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Research Article
Two novel Helicosporium species (Tubeufiaceae, Tubeufiales) from southern China based on morphological and molecular evidence
expand article infoSong Bai‡, Fang Wang‡, Su-Ran Wan‡, Xiao-Kang Lv‡, Li-Jun Chen‡, Rong Wu‡, Jian Ma‡§
‡ Guizhou Industry Polytechnic College, Guiyang, China
§ School of Food and Pharmaceutical Engineering, Guizhou Institute of Technology, Guiyang, China

Corresponding author: Li-Jun Chen ( clj102128@163.com )

Corresponding author: Jian Ma ( yanmajian@163.com )

Academic editor: Xin-Cun Wang
© 2025 Song Bai, Fang Wang, Su-Ran Wan, Xiao-Kang Lv, Li-Jun Chen, Rong Wu, Jian Ma.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Bai S, Wang F, Wan S-R, Lv X-K, Chen L-J, Wu R, Ma J (2025) Two novel Helicosporium species (Tubeufiaceae, Tubeufiales) from southern China based on morphological and molecular evidence. MycoKeys 124: 159-176. https://doi.org/10.3897/mycokeys.124.168933
Open Access

Abstract

During a survey of saprobic fungi, fresh specimens were collected from decaying wood in terrestrial habitats in Guizhou and Hainan provinces, southern China. Two novel species, Helicosporium qixianlingense and H. tongrenense, are introduced based on phylogenetic analyses of a combined dataset (ITS, LSU, tef1-α, and rpb2) and morphological evidence. Comprehensive descriptions, illustrations, notes, and phylogenetic analyses supporting the taxonomic placement of these new taxa are provided. These findings are significant for exploring the species diversity of Helicosporium in southern China.

Key words:

Dothideomycetes, helicosporous fungi, phylogeny, taxonomy, two novel species

Introduction

Tubeufiales (Pleosporomycetidae, Dothideomycetes) was established by Boonmee et al. (2014), with Tubeufiaceae designated as the type family based on both morphological characteristics and molecular DNA data. Currently, the order includes three families: Bezerromycetaceae, Tubeufiaceae, and Wiesneriomycetaceae, as determined through molecular DNA data and divergence time estimates (Boonmee et al. 2011, 2014, 2021; Liu et al. 2017; Wijayawardene et al. 2022). In the latest comprehensive revision by Lu et al. (2018b), based on morphological comparison and/or molecular data, 13 new genera, 52 new species, 16 new records, and 43 new combinations were added to Tubeufiales. Recent studies also revised the classification of previously misidentified Tubeufiaceae species and emphasized the importance of considering all morphological differences when identifying helicosporous asexual species (Lu et al. 2017a,b, 2018a, b, 2022a, 2023a, b; Lu and Kang 2020).

Tubeufiaceae, typified by Tubeufia (Barr 1979), is the largest family within Tubeufiales (Wijayawardene et al. 2022; Ma et al. 2024b), encompassing 53 accepted genera, namely Acanthohelicospora, Acanthophiobolus, Acanthostigma, Acanthostigmina, Acanthotubeufia, Acrogenihelicosporium, Aquaphila, Berkleasmium, Bifrontia, Boerlagiomyces, Camporesiomyces, Chaetosphaerulina, Chlamydotubeufia, Dematiohelicoma, Dematiohelicomyces, Dematiohelicosporum, Dematiotubeufia, Dictyospora, Discotubeufia, Excipulariopsis, Helicangiospora, Helicoarctatus, Helicodochium, Helicohyalinum, Helicoma, Helicomyces, Helicosporium, Helicotruncatum, Helicotubeufia, Hyalohelicoon, Hyalohelicotubeufia, Hyalohelisphora, Kamalomyces, Kevinhydea, Lichenotubeufia, Manoharachariella, Muripulchra, Neoacanthostigma, Neochlamydotubeufia, Neohelicoma, Neohelicomyces, Neohelicosporium, Neomanoharachariella, Neotubeufia, Parahelicomyces, Pleurohelicosporium, Pseudohelicoon, Pseudohelicosporium, Pseudotubeufia, Tamhinispora, Thaxteriella, Tubeufia, and Zaanenomyces (Boonmee et al. 2011, 2014; Hyde et al. 2016, 2017, 2024; Luo et al. 2017; Lu et al. 2018b; Tibpromma et al. 2018; Liu et al. 2019; Crous et al. 2021, 2023; Ma et al. 2023a, b, 2024b). Among them, Camporesiomyces and Tubeufia exhibit diverse asexual conidial morphology (Carris 1989; Matsushima 1993; Zhao et al. 2007; Hyde et al. 2020; Han et al. 2025).

