as the new species Zaanenomyces hilifer. In contrast, strain CBS 122759 is readily distinguished from P. parvisporum by flexuous to sinuous conidiophores with a pronounced zig-zag pattern, bearing solitary denticles scattered along the entire conidiophore axis, and by its 3–7-septate, subhyaline to pale olivaceous-brown conidia. It represents a novel lineage in the Tubeufiaceae and is introduced here as the new genus and species Skoliomycella flava. Biogeography assessment The biogeographic assessment revealed striking contrasts in distribution patterns among the studied taxa (Fig. 5). Some species, such as T. melanica, W. pallidopolaris, and Z. hilifer, are clearly cosmopolitan and ecologically 21 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum versatile, occurring across multiple continents, biomes, and climate zones. Their strong representation in soils, particularly croplands and grasslands, points to an ecological preference for terrestrial substrates and suggests that human activities, especially agriculture, may have played a role in their global dissemination. The high abundance of W. pallidopolaris and T. melanica in European croplands is particularly notable and supports the idea that these taxa thrive in disturbed and anthropogenic habitats. By contrast, other species appear much more restricted. Pleurophragmium parvisporum shows a concentration in East Asia with scattered European records, mainly from aquatic habitats, reflecting a narrower ecological niche. These findings provide a broader perspective and contrast with previously published studies, which have so far been based largely on European collections of this species (Costantin 1888; Ellis 1968; Holubová-Jechová 1972) and a few records from Japan (Matsushima 1975) and North America Figure 5. Geographical distribution of the studied species based on environmental DNA records from the GlobalFungi database. Each dot indicates a georeferenced sample in which the species was detected, with the number of samples (n) given in parentheses. 22 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum (Wang 2010). Skoliomycella flava is represented by a single environmental record in urban air from the Iberian Peninsula, while W. rhachidophora shows a disjunct distribution between India and subtropical North America, hinting at either true rarity or under-sampling. Finally, T. acropleurogena remains known only from its type locality, with no matches in environmental sequencing datasets. These patterns collectively illustrate how some lineages are widespread yet cryptic, while others are genuinely rare or overlooked, underscoring the importance of integrating eDNA surveys with traditional taxonomy to resolve their ecology and biogeography. Taxonomy Pleurophragmium Costantin, Les mucédinées simples: 100. 1888. Synonyms. Dactylaria sect. Pleurophragmium (Costantin) de Hoog, Stud. Mycol. 26: 36. 1985. Neomyrmecridium Crous, Persoonia 41: 287. 2018. Type species. Pleurophragmium parvisporum (Preuss) Hol.-Jech. Description. Sexual morph. Not observed. Asexual morph. Colonies effuse, hairy, brown to black, subhyaline, beige to pale brown when sporulating; vegetative hyphae immersed or semi-immersed. Conidiophores macronematous, mononematous, solitary or in small groups, erect, straight or slightly flexuous, unbranched or rarely branched, sometimes elongating percurrently, brown, septate. Conidiogenous cells integrated, terminal and/or intercalary resulting from the formation of a septum within the original cell during sympodial elongation, polyblastic, occasionally monoblastic, sympodially proliferating, bearing one to several denticles or forming a rachis with minute, pimple-like denticles or protrusions scattered over the surface; conidiogenesis holoblastic-denticulate. Conidia solitary, dry, acropleurogenous, obovoid or subclavate, fusiform, ellipsoidal, usually tapering towards the base, with a distinct hilum, hyaline, subhyaline or pale brown, sometimes with paler end cells, aseptate or transversely septate, wall smooth or finely ornamented, with an ephemeral mucoid sheath; conidial secession schizolytic. Notes. Morphological comparisons together with phylogenetic analyses of P. parvisporum CBS 770.83 support the view that genera Neomyrmecridium and Pleurophragmium are congeneric, leading to the proposal of 11 new combinations and the introduction of two new names. Given the substantial morphological variability among species currently placed in Pleurophragmium, the genus is unlikely to represent a single evolutionary lineage, rendering the traditional generic concept untenable. Accordingly, the present generic description is based on the morphological variability of species whose placement is supported by molecular data and refers to Pleurophragmium s. str. The following list of species is organised into three categories: (i) species accepted in Pleurophragmium s. str.; (ii) species of uncertain status that remain in Pleurophragmium s. lat.; and (iii) species excluded from Pleurophragmium and transferred to other genera. Names in bold indicate the currently accepted classification, accompanied by full synonymy and, where appropriate, brief explanatory notes. The interspecific variability of species listed in the first two categories is summarised in the synopsis table (Table 3). Key to species of Pleurophragmium was provided by D’Souza and Bhat (2012). 23 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Table 3. Synopsis table of species of Pleurophragmium. Species accepted in Pleurophragmium s. str. Taxon Substrate* Size (µm) Conidia End cells Ornamentation Conidiogenous cellsConidiophores Reference Septa Shape Sheath Colour Size (µm) Size (µm) Pleurophragmium asiaticum C(13–)15–16(–17) × (3.5–)4.5(–5) (2–)3 ellipsoid to obovoid, with truncate hilum at the base present pale brown smooth-walled 5–35 × 4–7 50–100 × 3–5 Crous et al. 2018a Pleurophragmium asymmetricum NS 12–15 × 2–3 1narrowly clavate or subclavate absent yellowish to subhyaline smooth-walled 8–35 × 3.5–5 40–210 × 3.5–9 Serrano et al. 2020 Pleurophragmium fluviale NS 14–16 × 4–6 3obovoid, rounded at the apex, pointed at the base absent subhyaline smooth-walled n/a 211–308 × 5–7 Luo et al. 2019 Pleurophragmium fusiforme NS 29–34 × 4–6 (0–)1–3 navicular to tapering, pointed at both ends absent pale brown paler smooth-walled n/a 214–285 × 4–5 Zhang et al. 2024 Pleurophragmium gaoligongense NS 16–24 × 5–7 0–3 clavate-cymbiform, truncate at the base absent subhyaline to pale brown smooth-walled 34–68 × 3–6 138–226 × 4–7 Xu et al. 2023 Pleurophragmium guizhouense NS 8.9–12.7 × 2.8–4.8 (2–)3 fusoid-ellipsoid, apex obtuse or tapering, with a subtruncate hilum at base absent subhyaline to pale brown smooth-walled 2.2–4.3 (width) 75–140 × 2–4.5 Hyde et al. 2020 Pleurophragmium jiulongheense NS 16–23 × 3.7–5.6 3clavate to fusiform, truncate at the base present subhyaline to pale brown smooth-walled n/a (144–)204–332 × 3.4–4.8 Cao et al. 2025 Pleurophragmium luguense NS 12–15 × 4–7 (0–)2–3 obovoid, tapering at the base absent subhyaline to pale brown smooth-walled 39–68 × 3–5 152–298 × 4–6 Xu et al. 2023 Pleurophragmium naviculare NS 16–24 × 5.5–7.5 (1–) 3 navicular to fusiform, tapering to a hilum towards the base, obtuse at the apex present hyaline, becoming pale brown paler smooth-walled n/a 100–200 × 4–5.6 Yang et al. 2023 Pleurophragmium parvisporum NS 10–18 × 3.5–6 (0–)3(–4) ellipsoid to subclavate, rounded at the apex, pointed at the base absent hyaline to very pale brown crumpled 26–47(–51) × 4.5–5.5 (80–)118–250(– 270) × (3.5–)4–5 This study Pleurophragmium pteridophytophilum NS 8.5–11 × 3–4 0–1 obovoid absent hyaline to pale brown rough-walled 20–55.5 × 2.5–4(– 4.5) 150–224 × 2.7–4.9 Zhang et al. 2025 Pleurophragmium septatum C(12–)14–16(–20) × (3.5–)4(–5) (1–) 3 fusoid-ellipsoid, apex obtuse, tapering in lower third to a truncate hilum present hyaline, becoming pale brown smooth-walled 30–40 × 4–5 40–70 × 4–5 Crous et al. 2018a Pleurophragmium sichuanense NS 9.5–14.5 × 4–4.5 0–1 fusiform or narrowly obovoid absent subhyaline to pale brown finely verrucose n/a 93–130 × 4–5 Chen et al. 2025 Pleurophragmium sorbicola C(7–)8–10(–15) × 4(–5) (0–)1(–3) obovoid, obtuse at the apex present hyaline, pale brown with age smooth-walled 20–65 × 3–4 50–200 × 4–7 Crous et al. 2018b Pleurophragmium s. lat.: species of uncertain status Pleurophragmium angamosense C24.5–40 × 5–8 3–7 mostly 5 fusiform, curved or straight absent pale brown paler smooth-walled 3–3.5 (width) 200 × 3.5–4.5 Matsushima 1995 Pleurophragmium aquaticum NS 25–30 × 6–6.5 3fusiform to clavate, sometimes navicular, truncated at the base absent brown, paler around septa paler smooth-walled n/a 200–390 × 5–11 Heredia et al. 2007 Pleurophragmium bitunicatum C20–35 × 4.5–7 3(–5) fusiform, narrowly truncated at the base absent brown smooth-walled n/a (50–) 100–220 × 3–5 Matsushima 1975 Pleurophragmium clavatum NS 9.5–16.5 × 3.5–4.5 0clavate to fusiform, narrower at the ends, tapered to a point at the base absent yellowish brown verrucose n/a 75–160 × 3.3–4 Ma et al. 2014 Pleurophragmium ellipsoideum NS 10–17 × 6–8.5 3ellipsoid to obovoid, rounded at the apex, pointed at the base absent pale brown smooth-walled n/a 150–260 × 3.5–6 Ma et al. 2014 24 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Species accepted in Pleurophragmium s. str. Taxon Substrate* Size (µm) Conidia End cells Ornamentation Conidiogenous cellsConidiophores Reference Septa Shape Sheath Colour Size (µm) Size (µm) Pleurophragmium harunganae NS 12–18 × 5–7 0fusoid, curved absent subhyaline to pale olivaceous smooth-walled n/a 150 × 4–5 Hansford 1946 Pleurophragmium indicum NS 20–30 × 4.5–11 3ellipsoidal to obovoid, rounded at the apex, pointed at the base, straight to slightly curved absent dark brown paler smooth-walled 15–23 × 4.5–9.5 100–160 × 7–22 D‘Souza and Bhat 2012 Pleurophragmium malaysianum C(20–) 40–75 × 4–5 3–10 pseudoseptate cylindrical to clavate, rounded at the apex, protruding at the base absent hyaline smooth-walled n/a 25–50 (–100) × 3.5–5 Matsushima 1996 Pleurophragmium miniumbonatum NS 16–19 × 6–7 (2–) 3 obovoid, pyriform to broadly clavate, sometimes slightly clavate, umbonate at the apex, truncate at the base absent dark brown to brown subhyaline basal cell smooth-walled 55–95 × 3–3.5 90–150 × 5.5–7 Castañeda 1999 Pleurophragmium naviculiforme C18–32 × 6.5–10 1navicular, pointy at the apex, pointed at the base absent hyaline smooth-walled n/a 70–140 × 6–8 Matsushima 1975 Pleurophragmium obcampanuloides C(10–)11.5–15.5(– 18) × 5–6.5(–7) (1–)2 obpyriform to turbinate absent pale olivaceous, olivaceous in mass smooth-walled n/a 100–150 × 3.8–5 Matsushima 1995 Pleurophragmium peruamazonicum var. peruamazonicum C10–30 × 4–6 2cylindrical, rounded at the apex, pointed at the base absent pale brown, fuscous in mass subhyaline smooth-walled n/a 100–300 × 3–4 Matsushima 1993 Pleurophragmium peruamazonicum var. inflatum C13–20 × (5–)6– 9(–10) (1–)2 ovoid, rounded at the apex, narrow at the base absent pale sooty subhyaline apical cell smooth-walled n/a (50–) 200–600 × 4–5.5 Matsushima 1993 Pleurophragmium subfusiforme C21–38 × 5.8–9.2 3–7 fusiform, straight to sometimes curved absent subhyaline, pale fuscous in mass smooth-walled n/a 200–500 × 4–5 Matsushima 1975 Pleurophragmium taiwanense C12–20 × 3–4.5 3cylindro-clavate, with a slightly inflated apical cell absent pale brown hyaline smooth-walled n/a 70–200 (–250) × 3.5–4.5 Matsushima 1987 Pleurophragmium tricolor NS 17–18 × 4–5 2ellipsoidal with, rounded at the apex, pointed at the base absent brown, light brown basal cell hyaline apical cell smooth-walled n/a 196–200 × 5–7 Rambelli 2009 Pleurophragmium varieseptatum NS 19–22 × 4–5 1–4 cylindrical, rounded at the apex, pointed at the base absent pale olivaceous, olivaceous-grey in mass smooth n/a 70–125 × 3.5–5 Matsushima 1975 Pleurophragmium verruculosum C10–16.5 × 3.3–5 1–3(–3) subclavate, clavate, pointed at the base absent subhyaline to pale brown verrucose n/a 8.2–26.4 (–15) × 1.6–4.5 (–3) Tiwari et al. 1969 Pleurophragmium yunnanense NS 10–15 × 6–7 (2–)3 broadly fusiform, narrowed at the ends, tapered to a point at the base absent pale brown smooth n/a 185–270 × 3.5–4.5 Ma et al. 2014 * natural substrate (NS); culture (C). 25 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Aquapteridospora (Distoseptisporales, Sordariomycetes) (Yang et al. 2015), is a dematiaceous hyphomycete genus that warrants comparison with Pleurophragmium. Both genera share rigid, erect, darkly pigmented, simple conidiophores on the natural substrate that bear terminal and frequently intercalary holoblastic-denticulate conidiogenous cells that proliferate sympodially, as well as septate conidia that may be surrounded by a mucoid sheath. However, they differ primarily in conidial pigmentation. In Pleurophragmium s. str., conidia are typically hyaline to subhyaline, becoming pale brown at maturity or when in mass, occasionally showing paler end cells, seldom with a mucoid sheath, and have thin-walled septa. Pleurophragmium fusiforme is an exception in having pale brown conidia with paler ends. However, in Aquapteridospora conidia are primarily pigmented, continuously brown or dark brown, often with paler end cells, with thick-walled septa and are often embedded in a mucoid sheath. The exception is A. hyalina described with hyaline conidia that become subhyaline to pale brown at maturity (Ma et al. 2022). Based on molecular evidence, Pleurophragmium bambusinum (Dai et al. 2016) has been transferred to Aquapteridospora by Bao et al. (2021). In addition, several other Pleurophragmium species of uncertain status (see below) and which lack molecular data, show close morphological similarity with members of this genus. Pleurotheciella (Réblová et al. 2012) is another genus warranting comparison with Pleurophragmium. Despite their close morphological resemblance in asexual characteristics, which makes them difficult to distinguish, the two genera are clearly separated phylogenetically. Pleurotheciella belongs to the Pleurotheciales, where it forms a robust and species-rich lineage. Species accepted in Pleurophragmium s. str. Here we list 14 species currently accepted in Pleurophragmium s. str. whose placement is supported by molecular data. These species are morphologically very similar, with conidial characters providing the primary basis for distinction. Several species of Pleurophragmium are known only from culture, with no confirmed wild type (the typical form of a species as it occurs in nature). This reliance on in vitro observations may complicate identification and lead to distortion of key traits, i.e. distribution of fertile regions on the conidiophore (apically or in nodulose swellings), conidiophore appearance, conidial dimensions and septation patterns, and even the presence of a mucoid sheath. In culture, the morphology of conidiophores and conidia may deviate from that observed in the wild type (Réblová et al. 2021, 2022). In addition, the conidial size of species currently referred to Pleurophragmium s. str. show considerable overlap, making it difficult to distinguish species based solely on conidial dimensions. Pleurophragmium asiaticum (Crous) Réblová & Hern.-Restr., comb. nov. MycoBank No: 860799 Basionym. Neomyrmecridium asiaticum Crous, Persoonia 41: 291. 2018. Typus. THAILAND • Ratchaburi Province; on leaves of unidentified vine; 2008; P. W. Crous HPC 2252 (holotype CBS H-23774, culture ex-type CPC 34535 = CBS 145080). 32 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum 8588 (PRA24034); • Olomouc Region, Hrubý Jeseník Mts., U Kříže forest between Bělá pod Pradědem and Vidly; on decaying wood of Fagus sylvatica; 8 Aug 1971; V. Holubová-Jechová 8584 (PRA-24035); • Ibid.; Bučina virgin forest above Františkova myslivna cabin near Kouty nad Desnou; on decaying wood of a branch of Fagus sylvatica; 4 Aug 1971; V. Holubová-Jechová 8586 (PRA-24036). JAPAN • Kyoto, Daitokuji Tempel D; on unidentified dead twigs; 28 Aug 1983; W. Gams & M. Tsuda (CBS H-3522, dried culture CBS H-3506, living culture CBS 770.83). SLOVAK REPUBLIC • Lúčanská Malá Fatra Mts., Šrámková National Nature Reserve, Kýčery; on decaying wood of Fagus sylvatica; 26 Sep 1983; V. Holubová-Jechová 8556 (PRA-24037). Habitat and geographical distribution. Saprobe on decaying wood and herbaceous stems of various hosts, such as Arctium lappa, Brassica oleracea, Campanula medium, Carpinus betulus, Conium maculatum, Epilobium hirsutum, Fagus sylvatica, Filipendula ulmaria, Heracleum sphondylium, Lithocarpus edulis, Polygonum sieboldii, Quercus sp., Sambucus ebulus, Urtica dioica in the Belgium, Czech Republic, Denmark, France, Germany, Japan, Mexico, Netherlands, New Zealand, Slovak Republic, UK and New Jersey and Washington, USA (Preuss 1852; Costantin 1888; Berkeley and Broome 1861; Ellis 1968; Holubová-Jechová 1972; Matsushima 1975; Wang 2010; GBIF Secretariat 2023). According to GlobalFungi database, P. parvisporum was detected in 40 samples. Most records originate from Asia, primarily from Japan and China, with a smaller contribution from South Korea. Additional records come from Europe, all from Spain. The species is most frequently detected in freshwater aquatic habitats (45%), followed by anthropogenic/urban sites (50%), shrubland (17.5%), forest (15%), and cropland (2.5%) biomes. It was detected from water (45%), soil (35%), air (17.5%), and dust (2.5%). These samples were collected across a wide elevational range, 26–948 m, suggesting no strong restriction to highor low-altitude habitats. Occurrences are associated with MAT ~15.4 °C and MAP ~1 059 mm/year. Notes. Pleurophragmium was originally described with a single species, P. bicolor, with the protologue and accompanying illustration (Costantin 1888: fig. 70) reproduced here in Fig. 6. The species was treated in detail under the synonymous names P. simplex by Ellis (1968), and P. parvisporum by Holubová-Jechová (1972) and Matsushima (1975). These authors reported a relatively broad conidial length range based on multiple collections from wood and herbaceous stems: 10–21 µm (mostly 15.2 µm) × 3.5–6 µm (mostly 4.5 µm) (Ellis 1968), and 12.5–22.5 (mostly 15–18.5 µm) × 3.5–6.5 µm (mostly 5 µm) (Holubová-Jechová 1972); for Matsushima’s measurements, see below. Collections from the Czech Republic and Japan examined in this study are consistent with these accounts, although the maximum conidial length observed was slightly shorter than in the published ranges. Our observations confirm that within a single species, conidial dimensions can be variable and are fully congruent with previous descriptions. In the Japanese material used in this study, conidial dimensions observed both in culture (CBS 770.83: 9.5–14 × 3–4.5 µm) and on the natural substrate (CBS H-3522: 10–13.5 × 3.5–4.5 µm) were highly similar and fall within the range of variability reported for this species from Europe (Ellis 1968; Holubová-Jechová 1972; this study). Although conidia from the Japanese material were somewhat shorter than those of the Czech specimens (10–18 × 3.5–6 µm on the natural 33 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Figure 8. Pleurophragmium parvisporum (PRA-24032). A Sporulating conidiophores B–F conidiophores with conidiogenous cells and conidia G Basal cell H–L conidia. Images: on natural substrate. Scale bars: 300 µm (A); 20 µm (B, C); 10 µm (D–L). 