Based on morphological evidence, the genus Helicosporium was established by Nees (1817), with H. vegetum as the type species. Currently, 30 species are accepted within the genus Helicosporium, occurring in freshwater and/or terrestrial habitats across diverse regions, including Austria, Belgium, Britain, British Guiana, Canada, China, Cuba, England, France, Germany, India, Thailand, and the USA (Nees 1817; Linder 1929; Samuels and Muller 1979; Barr 1980; Goos 1989; Morgan-Jones and Goos 1992; Spatafora et al. 2006; Tsui et al. 2006; Zhao et al. 2007; Boonmee et al. 2014, 2021; Lu et al. 2017a, 2018b, 2022a; Dong et al. 2020; Hsieh et al. 2021; Xiao et al. 2023; Ma et al. 2024b; Peng et al. 2025; Sun et al. 2025). Based on molecular DNA data, Helicosporium species exhibit similar morphological characteristics in both their sexual and asexual morphs (Boonmee et al. 2014, 2021; Lu et al. 2017a, 2018b, 2022a; Dong et al. 2020; Hsieh et al. 2021; Xiao et al. 2023; Ma et al. 2024b; Peng et al. 2025; Sun et al. 2025). For example, the sexual morph is characterized by solitary, greenish, reddish-yellow to brownish-yellow ascomata; cylindric-clavate, eight-spored bitunicate asci; and hyaline to yellowish-brown, elongate-fusiform ascospores (Boonmee et al. 2014, 2021; Brahmanage et al. 2017; Lu et al. 2022a; Peng et al. 2025; Sun et al. 2025). The asexual morph is characterized by pale yellow to yellow-green colonies, erect, setiferous conidiophores, and helicoid, hyaline to yellow-green conidia (Linder 1929; Zhao et al. 2007; Boonmee et al. 2014, 2021; Lu et al. 2017a, 2018b, 2022a; Dong et al. 2020; Hsieh et al. 2021; Xiao et al. 2023; Ma et al. 2024b).

In this study, four newly obtained fungal isolates, representing two distinct Helicosporium taxa within the family Tubeufiaceae (Tubeufiales, Dothideomycetes), were collected from decaying wood in southern China. Based on morphological comparisons, illustrations, and multigene phylogenetic analyses of combined ITS, LSU, tef1-α, and rpb2 sequence data, two novel species, Helicosporium qixianlingense and H. tongrenense, are introduced in the present study.

Materials and methods

Sample collection, examination, and isolation

Decaying wood was collected from Qixianling Hot Spring National Forest Park, Hainan Province, and Jiangkou County, Guizhou Province, in southern China. Samples were transported to the laboratory in plastic bags, accompanied by collection details such as locality, habitat, and date (Rathnayaka et al. 2024). Microscopic features were examined and photographed using a stereomicroscope (SMZ-168, Nikon, Japan) and an ECLIPSE Ni compound microscope (Nikon, Tokyo, Japan) equipped with a Canon 90D digital camera. Measurements were taken with Tarosoft Image Frame Work software, and photo plates were prepared using Adobe Photoshop CC 2019 (Adobe Systems, USA).

Single-spore isolations were conducted on potato dextrose agar (PDA) plates following the protocols of Chomnunti et al. (2014) and Senanayake et al. (2020). Germinated conidia or ascospores were aseptically transferred to fresh PDA plates. Morphological characteristics of fungal mycelia on PDA, including colony color, hyphal shape, and growth dimensions, were recorded. Dried fungal specimens were deposited in the Herbarium of Guizhou Academy of Agricultural Sciences (Herb. GZAAS), Guiyang, China, and pure cultures were preserved in the Guizhou Culture Collection (GZCC), Guiyang, China. MycoBank numbers were obtained following the procedures described at https://www.mycobank.org/.