34 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum substrate, this study), their overall morphology is indistinguishable, including the subtle characters such as the ornamentation of the outer conidial wall due to collapsing sheath, a feature not previously reported in the literature but clearly visible under DIC and PHC microscopy (Figs 7, 8). Notably, Matsushima (1975) reported three Japanese collections of P. parvisporum from decaying wood of Lithocarpus edulis and Quercus sp., with varying conidial sizes in culture: 14.5– 25 × 4.5–6.5 µm and 10–16 × 4–6 µm. We therefore concluded that CBS 770.83 and the Czech material of P. parvisporum listed above are conspecific and represent a reliable reference for this species. Attempts to extract DNA from the six herbarium specimens from the Czech Republic were unsuccessful. The ITS distribution data derived from the GlobalFungi database suggest that P. parvisporum has a broad ecological amplitude (Fig. 5), occurring across diverse biomes and climatic conditions, but with a notable concentration in aquatic environments in East Asia (China, Japan). Records from Spain confirm the distribution of the species in Europe, although based on published data (see above) the species has been repeatedly reported from several European countries. Records from North America, from where the species was also reported (Wang 2010), and other parts of the world (GBIF Secretariat 2023), are absent in the GlobalFungi database. This broad ecological and climatic range may indicate genuine ecological versatility, or alternatively, the existence of a species complex, as suggested by the variability in conidial length and occurrence on both herbaceous and woody hosts. To clarify its taxonomy, molecular data from additional strains collected across diverse geographical regions are essential for stabilising the species concept. Among Pleurophragmium species, P. parvisporum is most comparable to P. fluviale, P. luguense, and P. septatum in its conidial morphology. These taxa share obovoid to fusoid-ellipsoid conidia that are broadly rounded at the apex, tapering to a narrow base with a hilum, predominantly 3-septate, and hyaline to subhyaline, becoming pale brown at maturity. In the absence of molecular data, distinguishing among these species would be highly challenging. For detailed comparison, see Table 3. Interestingly, several biologically active compounds, collectively known as dactylfungins, were isolated from P. parvisporum (strain D500 obtained from a dead leaf in Japan) and were reported to exhibit antifungal activity against Candida pseudotropicalis and other fungi (Xaio et al. 1993). However, these findings should be interpreted with caution due to the simple morphology of the species and the recently recognised taxonomic ambiguity among preserved strains identified as P. parvisporum. Pleurophragmium pteridophytophilum (Jing Y. Zhang, K.D. Hyde & Y.Z. Lu) Réblová & Hern.-Restr., comb. nov. MycoBank No: 860808 Basionym. Neomyrmecridium pteridophytophilum Jing Y. Zhang, K.D. Hyde & Y.Z. Lu, Fungal Diversity 132: 375. 2025. Typus. CHINA • Guizhou Province, Zunyi City, Chishui County, Hushi Town, Chishui Alsophila Natural Reserve; on dead frond stalks of Pteridaceae sp. in terrestrial habitats; 28 Jul 2022; J. Y. Zhang BL9 (holotype GZAAS 23-0664, extype culture KUNCC 23-13858). 35 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Pleurophragmium septatum (Crous) Réblová & Hern.-Restr., comb. nov. MycoBank No: 860809 Basionym. Neomyrmecridium septatum Crous, Persoonia 41: 287. 2018. Typus. THAILAND • Ratchaburi Province; on leaves of unidentified vine; 2008, P. W. Crous HPC 2252 (holotype CBS H-23768, culture ex-type CPC 34585 = CBS 145073). Pleurophragmium sichuanense (Yan P. Chen & Maharachch.) Réblová & Hern.-Restr., comb. nov. MycoBank No: 860810 Basionym. Neomyrmecridium sichuanense Yan P. Chen & Maharachch., Phytotaxa 701(2): 134. 2025. Typus. CHINA • Sichuan Province, Chengdu City, Dujiangyan City, Longchi National Forest Park; elevation 702 m a.s.l.; 31°00.18'N, 103°38.77'E; on decaying branches of an unidentified herbaceous plant; 5 Oct 2021; Y. P. Chen & W. H. Tian LC64 (holotype HUEST 24.0068). Notes. There is a discrepancy between the protologue and the accompanying figures of P. sichuanense (Chen et al. 2025: fig. 3). While the conidia were described as subhyaline to pale brown, 0–1-septate, and finely verrucose, these features are not clear in the original image. The conidia appear hyaline, smooth, and 1–3-septate in the figures. Species with ornamented conidia are relatively rare in Pleurophragmium and currently include P. parvisporum, P. sichuanense along with two other species: P. clavatum (Ma et al. 2014) and P. verruculosum (Tiwari 1969). Pleurophragmium sorbicola (Crous & R.K. Schumach.) Réblová & Hern.- Restr., comb. nov. MycoBank No: 860811 Basionym. Myrmecridium sorbicola Crous & R.K. Schumach., Fungal Syst. Evol. 1: 191. 2018. Synonym. Neomyrmecridium sorbicola (Crous & R.K. Schumach.) Crous, Persoonia 41: 287. 2018. Typus. GERMANY • near Berlin; on branch of Sorbus aucuparia; 17 Feb 2016; R. K. Schumacher (holotype CBS H-23405, culture ex-type CPC 30455 = CBS 143433). Pleurophragmium s. lat.: species of uncertain status Altogether, 19 species and varieties are recognised as members of Pleurophragmium s. lat. in this study. This section comprises a morphologically heterogeneous assemblage of species historically described in Pleurophragmium, but for which molecular data are lacking. While some species conform to the generic concept and remain plausible candidates for inclusion in Pleurophragmium s. str., others diverge considerably from it. Notably, only three species produce hyaline to subhyaline conidia, occasionally appearing fuscous in mass, whereas the remaining 36 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum taxa bear pale brown, yellowish brown, brown, or dark brown conidia, often with distinctly paler end cell(s). To capture this variability and highlight inconsistencies in morphological characters, diagnostic features of Pleurophragmium s. lat. species currently retained in the genus are summarised in Table 3. However, the true affinities of these taxa cannot be reliably assessed until molecular data become available, emphasising the need for targeted recollection and sequencing. We provide details on the holotype to facilitate future recollection of these species by fellow mycologists. Pleurophragmium angamosense Matsush., Matshushima Mycol. Mem. 8: 30. 1995. Typus. PERU • Colonia Angamos; on the decayed petiole of palm; Jul 1994; T. Matsushima (holotype MFC-4P738). Notes. The species differs from the Pleurophragmium concept primarily in the morphology of its conidiogenous cells and conidia. Although the cells proliferate sympodially, the conidiogenous loci are conspicuous, appearing as distinct scars 3–3.5 µm wide. Conidia are fusiform, transversely septate, smooth, and brown, with the terminal cells noticeably paler, and with dark, thick-walled septa. The original illustration by Matsushima (1995) further suggests that conidial secession may be rhexolytic, as remnants of the outer wall appear to remain attached both to the conidiogenous locus and to the basal part of the conidium. These features clearly contrast with Pleurophragmium s. str., in which conidiogenous loci are reduced to minute denticles and conidial secession is schizolytic. Pleurophragmium aquaticum R.F. Castañeda, Heredia & R.M. Arias, Mycotaxon 101: 92. 2007. Typus. MEXICO • Veracruz, “Los Tuxtlas”; on decaying wood submerged in a stream; 19 May 2002; R. M. Arias & J. Y. C. Elizondo (holotype XAL CB743, isotype: MUCL 45625). Notes. Conidia formed on holoblastic-denticulate conidiogenous cells are fusiform to clavate, occasionally navicular, and often sub-umbonate at the apex. They are brown, becoming pale brown to subhyaline toward the ends and around the septa. By their shape and pigmentation, the conidia closely resemble those of P. miniumbonatum (Heredia et al. 2007). Pleurophragmium aquaticum also resembles P. fusiforme in having fusiform, pigmented conidia with paler ends, but the latter species differs in having narrower and uniformly pale brown conidia except the ends (without the paler bands at the septa). Pleurophragmium bitunicatum Matsush., Icon. microfung. Matsush. lect.: 115. 1975. Typus. JAPAN • Tokyo Prefecture, Hachijo Island; on decaying leaves of unidentified dicotyledon plant; Feb 1969; T. Matsushima (holotype MFC-1669). 37 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Notes. The species is remarkably similar to members of Aquapteridospora (Yang et al. 2015). Another species, P. bambusinum, was recently transferred to Aquapteridospora based on molecular data (Bao et al. 2021). Pleurophragmium clavatum L.G. Ma & X.G. Zhang, Mycotaxon 127: 216. 2014. Typus. CHINA • Yunnan Province, the Forbidden Forest of Banna; on dead branches of Beilschmiedia percoriacea (Lauraceae); 31 Oct 2011; L. G. Ma (holotype HSAUP H2090, isotype HMAS 243416). Notes. Ma et al. (2014) described three morphologically similar species, P. clavatum, P. ellipsoideum, and P. yunnanense, characterised by yellowish-brown to pale brown conidia and conidiophores that often terminate in a denticulate rachis and have distinct nodulose swellings along the whole length. These swellings, however, are not typical for Pleurophragmium based on observations of species on natural substrates, where the fertile region of the conidiophore is restricted to its upper part and bears terminal and intercalary conidiogenous cells. In culture, by contrast, we frequently observed such swellings in P. parvisporum. Conidia of P. clavatum are consistently aseptate, whereas in Pleurophragmium conidia are invariably septate, with both aseptate and multi-septate forms occurring within a single species. Molecular data are therefore needed to confirm the placement of these species in the genus. Pleurophragmium ellipsoideum L.G. Ma & X.G. Zhang, Mycotaxon 127: 214. 2014. Typus. CHINA • Yunnan Province, the Forbidden Forest of Banna; on dead branches of Bauhinia acuminata L. (Caesalpiniaceae); 17 Oct 2008; L. G. Ma (holotype HSAUP H0042, isotype HMAS 243411). Notes. See notes under P. clavatum. Pleurophragmium indicum D’Souza & Bhat, Mycotaxon 119: 477. 2012. Typus. INDIA • Goa, Molem Wildlife Sanctuary; on fallen dead and decaying leaves of Dendrocalamus strictus (Poaceae); 11 Mar 1999; M. A. D’Souza (holotype GUBH 367). Notes. The species is characterised by distinctly versicolorous conidia, with the middle cells dark brown, the end cells pale brown, and the septa conspicuously thick-walled and dark brown. In overall morphology, it bears resemblance to members of Aquapteridospora (Yang et al. 2015). Pleurophragmium harunganae Hansf., Mycol. Pap. 15: 211. 1946. Typus. UGANDA • Entebbe Road, on sori of Hemileia on leaves of Harungana madagascariensis; C. G. Hansford 2803, 3013. 38 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Notes. The species is parasitic on the sori of Hemileia (Pucciniales). It was described as having dematiaceous conidiophores with a terminal denticulate rachis, accompanied by additional, disconnected fertile regions along the conidiophore axis. Conidia are aseptate, curved, fusiform, and subhyaline to pale olivaceous. Based on these features, the species appears more consistent with morphologically similar genera such as Ramichloridium or Myrmecridium, pending recollection and the acquisition of molecular data to confirm its placement. Pleurophragmium malaysianum Matsush., Matsushima Mycol. Mem. 9: 20. 1996. Typus. MALAYSIA • Selangor Darul Ehsan, Ulu Gombak, The University of Malaya Field Stusy Centre; on decaying leaves of a deciduous tree; 12 Jun 1995; T. Matsushima (holotype MFC-5T054). Notes. The species is characterised by cylindrical to clavate, pseudoseptate conidia (Matsushima 1996), features that have not previously been observed in members of Pleurophragmium s. str. Pleurophragmium miniumbonatum (R.F. Castañeda, Iturr. & Guarro) R.F. Castañeda, Mycotaxon 101: 96. 2007. Basionym. Cordana miniumbonata R.F. Castañeda, Iturr. & Guarro, Mycotaxon 73: 5. 1999. Typus. VENEZUELA • Estado de Aragua, Parque Nacional “Henry Pittier”, Estación Rancho Grande, Camino de Interpretación de la Naturaleza “Andy Fields”, in undisturbed rain forest; on fallen decaying leaves on an unidentified plant; 25 Nov 1997; R. F. Castañeda & T. Iturriaga (holotype MUCL 40700). Notes. Heredia et al. (2007) transferred C. miniumbonata to Pleurophragmium based on its superficial resemblance to P. aquaticum (in the same study) and the presence of conidiophores with a terminal denticulate fertile region, a feature more consistent with Pleurophragmium than with Cordana (Preuss 1851; Hernández-Restrepo et al. 2014). The conidia of P. miniumbonatum are dark brown, except for the hyaline basal cell, and exhibit an umbonate apex with variable shapes ranging from obovoid and pyriform to broadly clavate. However, such dark brown conidia are atypical of Pleurophragmium under its current circumscription. Pleurophragmium naviculiforme Matsush., Icon. microfung. Matsush. lect.: 115. 1975. Typus. JAPAN • Okinawa Prefecture, Iriomote island; on decaying leaves of an unidentified deciduous tree; Feb 1972; T. Matsushima (holotype MFC-4355). Notes. The species is characterised by 1-septate, navicular conidia acute at the apex and remain hyaline, both individually and in mass. The fertile 39 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum region of the conidiophore is apical, relatively small, and often appears inflated and curved. Morphologically, the species resembles P. naviculare (Yang et al. 2023), but the latter differs in possessing a longer fertile region and smaller, verrucose conidia that become pale brown with hyaline end cells at maturity. Molecular data are needed to determine whether P. naviculiforme represents a species belonging to a distinct genus or falls within the interspecific variability of Pleurophragmium. Pleurophragmium obcampanuloides Matsush., Matsushima Mycol. Mem. 8: 29. 1995. Typus. JAPAN • Okinawa Prefecture, Iriomote island; in litter on the forest floor; 1992; T. Matsushima (holotype MFC-2J039). Notes. Due to its unique obpyriform to turbinate conidia, which are septate, pale olivaceous, and olivaceous in mass (Matsushima 1995), this species is readily distinguished from other members of the genus. Pleurophragmium peruamazonicum Matsush., Matsushima Mycol. Mem. 7: 61. 1993. Typus. PERU • Loreto, forest Rio Negro; on decaying palm leaves; Jun 1992; T. Matsushima (holotype MFC-2P079). Notes. The species is characterised by versicolorous, 3-celled conidia with a brown median cell and hyaline end cells. Its conidiophores bear an apical fertile region that often extends along the upper half of the conidiophore axis, with denticles sparsely distributed (Matsushima 1993). Morphologically, it is strikingly similar to P. tricolor (nom. inval., Art. 40.1; Rambelli 2009); for a detailed comparison, see Table 3. While P. peruamazonensis was described from culture, measurements of P. tricolor were derived from natural material. Consequently, features that distinguish the two taxa, such as conidial size and the extent of the apical fertile region, may simply reflect differences between in vitro and in vivo observations. Pleurophragmium peruamazonicum var. inflatum Matsush., Matsushima Mycol. Mem. 7: 61. 1993. Typus. PERU • Loreto, forest Rio Negro; on decaying leaves of a deciduous tree; Nov 1990; T. Matsushima (holotype MFC-0P570). Notes. The variety inflatum differs from the type variety by the conspicuous inflation of the basal portion of the conidium (Matsushima 1993). It is further characterised by thick-walled septa, with the two basal cells pale brown. However, it remains uncertain whether these distinctions fall within the natural intraspecific variability of the species or represent a separate species. 40 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Pleurophragmium subfusiforme Matsush., Icon. Microfung. Matsush. lect.: 116. 1975. Typus. JAPAN • Mie Prefecture, University of Mie, Hirakura Exp. Forest; forest; Sep 1965; T. Matsushima (holotype MFC-1551). Notes. Pleurophragmium subfusiforme is readily distinguished from other species of the genus by its large (21–38 × 5.8–9.2 µm), fusiform conidia, which are 3–7-septate and sometimes slightly constricted at the septa. The conidia are subhyaline, becoming pale fuscous in mass (Matsushima 1975). Pleurophragmium taiwanense Matsush., Matsushima Mycol. Mem. 5: 24. 1987. Synonym. Pleurophragmium bicolor Matsush., Icon. microfung. Matsush. lect.: 114. 1975. Nom. illegit. (ICN, Art. 53.1) non Pleurophragmium bicolor Costantin, Costantin, Mucéd. Simpl.: 100. 1888. Typus. TAIWAN • Ken-Ting Park; on rotten leaf-rachis of Arenga engleri; 27 May 1980; T. Matsushima (holotype MFC-10120). Notes. This species produces cylindro-clavate to somewhat turbinate conidia with a slightly inflated apical cell. Observations from culture show that the fertile regions of the conidiophore are discontinuous, occurring apically as well as at several points along the conidiophore axis, sometimes associated with nodose swellings (Matsushima 1987). Morphologically, it resembles P. peruamazonicum but can be distinguished by its shorter conidia and fewer septa. For additional comparison, see notes to P. asiaticum. Pleurophragmium tricolor Rambelli, Flora Mediterranea 19: 82. 2009. Nom. inval. (ICN, Art. 40.1). Typus. ITALY • Pantelleria, Montagna Grande; on dead leaves of Arbutus unedo (a7); (holotype PAL). Pleurophragmium varieseptatum Matsush., Icon. microfung. Matsush. lect.: 117. 1975. Typus. JAPAN • Kyoto City; on decaying stem of Phyllostachys edulis; Feb 1966; T. Matsushima (holotype MFC-1964). Notes. The species is readily distinguished by its cylindrical, pale olivaceous conidia, which display considerable variability in both size and septation, ranging from one to four septa (Matsushima 1975). Pleurophragmium verruculosum D.P. Tiwari, Indian Phytopath.: 513. 1970. Typus. INDIA • Madhya Pradesh, Sagar; rhizosphere soil of Piper betle; Dec 1965; D. P. Tiwari (holotype IMI 134426, isotype ITCC New Delhi 1375). 41 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Notes. Verrucose conidia are rare in Pleurophragmium and are known only from a few species, namely P. clavatum, P. sichuanense, and P. verruculosum. Ornamented conidia are also observed in P. pteridophytophilum as “rough-walled” (Zhang et al. 2025), and in P. parvisporum (this study), where the conidial wall appears somewhat longitudinally crumpled. Among these taxa, molecular data are currently available only for P. sichuanense and P. parvisporum. Morphologically, P. verruculosum is most similar to P. clavatum, but can be distinguished by its septate conidia and shorter conidiophores. Pleurophragmium yunnanense L.G. Ma & X.G. Zhang, Mycotaxon 127: 215. 2014. Typus. CHINA • Yunnan Province, the Forbidden Forest of Banna; on dead branches of Machilus salicina (Lauraceae); 18 Oct 2008; L. G. Ma (holotype HSAUP H0086, isotype HMAS 243412). Notes. The species produces pale brown, broadly fusiform conidia tapering at both ends, which are mostly 3-septate (Ma et al. 2014). In conidial size and septation, it resembles P. ellipsoideum from the same study, but the latter differs in having ellipsoidal conidia that are broadly rounded at the apex. For additional information, see the notes under P. clavatum. Species excluded from Pleurophragmium and described in other genera Here, we provide a list of species that have been excluded from Pleurophragmium and subsequently transferred to other genera by various authors. Synonymy follows the records in MycoBank, unless stated otherwise. Aquapteridospora bambusinum (D.Q. Dai & K.D. Hyde) D.F. Bao, J. Fungi 7(669): 10. 2021. Basionym. Pleurophragmium bambusinum D.Q. Dai & K.D. Hyde, Fungal Diversity 82: 92. 2016. Dactylaria arecae (Matsush.) R.F. Castañeda & W.B. Kendr., Univ. Waterloo, Biol. Ser. 35: 26. 1991. Basionym. Pleurophragmium arecae Matsush., Matsushima Mycol. Mem. 5: 23. 1987. Dactylaria cylindrospora (Matsush.) R.F. Castañeda & W.B. Kendr., Univ. Waterloo, Biol. Ser. 35: 27. 1991. Basionym. Pleurophragmium cylindrosporum Matsush., Icon. microfung. Matsush. lect.: 115. 1975. 