DNA extraction, PCR amplification, and sequencing

Fresh fungal mycelia were scraped from PDA-grown colonies and transferred to 1.5-mL microcentrifuge tubes using sterilized lancets for genomic DNA extraction. DNA was extracted with the Biospin Fungus Genomic DNA Extraction Kit (BioFlux, China). The following primer pairs were used for PCR amplification: ITS5/ITS4 for the internal transcribed spacer region (ITS; White et al. 1990), LR0R/LR5 for the large subunit ribosomal RNA gene (LSU; Vilgalys and Hester 1990), EF1-983F/EF1-2218R for translation elongation factor 1-α (tef1-α; Rehner and Buckley 2005), and fRPB2-5F/fRPB2-7cR for RNA polymerase II second largest subunit (rpb2; Liu et al. 1999). Each 50-μL PCR reaction contained 2 μL of template DNA, 2 μL of each primer, and 44 μL of 1.1× T3 Super PCR Mix (Qingke Biotech, Chongqing, China). Polymerase chain reaction (PCR) was performed using the cycling conditions described by Ma et al. (2024a). Amplification products were purified and sequenced using the same primers at Beijing Tsingke Biotechnology Co., Ltd.

Phylogenetic analyses

The newly obtained sequences were quality-checked and assembled using BioEdit v.7.0.5.3 (Hall 1999) and SeqMan v.7.0.0 (DNASTAR, Madison, WI, USA; Swindell and Plasterer 1997). The sequences used in this study were retrieved from GenBank (Table 1; https://www.ncbi.nlm.nih.gov/). Sequence matrices for each gene were aligned using MAFFT v.7.473 (Katoh et al. 2019; https://mafft.cbrc.jp/alignment/server/). Alignments were trimmed with trimAl v.1.2rev59 (Capella-Gutiérrez et al. 2009) and concatenated using SequenceMatrix v.1.7.8 (Vaidya et al. 2011).

Table 1.
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Taxa used in this study and their GenBank accession numbers.