48 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum unbranched, sometimes percurrently elongating, flexuous to sinuous, sometimes becoming geniculate exhibiting a zig-zag pattern due to local bending above the septum, each bend is associated with the formation of a single denticle on the ‘outside’ giving the appearance of irregular or dichotomous branching, pale brown to golden brown, smooth-walled, septate. Conidiogenous cells 10–24 × 3–4 µm, integrated, terminal, form transverse septa during sympodial extension and become intercalary, occasionally lateral growing directly on hyphae, monoblastic or polyblastic with 1–4 peg-like denticles, cylindrical or subulate, subhyaline to pale olivaceous-brown when in the terminal position, golden-brown when intercalary, smooth-walled; conidiogenesis holoblastic-denticulate. Conidia 18.5–36(–45) × (3–)4–5.5 µm (mean ± SD = 27.1 ± 5.6 × 4.9 ± 0.1 μm), solitary, dry, acropleurogenous, oblong to cylindrical, elongate fusiform to narrowly ellipsoidal, tapering at both ends, truncate at the base 1–1.5 µm wide, with a conspicuous basal scar, usually straight, occasionally slightly curved, often with guttules or granules visible inside the cells, smooth-walled, aseptate and hyaline when young, at maturity with 3–7 transverse septa and subhyaline to pale olivaceous-brown, olivaceous-grey in mass; conidial secession schizolytic. Habitat and geographical distribution. Skoliomycella flava is a saprobe occurring on plant remnants. To date, two confirmed records originate from the Iberian Peninsula, specifically from Portugal (this study) and Spain (GlobalFungi), within a temperate, Mediterranean climate. According to GlobalFungi, it was detected in a single air sample from an anthropogenic biome, MAT ~14.5 °C, MAP ~575 mm/year. Notes. Skoliomycella flava is readily distinguished from the morphologically similar species attributed to Camporesiomyces (Hyde et al. 2020) and Zaanenomyces (Crous et al. 2021) by the absence of a rachis in the fertile apical portion of the conidiogenous cell. Instead, its conidiophores are flexuous to sinuous, sometimes exhibit a zig-zag pattern, with a single denticle and/or a slightly prolonged conidiogenous cell formed at each bend on the outer side of the conidiophore, giving the impression of irregular or dichotomous branching. The detection of S. flava in urban air highlights its capacity for aerial dispersal, although it occurs at very low relative abundance. However, its global distribution remains uncertain, as no additional records outside the Iberian Peninsula are available. Whether S. flava represents a rare, geographically restricted lineage, or whether its scarcity reflects limited sampling, or under-detection due to low environmental abundance, remains to be determined. Thysanorea Arzanlou, W. Gams & Crous, Stud. Mycol. 58: 80. 2007. emend. Hern.-Restr. & Crous, Fungal Syst. Evol. 6: 17. 2020. Description. See Arzanlou et al. (2007) and Hernández-Restrepo et al. (2020). Notes. Thysanorea is characterised by microto macronematous, dematiaceous conidiophores that are simple or apically branched, bearing terminal or intercalary, holoblastic, polyblastic conidiogenous cells. Conidia are solitary, acropleurogenous, transversely septate, pale brown, and variable in shape, most commonly oblong, obovoid, or fusiform. In contrast, the associated synasexual morph is phialidic. 49 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Three species previously classified in Uncispora were shown to be congeneric with Thysanorea and are here formally transferred to the genus, with new combinations proposed below. Thysanorea acropleurogena Réblová & Hern.-Restr., sp. nov. MycoBank No: 860719 Fig. 10 Etymology. From Greek akros (apex), pleurá (side) and genēs (born, produced). Referring to the mode of conidium development, in which conidia are formed both terminally and laterally on the conidiogenous cells. Typus. PAPUA NEW GUINEA • Madang Province, foothills of Finisterre Range, 40.8 km along road to Lae; 200 m a.s.l.; on unidentified wood; 2 Nov 1995; A. Aptroot 36665 (holotype CBS H-25780 dried culture, ex-type culture CBS 215.96). Culture characteristics. On CMD colonies 44–45 mm diam., circular, flat, margin entire, lanose, pinkish-brown to brown, darker at the margin, reverse dark brown. On MLA colonies 36–38 mm diam., circular, convex, margin entire, floccose to lanose, olivaceous-grey to mouse grey, dark olivaceous at the margin, reverse dark olivaceous. On OA colonies 40–41 mm diam., circular, flat, margin entire, lanose at the centre and on the inoculation block, cobwebby towards the periphery, with a well-defined grey-brown central zone, dark brown towards the periphery, with a diffuse lighter halo at the margin, reverse dark brown. On PCA colonies 40–42 mm diam., circular, flat, margin entire, lanose, grey-brown, cobwebby and dark brown towards the periphery, reverse dark brown. With a prominent submerged growth on all media. Sporulation on PCA, absent on CMD, MLA and OA. Description in culture. Colonies on PCA effuse. Sexual morph. Not observed. Asexual morph. Mycelium composed of subhyaline to pale olivaceous brown, septate hyphae, 1.5–2.5 µm wide. Conidiophores 98–145 × 3.5–4.5 µm, basal cells sometimes slightly inflated 5–6.5 µm wide, macronematous, mononematous, loosely scattered to densely fasciculate, mostly erect, straight to slightly flexuous, cylindrical, subtly undulate, unbranched to loosely branched in the upper part, pale to medium brown to pale olivaceous brown, darker at the base, smooth-walled, septate. Conidiogenous cells 23.5–28(–40) × 3.5–5(–5.5) µm, integrated, terminal, forming transverse septa during sympodial proliferation and often becoming intercalary, arranged either along most of the conidiophore length or restricted to the upper part, forming a rachis with numerous, closely spaced, minute denticles, polyblastic, cylindrical, pale brown to pale olivaceous brown, end cells tend to be paler, smooth-walled; conidiogenesis holoblastic-denticulate. Conidia (13.5–)15–21 × (3.5–)4–5.5 µm (mean ± SD = 17.2 ± 1.8 × 4.5 ± 0.3 μm), solitary, dry, acropleurogenous, subcylindrical to oblong or fusiform to fusiform-clavate, tapering towards both ends, truncate at the base 1–1.5 µm wide, with a conspicuous basal scar, straight or slightly curved, pale brown to pale olivaceous brown, darker at the base, dark olivaceous brown in mass, smooth-walled, (1–)3-septate; conidial secession schizolytic. 50 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Figure 10. Thysanorea acropleurogena (ex-type CBS 215.96). A Sporulating conidiophores B, C upper part of the conidiophore conidiogenous cells forming a rachis and attached conidia D–G conidiophores, conidiogenous cells and conidia H conidia I diversity of colony morphology on CMD, MLA, OA, and PCA, respectively (from left to right) after 4 wk. Images: on PCA (A–H). Scale bars: 300 µm (A); 10 µm (B–H); 1 cm (I). 51 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Habitat and geographical distribution. The species is a saprobe occurring on decaying wood and is currently known only from Papua New Guinea. No identical ITS sequences were recovered in the GlobalFungi database. Notes. Phylogenetic analyses (Fig. 2) revealed that T. acropleurogena forms a sister relationship to a subclade comprising three closely related species, T. hainanensis, T. sinensis, and T. wuzhishanensis. They belong to a subclade that also includes T. seifertii and T. cantrelliae. Among known species, T. acropleurogena is closest to T. obscura and T. seifertii in having lightly pigmented, subcylindrical to oblong, predominantly 3-septate conidia. However, both species are readily distinguishable in conidial characters and having unbranched conidiophores. Thysanorea obscura differs from the new species in subhyaline to yellowish brown, larger conidia, 20–31 × 5–8 µm (Matsushima 1983). Thysanorea seifertii produces similar pale brown, subcylindrical to clavate or oblong conidia, but these are considerably shorter, 7–15 × 1.5–3 µm (Hernández-Restrepo et al. 2020). In addition, T. seifertii produces a phialidic synasexual morph in culture, a feature so far unique among Thysanorea species. In the phylogenetic tree, all three species are resolved as distinct lineages. The original material, CBS H-6267, from which the axenic culture was derived, consists of multiple twigs. However, we could not locate the target fungus. Several twigs are covered with aerial mycelium, while others carry fertile conidiophores of a dendryphiella-like fungus. Accordingly, a dried culture was designated as the holotype of T. acropleurogena. Thysanorea acropleurogena appears to represent a rare species, confirmed thus far from a single collection on decaying wood. The absence of identical ITS sequences in the GlobalFungi database indicates that it has not yet been detected in environmental sequencing datasets, suggesting that the species may be geographically restricted or currently overlooked due to under-sampling. Thysanorea hainanensis (Jian. Y. Li & Z.F. Yu) Réblová & Hern.-Restr., comb. nov. MycoBank No: 860812 Basionym. Uncispora hainanensis Jian. Y. Li & Z.F. Yu, Mycotaxon 129: 474. 2015. Typus. CHINA • Hainan Province, Wuzhishan National Nature Reserve; 754 m a.s.l., isolated from decayed leaves; Dec 2011; G. Z. Yang (holotype YMF 1.04038, ex-type culture YMF1.040381). Thysanorea melanica (Hong Y. Su, Udayanga & K.D. Hyde) Hern.-Restr. & Crous, Fungal Syst. Evol. 6: 18. 2020. Basionym. Minimelanolocus melanicus Hong Y. Su, Udayanga & K.D. Hyde, Fungal Biol. 119: 1056. 