Taxon Strain GenBank Accessions
ITS LSU tef1-α rpb2
Acanthohelicospora aurea GZCC 16-0060 KY321323 KY321326 KY792600 MF589911
Acanthostigma chiangmaiensis MFLUCC 10-0125T JN865209 JN865197 KF301560 N/A
Acanthostigma perpusillum UAMH 7237 AY916492 AY856892 N/A N/A
Berkleasmium aquaticum MFLUCC 17-0049T KY790444 KY790432 KY792608 MF535268
Berkleasmium fusiforme MFLUCC 17-1978T MH558693 MH558820 MH550884 MH551007
Boerlagiomyces macrospora MFLUCC 12-0388 KU144927 KU764712 KU872750 N/A
Botryosphaeria agaves MFLUCC 10-0051 JX646790 JX646807 N/A N/A
Botryosphaeria dothidea CBS 115476 KF766151 DQ678051 DQ767637 DQ677944
Camporesiomyces bhatii GMBCC 1120T PQ763360 PQ842543 PV388894 PV388888
Chlamydotubeufia cylindrica MFLUCC 16-1130T MH558702 MH558830 MH550893 MH551018
Chlamydotubeufia huaikangplaensis MFLUCC10-0926T JN865210 JN865198 N/A N/A
Dematiohelicomyces helicosporus MFLUCC 16-0213T KX454169 KX454170 KY117035 MF535258
Dematiohelicosporum guttulatum MFLUCC 17-2011T MH558705 MH558833 MH550896 MH551021
Dematiotubeufia chiangraiensis MFLUCC 10-0115T JN865200 JN865188 KF301551 N/A
Helicangiospora lignicola MFLUCC 11-0378T KF301523 KF301531 KF301552 N/A
Helicoarctatus aquaticus MFLUCC 17-1996T MH558707 MH558835 MH550898 MH551024
Helicohyalinum aquaticum MFLUCC 16-1131T KY873625 KY873620 KY873284 MF535257
Helicohyalinum infundibulum MFLUCC 16-1133T MH558712 MH558840 MH550903 MH551029
Helicoma guttulatum MFLUCC 16-0022T KX454171 KX454172 MF535254 MH551032
Helicoma hongkongense MFLUCC 17-2005 MH558716 MH558843 MH550907 MH551033
Helicosporium acropleurogenum CGMCC 3.25563T PP626574 PP639430 PP596333 PP596460
Helicosporium aquaticum MFLUCC 17-2008T MH558733 MH558859 MH550924 MH551049
Helicosporium brunneisporum CGMCC 3.25542T PP626577 PP639433 PP596336 PP596463
Helicosporium changjiangense GZCC 22-2113T PP626578 PP639434 PP596337 PP596464
Helicosporium flavisporum MFLUCC 17-2020T MH558734 MH558860 MH550925 MH551050
Helicosporium flavum MFLUCC 16-1230T KY873626 KY873621 KY873285 N/A
Helicosporium hainanense GZAAS 22-2006T OP508730 OP508770 OP698081 OP698070
Helicosporium jiangkouense HKAS 128933T PP626580 PP639436 PP596339 PP596466
Helicosporium latisporum HKAS 128960T PP626582 PP639437 PP596340 PP596467
Helicosporium liuzhouense GZCC 22-2014T OQ981394 OQ981402 OQ980476 OQ980474
Helicosporium luteosporum MFLUCC 16-0226T KY321324 KY321327 KY792601 MH551056
Helicosporium multidentatum GZCC 22-2013T OQ981395 OQ981403 OQ980477 OQ980475
Helicosporium multiseptatum GUCC 24-0090T PQ570843 PQ570860 PQ761135 N/A
Helicosporium nanningense GZCC 22-2175T OR066418 OR066425 OR058864 OR058857
Helicosporium qixianlingense GZCC 25-0641T PX111181 PX111188 PX102605 PX102599
Helicosporium qixianlingense GZCC 25-0642 PX111182 PX111189 PX102606 PX102600
Helicosporium ramosiphorum CGMCC 3.25541T PP626576 PP639432 PP596335 PP596462
Helicosporium rubrum MFLUCC 24-0090T PQ098477 PQ098514 PQ490681 PQ490675
Helicosporium setiferum MFLUCC 17-1994T MH558735 MH558861 MH550926 MH551051
Helicosporium sexuale MFLUCC 16-1244T MZ538503 MZ538537 MZ567082 MZ567111
Helicosporium sp. NBRC 9014 AY916489 AY856903 N/A N/A
Helicosporium sp. Z17 PX220122 PX220124 N/A N/A
Helicosporium thailandense MFLUCC 18-1407T MT627698 MN913718 MT954371 N/A
Helicosporium tongrenense GZCC 23-0026T PQ098476 PQ098513 PX102603 PX102597
Helicosporium tongrenense GZCC 25-0640 PX111180 PX111187 PX102604 PX102598
Helicosporium vegetum GZCC 23-0060 PP626584 PP639439 PP596342 PP596469
Helicosporium vesicarium MFLUCC 17-1795T MH558739 MH558864 MH550930 MH551055
Helicosporium viridiflavum MFLUCC 17-2336T MH558738 N/A MH550929 MH551054
Helicosporium viridisporum GZCC 22-2008T OP508736 OP508776 OP698087 OP698076
Helicotubeufia hydei MFLUCC 17-1980T MH290021 MH290026 MH290031 MH290036
Helicotubeufia jonesii MFLUCC 17-0043T MH290020 MH290025 MH290030 MH290035
Muripulchra aquatica MFLUCC 15-0249T KY320532 KY320549 N/A N/A
Neoacanthostigma fusiforme MFLUCC 11-0510T KF301529 KF301537 N/A N/A
Neochlamydotubeufia fusiformis MFLUCC 16-0016T MH558740 MH558865 MH550931 MH551059
Neohelicomyces acropleurogenus CGMCC 3.25549T PP626594 PP639450 PP596351 PP596478
Neohelicomyces aquaticus MFLUCC 16-0993T KY320528 KY320545 KY320561 MH551066
Neohelicosporium acrogenisporum MFLUCC 17-2019T MH558746 MH558871 MH550937 MH551069
Neohelicosporium aquaticum MFLUCC 17-1519T MF467916 MF467929 MF535242 MF535272
Neomanoharachariella xizangensis KUNCC 23-15799T OR803724 OR803722 OR813978 OR813975
Parahelicomyces quercus MFUCC 17-0895T MK347720 MK347934 MK360077 MK434906
Parahelicomyces talbotii MFLUCC 17-2021T MH558765 MH558890 MH550957 MH551091
Tubeufia guttulata GZCC 23-0404T OR030841 OR030834 OR046678 OR046684
Tubeufia hainanensis GZCC 22-2015T OR030842 OR030835 OR046679 OR046685

Note: Newly generated sequences are in bold. The superscript “T” indicates ex-type strains. “N/A” indicates unavailable data in GenBank.

Maximum likelihood (ML) analysis was conducted using the IQ-TREE web server (http://iqtree.cibiv.univie.ac.at/) with the best-fit substitution model automatically selected based on the Bayesian Information Criterion (BIC) (Nguyen et al. 2015). Bayesian inference (BI) analysis was performed using MrBayes on XSEDE (v.3.2.7a) via CIPRES (Stamatakis 2014). The aligned FASTA file was converted to NEXUS format using AliView (Daniel et al. 2010). The optimal substitution model for each dataset was selected with MrModeltest v.2.3 (Nylander et al. 2008). Posterior probabilities (BYPP) were determined based on Bayesian Markov chain Monte Carlo (BMCMC) sampling (Huelsenbeck and Ronquist 2001). Four simultaneous Markov chains were run for 10,000,000 generations, and trees were sampled every 1,000th generation. The burn-in phase was set at 25%, and the remaining trees were used for calculating posterior probabilities (BYPP).

Phylogenetic trees were visualized using FigTree v.1.4.4 and subsequently edited with Adobe Illustrator CC 2019 (v.23.1.0; Adobe Systems, USA).

Phylogenetic results

The phylogenetic positions of the four novel strains were assessed using a multilocus phylogenetic approach. The concatenated sequence matrix comprised 3,346 nucleotide positions (ITS: 1–539, LSU: 540–1389, tef1-α: 1390–2301, and rpb2: 2302–3346), incorporating 61 ingroup taxa and two outgroup taxa, Botryosphaeria agaves (MFLUCC 10–0051) and B. dothidea (CBS 115476). Both maximum likelihood (ML) and Bayesian inference (BI) analyses of the combined ITS, LSU, tef1-α, and rpb2 datasets yielded congruent tree topologies. The best-scoring ML tree (Fig. 1) exhibited a final log-likelihood value of –34,578.071.

Figure 1. 

Phylogenetic tree generated using RAxML analysis based on the concatenated ITS, LSU, tef1-α, and rpb2 sequence data. Bootstrap support values (MLBS) ≥ 75% and Bayesian posterior probabilities (BYPP) ≥ 0.95 are indicated near the nodes as MLBS/BYPP, respectively. A hyphen (–) denotes support values below 75% for ML and posterior probabilities below 0.95 for BI. Botryosphaeria agaves (MFLUCC 10–0051) and B. dothidea (CBS 115476) were used as outgroups. Ex-type strains are marked with “T,” and newly obtained isolates are highlighted in bold black font.

Phylogenetic analyses of the phylogram (Fig. 1) revealed that our collections include two new species of Helicosporium within the Tubeufiaceae (Tubeufiales, Dothideomycetes). Isolates GZCC 25–0641 and GZCC 25–0642 formed a sister clade to Helicosporium hainanense (GZAAS 22–2006), with robust support of 93% MLBS and 1.00 BYPP. Furthermore, GZCC 23–0026 and GZCC 25–0640 clustered together, and this clade was sister to H. jiangkouense (HKAS 128933), with 100% MLBS and 1.00 BYPP support.