2015. Culture characteristics. On CMD colonies 51–52 mm diam., circular, flat to slightly raised in the centre, margin diffuse, entire, zonate, floccose to velvety, white-beige in the centre, brown to dark brown towards the margin, reverse dark brown to black. On MLA colonies 49–51 mm diam., circular, flat, raised at the centre, margin entire, zonate, lanose to floccose centrally, becoming mucoid and glossy towards the periphery, lanose at the margin, cream to buff in the 52 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum central region, sharply contrasting with the surrounding, dark grey to blackish mucoid mycelium, olivaceous-grey at the margin, reverse dark olivaceous-grey to black. On OA colonies 47–50 mm diam., circular, flat, margin diffuse, entire, floccose to slightly lanose at the centre, becoming cobwebby towards the periphery, whitish to pale pinkish-buff, surrounded by a wide zone of submerged, dark olivaceous-grey mycelium that diffuses into the agar, paler at the margin, reverse uniformly dark olivaceous-grey to black. On PCA colonies 41–42 mm diam., circular, flat to slightly raised in the centre, margin diffuse, entire, floccose to somewhat lanose, central zone pale pink-brown-buff, surrounded by a dark brown submerged zone, paler at the margin, reverse dark olivaceous-brown to nearly black. Sporulation absent on all media. Description in culture. Colonies on PCA effuse. Sexual morph. Not observed. Asexual morph. Mycelium composed of subhyaline to pale brown, septate hyphae, 1.5–3 µm wide. Conidiophores, conidiogenous cells and conidia absent. Specimen examined. THE NETHERLANDS • North Holland Province, Wieringermeer Polder, Van Bemmelen Hoeve; isolated from wheat field soil; May 1966; W. Gams (living culture CBS 862.68). Habitat and geographical distribution. Thysanorea melanica was originally described from decaying wood in China (Liu et al. 2015), with an additional record from wheat field soil in the Netherlands (this study). According to the GlobalFungi database, the species has been detected in 266 environmental samples across three continents. Its ITS dataset is strongly Eurocentric, with ~90% of detections originating from Europe, where it is recorded in multiple countries, showing notable hotspots in Estonia, Switzerland, and Germany. Asia contributes ~8% (driven largely by China), and North America is sparsely represented by ~2%. It is most frequently detected in cropland (50%), followed by grassland (20%), forest (17%), anthropogenic habitats (9.4%), woodland (2.3%) and shrubland (0.4%) biomes. Most records are from soil samples including topsoil and rhizosphere soil (93.2%), with minor representation in roots and shoots. Occurrences are associated with MAT ~8.4 °C and MAP ~749 mm/year. Notes. Although strain CBS 862.68 did not sporulate on any of the culture media tested, molecular data enabled its placement in the genus Thysanorea and as conspecific with T. melanica (Fig. 2). Thysanorea melanica is nested within a well-supported subclade comprising six additional species. It is characterised by unbranched, dark brown conidiophores bearing terminal and intercalary conidiogenous cells, and by pale brown conidia that are 1–3-septate when young and 4–6-septate at maturity, measuring (9–)13–37(–45) × (2.5– )3.5–6.5(–8) μm (Liu et al. 2015). Thysanorea melanica is a cosmopolitan, soil-dwelling saprobe, occasionally associated with plant tissues, that thrives across a wide range of environments with a preference for temperate to cool-temperate climates with moderate rainfall. Its frequent occurrence in croplands, grasslands, and forests highlights its ecological versatility and ability to persist in both natural and anthropogenic ecosystems. The predominance of records from cropland soils further implies that agricultural activities may have facilitated its spread and persistence, potentially through soil transport or crop-associated dispersal. This hypothesis is consistent with the occurrence of CBS 862.68 strain, which was isolated from wheat field soil. 53 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Thysanorea sinensis (G.Z. Yang & Z.F. Yu) Réblová & Hern.-Restr., comb. nov. MycoBank No: 860813 Basionym. Uncispora sinensis G.Z. Yang & Z.F. Yu, Mycotaxon 116: 172. 2011. Typus. CHINA • Yunnan province, Mengla County, Xishuangbanna Tropical Botanical Garden; on submerged leaves of an unidentified dicotyledonous plant; Sep 2010; G. Z. Yang (holotype YMF 1.03683; ex-type culture YMF 1.03683). Thysanorea wuzhishanensis (L.P. Chen & Z.F. Yu) Réblová & Hern.-Restr., comb. nov. MycoBank No: 860814 Basionym. Uncispora wuzhishanensis L.P. Chen & Z.F. Yu, Sydowia 70: 255. 2018. Typus. CHINA • Hainan Province, Wuzhishan National Nature Reserve; 754 m a.s.l.; on submerged decaying leaves in a stream; 30 Jun 2011; Z. F. Yu (holotype YMF 1.04080). Wongia Khemmuk, Geering & R.G. Shivas, IMA Fungus 7: 249. 2016. Description. See Khemmuk et al. (2016). Notes. To date, 11 species have been described in Wongia. The asexual morphs produce macronematous conidiophores with holoblastic, polyblastic, sympodially elongating conidiogenous cells that generate septate, subhyaline to dark brown, transversely septate or rarely aseptate conidia (e.g. Bao et al. 2021; Crous et al. 2022; Manawasinghe et al. 2024; Wang et al. 2025). Wongia pallidopolaris Réblová & Hern.-Restr., sp. nov. MycoBank No: 860720 Fig. 11 Etymology. From Latin pallidus (pale), and polaris (of or relating to the poles). Referring to the conidial pigmentation, in which the apical and basal cells are distinctly paler than the central, more pigmented cells, creating a noticeable bipolar contrast. Typus. THE NETHERLANDS • Gelderland Province, Wageningen; isolated from sandy soil under continuous wheat; Jan 1970; J. H. van Emden No. 4118, 30 (holotype CBS H-25781 dried culture, ex-type culture CBS 440.70). Culture characteristics. On CMD colonies 40–41 mm diam., circular, flat, margin entire to slightly fimbriate, diffuse, lanose, with a subtle concentric zoning, beige to grey-beige at the centre, brown at the margin, reverse dark brown. On MLA colonies 38–40 mm diam., circular, raised, margin entire, lanose, composed of camel brown and pale olivaceous brown concentric zones, reverse dark brown. On OA colonies 49–51 mm diam., circular, raised, margin entire, lanose, composed of beige, camel brown, dark brown and cinnamon concentric zones, brown at the margin, aerial hyphae at the centre and margin bearing numerous colourless exudates, reverse brown. On PCA colonies 44–45 mm diam., 54 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum circular, convex, margin entire, lanose, whitish-grey at the centre surrounded with a thin smoke-grey zone, pale beige to light fawn towards the margin, reverse dark brown. Sporulation abundant on CMD, MLA, and PCA, absent on OA. Description in culture. Colonies on MLA effuse. Sexual morph. Not observed. Asexual morph. Mycelium composed of pale brown, septate, sparsely branched hyphae, 1.5–3 µm wide. Conidiophores 24–70 × 3.5–5(–5.5) µm, macronematous, mononematous, solitary or aggregated, erect, straight to slightly flexuous, apically almost sinuous, cylindrical, unbranched, occasionally proliferating sympodially, brown, dark brown in the lower part, smooth-walled, septate. Conidiogenous cells 12–30 × (4.5–)5–6 µm, integrated, terminal, monoor polyblastic, with one to several denticles, extending sympodially, cylindrical, tapering, sometimes slightly swollen at the apex, pale brown, paler at the apex, smooth-walled; conidiogenesis holoblastic-denticulate. Conidia (20–)22–28(–30) × 5.5–6.5(–5.5) µm (mean ± SD = 25.2 ± 1.7 × 6.1 ± 0.3 μm), solitary, dry, acropleurogenous, ellipsoid to fusiform to fusiform-clavate, tapering towards both ends, truncate at the base 2–2.5 µm wide, with a conspicuous basal scar, mostly straight, occasionally slightly curved, brown to dark brown, end cells paler then the middle ones, apical cell often with a dark brown tip, smooth-walled to finely roughened, thick-walled, the outer wall partly detaches from the conidium, the detached segments appear as apical or side pocket or wings, sometimes the outer wall is detached around the base imitating a minute frill, 3-septate, mucoid sheath absent; conidial secession schizolytic. Habitat and geographical distribution. The examined strain was isolated from sandy agricultural soil in the Netherlands. According to the GlobalFungi database, W. pallidopolaris was detected in 812 environmental samples. It is cosmopolitan, widely distributed in temperate to subtropical regions, with most records from North America, Europe, and Asia. The main hotspots are in the USA (Michigan, New York and North Carolina), China (Provinces Fujian, Guizhou, Hebei, Jiangxi, Jilin and Yunnan), and central Europe (particularly Switzerland and the Netherlands), while additional, less frequent records originate from Australia and Africa. The species is primarily soil-associated (86.2%), with occasional detection in roots, shoots and rare occurrence in air or water. It is predominantly associated with cropland (59.1%) and grassland (28%) ecosystems, followed by occasional occurrence in the anthropogenic habitats (6.4%), forest (4.2%) and woodland, aquatic, shrubland and wetland biomes. In both USA and China, W. pallidopolaris is strongly associated with cropland ecosystems, particularly cereals and legumes (Zea mays, Glycine max, Oryza sativa, Chenopodium quinoa). However, in China the species exhibits a broader ecological amplitude, occurring not only in croplands but also in forest soils, rhizospheres, and even aerosols, whereas in the USA it appears to be more restricted to cropland and anthropogenic soils. Occurrences are associated with MAT ~11.8 °C and MAP ~926 mm/year. Notes. Wongia pallidopolaris closely resembles W. aquatica (Luo et al. 2019) in having 3-septate, brown conidia with paler end cells. However, W. aquatica differs in possessing shorter conidia, measuring 17–21 × 5–7 µm. Phylogenetically, both species form a strongly supported sister relationship within a monophyletic clade that is basal to all remaining Wongia species (Fig. 3). Most conidia lacked any sheath-like structure; however, in a few cases, a similar feature was observed (Fig. 11O). It appeared on one or both sides 55 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Figure 11. Wongia pallidopolaris (ex-type CBS 440.70). A Sporulating conidiophores B–K conidiophores, conidiogenous cells and conidia L–O conidia P diversity of colony morphology on CMD, MLA, OA, and PCA, respectively (from left to right) after 4 wk. Images: on MLA (A–O). Scale bars: 300 µm (A); 10 µm (B–O); 1 cm (P). 56 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum near the base or apex, only at the apex, or occasionally at several points on the same conidium. This structure is slightly pigmented and resembles an outer wall layer that detaches in one or several places rather than a mucoid sheath. In other Wongia species, a mucoid sheath has not been reported. It is possible that in culture, where osmotic conditions differ from those in nature, the outer conidial wall deteriorates and partially detaches. In contrast, a similar structure observed in P. parvisporum represents a true mucoid but ephemeral sheath that is practically invisible on conidia from natural material yet clearly visible in culture, positioned laterally around the middle of the conidium (Fig. 7N–P) or covering the upper two-thirds (Fig. 7D, E). Based on eDNA data, W. pallidopolaris is regarded as a cosmopolitan soil saprobe with strong ecological associations to cropland and grassland ecosystems. Its prevalence in agricultural soils suggests that human activity, particularly through agricultural practices, may have facilitated its dissemination. Its occasional detection in forest soils, rhizospheres, and even aerosols indicates that it can exploit a broader range of habitats. Despite its widespread occurrence in environmental samples, W. pallidopolaris represents a morphologically cryptic fungal lineage that has likely been overlooked in traditional surveys. Wongia rhachidophora Réblová & Hern.-Restr., sp. nov. MycoBank No: 860721 Fig. 12 Etymology. From Greek rhachis (spine, axis) and phoros derived from phora (bearing). Referring to the fertile, rachis-like upper part of the conidiophore, which bears conidia along its length. Typus. INDIA • Bangalore, Arboretum of Forestry Department; on dead leaf of Bambusa sp.; Jun 1973; W. Gams (holotype CBS H-11671 dried culture, extype culture CBS 531.73, paratype CBS H-5414). Culture characteristics. On CMD colonies 45–51 mm diam., circular, flat, powdery, sienna, margin paler, cinnamon to fawn, diffuse, fimbriate to slightly lobate, reverse brown with different tones of dark brick, brick to sienna in concentric zones. On MLA colonies 65–67 mm diam., circular, flat, margin entire to fimbriate, velvety to powdery, centre rosy buff, cinnamon to saffron towards the periphery, reverse brown. On OA colonies 58–60 mm diam., circular, flat, margin entire to fimbriate to floccose, velvety, with concentric brown zones of different tones such as fulvous, sienna to cinnamon, ochreous towards the margin, reverse dark brown (umber) in the centre with ochreous margin. On PCA colonies 53–54 mm diam., circular, flat, margin entire fimbriate to diffuse, velvety to sandy, centre fulvous to ochreous, isabelline towards the margin, with numerous exudates at the centre, reverse of the same colour. Sporulation abundant on all media. Description in culture. Colonies on OA effuse. Sexual morph. Not observed. Asexual morph. Mycelium composed of hyaline to pale brown, smooth hyphae, verrucose close to the conidiophore base, 1–2 μm wide. Conidiophores up to 63 μm long, 2.5–3.5 μm wide at the base, macronematous, mononematous, solitary, erect, straight to slightly flexuous, cylindrical, unbranched, pale brown to brown-orange. Conidiogenous cells 13–52 × 3–4 μm, integrated, terminal, 57 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Figure 12. Wongia rhachidophora (ex-type CBS 531.73). A–C Sporulating conidiophores D–G conidiophores, conidiogenous cells and conidia H–K conidia L diversity of colony morphology on CMD, MLA, OA, and PCA, respectively (from left to right) after 4 wk. Images: on OA (A–K). Scale bars: 50 µm (A, C); 100 µm (B); 10 µm (D–K); 1 cm (L). 64 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum historically placed in Pleurophragmium continues to hinder a robust taxonomic reassessment of the genus. To clarify the taxonomy of Pleurophragmium, we conducted a morphological comparison of the accepted species, which provides additional insights into the morphological affinities and distinctions of its species (Table 3). Based on this evaluation, we did not identify any Neomyrmecridium species that could be regarded as conspecific with already known Pleurophragmium species, although some taxa merit closer examination. Herpotrichiellaceae: novel lineages in Thysanorea The holoblastic-denticulate mode of conidiogenesis, with conidia often borne on a sympodially elongating rachis, occurs in several genera of the Herpotrichiellaceae, including Aciculomyces (Torres-Garcia et al. 2023), Fonsecaea (Najafzadeh et al. 2009), Neoveronaea (Qiu et al. 2022), Rhinocladiella (Melin and Nannfeldt 1934; de Hoog and Hermanides-Nijhof 1977; Müller et al. 1987), Veronaea (Ciferri and Montemartini 1957), and Thysanorea. In addition, Hernández-Restrepo et al. (2020) reported a phialidic synasexual morph of Thysanorea seifertii, which is produced only in culture. Fonsecaea and Rhinocladiella also display marked pleomorphism, exhibiting considerable variation in asexual morphology, including several types of synasexual morphs, and conidial ontogeny under varying conditions (Schol-Schwarz 1968; McGinnis 1983; de Hoog and Hermanides-Nijhof 1977; Tsuneda et al. 1986). The pleomorphic nature of several genera in this family further complicates identifications based solely on morphology and has historically contributed to their polyphyly within the Herpotrichiellaceae. The genus Thysanorea, based on T. papuana, was established for fungi with mature conidiophores bearing several tiers of branchlets arranged apically in a compact cluster, imparting a distinctly arboreous appearance (Arzanlou et al. 2007). The sympodially elongating conidiogenous cells are terminal or intercalary on branchlets, occasionally discrete, with a conspicuous denticulate rachis. Recent studies have shown that the traditionally diagnostic branching pattern of Thysanorea conidiophores, used to distinguish the genus from Periconiella (Saccardo and Berlese 1885), can vary with culture conditions and may be less pronounced in nature or in young cultures (Kirschner 2016; Wang et al. 2019; Hernández-Restrepo et al. 2020). Hernández-Restrepo et al. (2020) pointed out that Thysanorea is closely related to Minimelanolocus (Castañeda-Ruiz 2001). However, the placement of the M. navicularis, the type species of the genus, remains uncertain due to the absence of DNA sequence data, with its presumed phylogenetic position in the Herpotrichiellaceae inferred from other species currently accepted in the genus (e.g. Liu et al. 2015; Wang et al. 2019). Hernández-Restrepo et al. (2020) highlighted morphological differences between M. navicularis, and species currently assigned to Minimelanolocus based on molecular data. Consequently, these authors transferred sequenced Minimelanolocus species to Thysanorea and emended the genus. In addition, molecular data support the transfer of three Uncispora species to Thysanorea, for which we propose new combinations. Our phylogenetic analyses resolved ex-type strains of U. hainanensis (Li et al. 2015), U. sinensis (Yang et al. 2011), and U. wuzhishanensis (Liu et al. 2018a), within the Thysanorea clade. 65 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum This evidence expands the morphological concept of Thysanorea by incorporating new conidial characteristics. These taxa share distinctive apically tapering, cylindrical conidia, curved in the upper portion or terminating in a hooked apical cell, and are morphologically incompatible with U. harroldiae, the type species of Uncispora (Sinclair and Morgan-Jones 1979). The latter is characterised by holoblastic, determinate conidiogenous cells with a single flat locus, whereas the analysed species possess polyblastic, holoblastic cells that elongate sympodially, and conidia are borne on denticles. The phylogenetic position of Uncispora s. str. remains unresolved. These findings emphasise the need for careful identification of Uncispora species, particularly considering the diagnostic value of conidiogenous cell morphology and shape of conidia, but also suggest the polyphyly of Uncispora, warranting a narrower circumscription of the genus based on freshly recollected material. Papulosaceae: expanding the diversity of Wongia Although Wongia was originally established for two sexually reproducing species (Khemmuk et al. 2016), several subsequently described taxa have been defined solely by asexual features (Luo et al. 2019; Bao et al. 2021; Manawasinghe et al. 2024; Yu et al. 2024; Wang et al. 2024). Both new species, W. pallidopolaris and W. rhachidophora, conform to the generic concept of Wongia in producing macronematous, unbranched, dark brown conidiophores with polyblastic, sympodially proliferating conidiogenous cells and septate, brown conidia. Their phylogenetic placement within the genus expands the known diversity of Wongia. Currently, the genus Wongia comprises 13 species. Wongia pallidopolaris and W. aquatica formed a strongly supported subclade and include species with relatively short conidiophores (≤ 90 µm long) and dark brown conidia with distinctly paler end cells. By contrast, all other known species, including W. rhachidophora, represent morphologically distinct and phylogenetically strongly supported group characterised by subhyaline to pale brown conidia that are mostly evenly pigmented, and only rarely have end cells slightly paler than the median cells. Within the second subclades are grouped the only species known that reproduce also sexually, namely W. ficherai, W. griffinii, and W. guttulata (Khemmuk et al. 2016; Wang et al. 2024). They are characterised by perithecial, non-stromatic ascomata, filiform paraphyses, unitunicate asci with a non-amyloid apical ring, and 3-septate, dark brown ascospores with pale brown to subhyaline end cells. Wongia rhachidophora was nested within a complex comprising four other closely related species. While W. suae, W. bandungensis, and W. bambusae are morphologically indistinguishable, differing only by subtle variation in conidial size, W. rhachidophora and W. fusiformis are clearly separated from them by conidial colour, septation, and shape. Comparisons of ITS, tef1, and rpb2 sequences further support W. rhachidophora as a distinct species. In contrast, W. bandungensis and W. suae show almost complete sequence identity in tef1 and rpb2 and differ by 1.4% in ITS. Given their morphological indistinguishability and minimal genetic divergence, these two taxa may represent very recently diverged populations or are likely conspecific, pending further population-level sampling. 