Taxonomy

Helicosporium qixianlingense S. Bai, L.J. Chen, & J. Ma, sp. nov.

MycoBank No: 904205
Fig. 2

Etymology.

‘‘qixianlingense” refers to the place ‘‘Qixianling Hot Spring National Forest Park,” from where the holotype was collected.

Holotype.

GZAAS 25–0671

Description.

Saprobic on decaying wood in a terrestrial habitat. Sexual morph Undetermined. Asexual morph Hyphomycetous, helicosporous. Colonies on natural substrate superficial, effuse, gregarious, yellowish green to brown. Mycelium partly immersed, partly superficial, composed of pale brown to brown, branched, septate, guttulate, smooth hyphae. Conidiophores 132–144 × 3.7–4.8 μm (x̄ = 139 × 5.5 μm, n = 30), macronematous, mononematous, erect, cylindrical, long, straight or slightly flexuous, simple, septate, brown to dark brown at base, paler towards the apex, smooth, thick-walled. Conidiogenous cells 4.5–10 × 3–4.4 μm (x̄ = 8 × 3.6 μm, n = 30), holoblastic, monoblastic, or polyblastic, integrated, intercalary, cylindrical, denticulate, arising laterally from the lower portion of conidiophores as tiny bladder-like protrusions (3–4 µm long, 2–2.7 µm wide), with each bearing 1–2 tiny sporogenous conidiogenous loci, hyaline to pale brown, smooth. Conidia solitary, pleurogenous, helicoid, tapering towards the rounded ends, developing on bladder-like protrusion, 11.7–12 μm in diameter, and conidial filament 1.2–2.1 μm wide (x̄ = 11.9 × 1.7 μm, n = 30), 72.5–84 μm long (x̄ = 76.5 μm, n = 35), up to 41/2 times, becoming loosely coiled when the conidia are young and not becoming loose when mature in water, indistinctly multi-septate, guttules, hyaline, smooth-walled.

Figure 2. 

Helicosporium qixianlingense (GZAAS 25–0671, holotype). a, b. Colonies on the host surface; c–f. Conidiophores and conidiogenous cells; g–l. Conidiogenous cells; m–q. Conidia; r. A germinated conidium; s, t. Colonies on PDA from above and below after 44 days of incubation at room temperature. Scale bars: 30 μm (c–f); 10 μm (r); 5 μm (g–q).

Culture characteristics.

Conidia germinating on PDA within 11 hours, producing germ tubes from the conidial body. Colony on PDA reaching 3.2 cm in diameter after 44 days at room temperature (approximately 25 °C), circular or irregular, umbonate, with an undulate margin, and pale brown to light pinkish in color.

Material examined.

• China, Hainan Province, Baoting Li and Miao Autonomous County, Qixianling Hot Spring National Forest Park, on decaying wood in a terrestrial habitat, 2 November 2024, Jian Ma Q23 (GZAAS 25–0671, holotype), ex-type living cultures GZCC 25–0641; Ibid., Q39 (GZAAS 25–0672, paratype), living culture GZCC 25–0642.

Notes.

In the phylogenetic analyses (Fig. 1), our isolates (GZCC 25–0641 and GZCC 25–0642) formed a sister clade to Helicosporium hainanense (GZAAS 22–2006) with 93% MLBS and 1.00 BYPP support. However, nucleotide polymorphisms in the ITS, LSU, tef1-α, and rpb2 sequence data between GZCC 25–0641 and H. hainanense (GZCC 22–2006) revealed nucleotide differences of 11/432 bp (2.5%, with no gaps), 5/814 bp (0.6%, with no gaps), 11/906 bp (1.2%, with no gaps), and 39/1,045 bp (3.7%, with no gaps), respectively. Furthermore, Helicosporium qixianlingense (GZAAS 25–0671) differs from H. hainanense (GZAAS 22–2006, ex-type) by its longer conidia (72.5–84 μm vs. 55–60 μm), narrower conidial filament (1.2–2.1 μm vs. 2–3 μm), and more coiled times (up to 41/2 times vs. 21/4–23/4 μm times) (Lu et al. 2022a). Therefore, based on both molecular and morphological evidence, we introduce Helicosporium qixianlingense as a novel species.