66 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Tubeufiaceae: new genera and species with holoblastic-denticulate conidiogenesis Skoliomycella flava and Z. hilifer are new additions to the family Tubeufiaceae. The holoblastic conidiogenesis on denticulate conidiogenous cells is a feature that is widespread and convergent across the family. Skoliomycella and Zaanenomyces are members of a well-supported subclade comprising fungi characterised by holoblastic (mostly holoblastic-denticulate) conidiogenesis and three conidial morphotypes, including Acanthostigma (De Notaris 1863; Réblová and Barr 2000; Boonmee et al. 2014) and Helicosporium (Nees von Esenbeck 1816; Linder 1929; Lu et al. 2018a) with helicosporous conidia, Neodictyospora (Zhang et al. 2023b) with dictyosporous conidia, and Camporesiomyces (Hyde et al. 2020) and Zaanenomyces (Crous et al. 2021) with straight, rarely helicosporous conidia. Among these genera, Skoliomycella is morphologically most similar to Camporesiomyces and Zaanenomyces, but differs in having sinuous to geniculate conidiophores bearing scattered denticles along their entire length. Zaanenomyces, typified by Z. quadripartis, was established by Crous et al. (2021) for dematiaceous hyphomycetes with simple, erect conidiophores, terminal conidiogenous cells that extend sympodially forming a rachis and dry, solitary, hyaline, narrowly obclavate, septate conidia. All three Zaanenomyces species have been recorded from dead culms of Juncus spp. (Crous et al. 2021). Among these, Z. hilifer is closely related to Z. quadripartis. Both species exhibit comparable conidiophore and conidiogenous cell architecture; however, they can be distinguished by differences in conidial morphology. In our preliminary phylogenetic analyses of ITS–LSU sequences (data not shown), Z. moderatricis-academiae (ex-type CBS 148312, and CBS149453) and Z. versatilis (ex-type CBS 148315), formed either a lineage distinct from Z. quadripartis–Z. hilifer subclade, or a monophyletic but statistically unsupported clade in the ML analyses. Therefore, additional loci (rpb2 and tef1) were sequenced for these three species to complete their dataset. In the final four-gene analyses (Fig. 4), all four species were recovered as a monophyletic clade with moderate statistical support in the ML analysis. Given the variable topology of the Zaanenomyces clade observed in several phylogenetic analyses, ex-type strains of three Zaanenomyces species were cultivated and examined for cultural characteristics and micromorphology. Comparative analysis showed that Z. moderatricis-academiae and Z. versatilis differ from Z. hilifer and Z. quadripartis in several distinct morphological and cultural features. Zaanenomyces moderatricis-academiae and Z. versatilis have much paler, often hyaline and micronematous conidiophores, particularly pronounced in the former species, and their conidia germinate rapidly. In contrast, Z. hilifer and Z. quadripartis produce brown, macronematous conidiophores with a conspicuous, pigmented basal cell. Colony morphology also varies; colonies of Z. moderatricis-academiae and Z. versatilis are uniformly dark from the centre to periphery, whereas Z. quadripartis and Z. hilifer display a more distinct, paler margin and certain zonation. The instability in phylogenetic resolution of the Zaanenomyces clade across analyses suggests that its current circumscription may not adequately capture the full extent of morphological variability among its species. This warrants further investigation through expanded taxon sampling, the inclusion of 67 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum additional molecular loci, and consideration of a possible revision of the generic concept. Correspondingly, our morphological observations indicate that Z. moderatricis-academiae and Z. versatilis form a morphologically cohesive subgroup within Zaanenomyces, distinguishable from the lineage comprising Z. hilifer and Z. quadripartis. This raises the possibility that they may represent a separate, yet closely related taxon. Morphologically, Zaanenomyces is remarkably similar to Camporesiomyces (Hyde et al. 2020), making both genera difficult to distinguish. Camporesiomyces was originally described for two sexually reproducing species and the asexual Ca. vaccinii (syn. Helicoma vaccinii, Carris 1989), which has helicosporous conidia borne on denticulate conidiogenous cells. Han et al. (2025) expanded the genus by adding three species defined solely by asexual characteristics, namely dematiaceous, macronematous conidiophores often in small clusters, holoblastic-denticulate conidiogenous cells, and hyaline to pale brown, septate, straight conidia, thereby broadening the generic concept to include a non-helicosporous conidial shape. The ex-type strain of H. vaccinii (ATCC 66068) is also preserved as CBS 216.90, and the latter is the source of the DNA sequences available in GenBank (Tsui et al. 2006). In this study, we examined CBS 216.90 in vitro, and we confirm that it matches the protologue and produces the characteristic helicosporous conidia. However, helicoid conidial morphology was omitted from the protologue of Camporesiomyces (Hyde et al. 2020) and was not adequately addressed by Han et al. (2025), who suggested that the occurrence of two conidial types within the genus might be the result of geographical isolation in Ca. vaccinii. Camporesiomyces vaccinii is readily distinguished from Helicoma by its conidiophores and conidial morphology. In Ca. vaccinii, conidiophores terminate in an integrated, denticulate conidiogenous cell, and the conidia are coiled but asymmetrical with a distinct basal cell that is elongated, tapering, and truncate at the base. In contrast, Helicoma develops setiform conidiophores with numerous intercalary conidiogenous cells, usually bearing one or two denticles, and produces symmetrical, coiled conidia with cells that often become progressively smaller from the centre toward both ends. The conidiophore architecture of Ca. vaccinii is consistent with other Camporesiomyces species that form straight conidia. Notably, the straight, distinctly truncate basal cell of the otherwise coiled conidium of Ca. vaccinii may reflect an intermediate stage in the morphogenetic diversification of conidial forms within Camporesiomyces, pointing to a broader morphological plasticity in the genus than previously recognised. Conidial morphology within the Tubeufiaceae is highly diverse. The helicoid, septate, hyaline to pale brown conidial type is the most common morphotype that occurs in the majority of genera in the family. In addition to the straight, hyaline, septate conidia of Camporesiomyces, Skoliomycella, and Zaanenomyces, other forms include darkly pigmented dictyoconidia of monodictys-like asexual morphs, described in the life cycles of Chlamydotubeufia (Boonmee et al. 2011), Dictyospora (Brahmanage et al. 2017), Manoharachariella (Bagyanarayana et al. 2009), Muripulchra (Luo et al. 2017), Neochlamydotubeufia (Lu et al. 2018a), Neodictyospora (Zhang et al. 2023b), and Tubeufia amazonensis (Samuels et al. 1987). Whereas the helicosporous and straight conidia are produced on holoblastic-denticulate conidiogenous cells, the dictyoconidia arise from holoblastic conidiogenous cells with a flat, non-denticulate locus. 68 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Tubeufia amazonensis also produces a pycnidial, asteromella-like synasexual morph (synanamorph). The systematic placement of Tu. amazonensis remains uncertain. Based on ascomatal wall characters, Crane et al. (1998) transferred this species to Thaxteriella, a genus characterised by helicosporous conidia. However, no molecular data are currently available for Thaxteriella spp., leaving its phylogenetic position unresolved. Conclusions This study highlights the importance of routine molecular and morphological verification of strains obtained even from well-curated culture collections. The case of Pleurophragmium parvisporum demonstrates how subtle morphological characters, if misinterpreted, can obscure true phylogenetic relationships. Our findings indicate that historical misidentifications persist and may perpetuate taxonomic inaccuracies if left unverified. Phylogenetic analyses revealed that seven strains deposited under the name P. parvisporum represent a polyphyletic assemblage distributed across four families and/or orders in three classes, namely in the Dothideomycetes (Tubeufiales), Eurotiomycetes (Chaetothyriales), and Sordariomycetes (Myrmecridiales and Papulosaceae). Only one strain, CBS 770.83, conforms to the species concept of P. parvisporum, confirming its placement within the Myrmecridiales. These results led to the synonymisation of Neomyrmecridium with Pleurophragmium and the proposal of one new genus, several new species, new names and combinations. Acknowledgements We sincerely thank Gary J. Samuels for reading the manuscript and for his valuable comments and suggestions. We thank the culture collection manager Gerard Verkleij (CBS) for availability of the studied strains, and culture and collection curators Trix Merkx (CBS) and Dana Lančová (PRA) for their invaluable assistance in obtaining living cultures and facilitating the deposition of new herbarium specimens and dried strains. We are grateful to Kazuaki Tanaka for checking taxa and their GenBank Accessions used in this study. 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. 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). 69 IMA Fungus 16: e173033 (2025), DOI: 10.3897/imafungus.16.173033 Martina Réblová et al.: Pleurophragmium parvisporum Funding This study was supported by the project of the Czech Academy of Sciences ‘Strategie AV21 MycoLife – svět hub’ (M.R.), and long-term research development projects, no. RVO 67985939 of the Czech Academy of Sciences (M.R., L.B.) and MH CZ – DRO (UHHK, 00179906) of the University Hospital Hradec Králové (J.N.). Author contributions Conceptualization: MR, MHR. Formal analysis: JN, MR, MHR, LB. Funding acquisition: MR. Investigation: JN, MR, LB, MHR. Methodology: JN, MR, MHR. Project administration: MR. Resources: MR. Supervision: MR. Validation: MR. Writing - original draft: MR, MHR. Writing - review and editing: JN, MHR, MR. 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Link: https://doi.org/10.3897/imafungus.16.173033.suppl1 Supplementary material 2 Biogeographical distribution, sample type, habitat and other detailed metadata for newly described species inferred from the GlobalFungi database Authors: Martina Réblová Data type: xlsx Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/imafungus.16.173033.suppl2