Helicosporium tongrenense S. Bai, L.J. Chen, & J. Ma, sp. nov.

MycoBank No: 904206
Fig. 3

Etymology.

‘‘tongrenense” refers to the place ‘‘Tongren City,” from where the holotype was collected.

Holotype.

HKAS 128925

Description.

Saprobic on decaying wood in a terrestrial habitat. Asexual morph Undetermined. Sexual morph: Ascomata superficial, seated on a subiculum, solitary, scattered, globose to subglobose, bright reddish yellow to brown yellow, with central narrow ostiole, no observed setae. Peridium composed of several layers of brown to dark brown cells of textura angularis, outer layer brown cells, and inner layer pale brown to hyaline cells. Hamathecium comprising numerous, filiform, septate, branched, hyaline pseudoparaphyses. Asci 90.5–143 × 11.5–14.5 µm (x̄ = 107.5 × 13 μm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical to clavate, short-pedicellate, apically rounded, basally flexious. Ascospores 37–48 × 3.5–5.5 µm (x̄ = 43 × 4 μm, n = 20), overlapping 2–3 seriate, elongate-fusiform, tapering towards narrow, widest at the central part, subacute ends, straight to slightly curved, guttules, multi-septate, hyaline, smooth-walled.

Figure 3. 

Helicosporium tongrenense (HKAS 128925, holotype). a. Ascomata on host surface; b–d. Vertical sections of ascomata and peridium; e–h. Asci; i–n. Ascospores; o, p. Colonies on PDA from above and below after 3 months of incubation at room temperature. Scale bars: 50 µm (b–e), 20 µm (f–h), 10 µm (i–n).

Culture characteristics.

Ascospores germinating on PDA, producing germ tubes within 19 hours. Colony on PDA reaching 0.7 cm diameter after 90 days at room temperature (approximately 25 °C), circular or irregular, flat, with an entire margin, and pale brown to yellowish brown in color.

Material examined.

• China, Guizhou Province, Tongren City, Jiangkou County, on decaying wood in a terrestrial habitat, 20 May 2022, Xia Tang, JK1.1 (HKAS 128925 = GZAAS 23–0027, holotype), ex-type living cultures GZCC 23–0026; Ibid., JK4 (GZAAS 25–0670, paratype), living culture GZCC 25–0640.

Notes.

Morphologically, Helicosporium tongrenense (HKAS 128925) resembles H. flavum (MFLU 17–0704) in having solitary, scattered, globose to subglobose, bright reddish-yellow to brown-yellow ascomata; 8-spored, bitunicate, fissitunicate, cylindrical to clavate asci; and elongate-fusiform, straight to slightly curved, multi-septate, hyaline ascospores (Brahmanage et al. 2017). However, H. tongrenense (HKAS 128925) differs from H. flavum (MFLU 17–0704) by shorter ascospores (37–48 μm vs. up to 60 μm) (Brahmanage et al. 2017). Additionally, H. flavum possesses short, setae-like projections, which are absent in H. tongrenense (Brahmanage et al. 2017). Phylogenetically, our isolates (GZCC 23–0026 and GZCC 25–0640) form a sister clade to H. jiangkouense (HKAS 128933), with 100% MLBS and 1.00 BYPP support. However, comparison of the ITS, LSU, tef1-α, and rpb2 sequence data between H. tongrenense (GZCC 23–0026) and H. jiangkouense (HKAS 128933) revealed nucleotide base differences of 26/526 bp (4.9%, including 1 gap), 51/834 bp (6.1%, including 3 gaps), 1/884 bp (0.1%, including 1 gap), and 1/916 bp (0.1%, with no gaps), respectively. Notably, the asexual morph of H. jiangkouense is the only form observed to date (Ma et al. 2024b). Therefore, based on both multigene phylogenetic analyses and morphological differences, we introduce Helicosporium tongrenense as a novel species.

Discussion

Based on molecular data and/or morphological characteristics, the genus Helicosporium currently comprises 32 species, including our new species, H. qixianlingense and H. tongrenense (Hsieh et al. 2021; Xiao et al. 2023; Ma et al. 2024b; Peng et al. 2025; Sun et al. 2025). Among the known Helicosporium species, 26 are reported exclusively from the asexual morph, three (H. multiseptatum, H. rubrum, and H. tongrenense) solely from the sexual morph, and three (H. flavum, H. sexuale, and H. vegetum) have been documented in both sexual and asexual morphs (Nees 1817; Linder 1929; Samuels and Muller 1979; Barr 1980; Goos 1989; Morgan-Jones and Goos 1992; Spatafora et al. 2006; Tsui et al. 2006; Zhao et al. 2007; Boonmee et al. 2014, 2021; Lu et al. 2017a, 2018b, 2022a; Dong et al. 2020; Hsieh et al. 2021; Xiao et al. 2023; Ma et al. 2024b; Peng et al. 2025; Sun et al. 2025). Seven species—Helicosporium albidum, H. casuarinae, H. decumbens, H. favidum, H. melghatianum, H. murinum, and H. neesii—are currently known only from morphological descriptions but lack molecular data (Grove 1886; Linder 1929; Moore 1957; Matsushima 1985; Goos 1989; Zhao et al. 2007; Dharkar et al. 2010; Lu et al. 2018b; Hsieh et al. 2021; Ma et al. 2024b).

Previous studies have reported that secondary metabolites of Helicosporium species exhibit antibacterial activity against both bacteria and fungi (Kim et al. 2003; Lee et al. 2006, 2013; Choi et al. 2012). Our strains, GZCC 23–0026 and GZCC 25–0640, were successfully isolated using the single-spore isolation method of Lu et al. (2022b); however, the colonies exhibited slow growth, reaching only 0.7 cm in diameter after 90 days of incubation. Similarly, Ma et al. (2024b) employed the same method for two newly described asexual Helicosporium species, but the cultures yielded only sufficient material for DNA extraction. The alternative method of crushing the substrate and incorporating it into PDA medium to mimic natural growth conditions has proven useful for cultivating helicosporous fungi lacking available strains, as well as species that are difficult to grow on artificial media (Lu et al. 2022b; Ma et al. 2024b). Nonetheless, further optimization of culture media is necessary to accelerate mycelial growth in helicosporous fungi—thereby improving the acquisition of viable strain resources.

The asexual morphs of Helicosporium exhibit considerable variation in conidial filament width (Linder 1929; Boonmee et al. 2014, 2021; Lu et al. 2017a, 2018b, 2022a; Dong et al. 2020; Hsieh et al. 2021; Xiao et al. 2023; Ma et al. 2024b). For example, the conidial filaments of H. setiferum (GZAAS 23–0154) measure 1.5–2.5 μm in width, those of H. flavum (GZAAS 23–0491) are 3–4 μm wide, and those of H. jiangkouense (HKAS 128933) range from 4.5–6.5 μm (Ma et al. 2024b). Moreover, we observed that conidia with wider filaments maintain a tightly coiled structure in water, whereas those with narrower filaments tend to be loosely coiled or uncoiled (Linder 1929; Boonmee et al. 2014, 2021; Lu et al. 2017a, c, 2018b, 2022a; Dong et al. 2020; Hsieh et al. 2021; Xiao et al. 2023; Ma et al. 2024b). These observations suggest that the width of the conidial filaments is a key factor influencing whether Helicosporium conidia remain tightly coiled, become loosely coiled, or uncoil in water.

Acknowledgments

We thank Shaun Pennycook (Manaaki Whenua Landcare Research, New Zealand) for his valuable suggestions on fungal nomenclature.

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.

Funding

This work was supported by the Guizhou Industry Polytechnic College Faculty-level Research Project (Grant No. 2024ZK18), the Science and Technology Planning Project of Guizhou Province (Grant No. Qian Ke He Ji Chu ZK [2022] Zhong Dian 025), and the High-Level Talent Initial Funding of Guizhou Industry Polytechnic College (Grant No. 2023-RC-01).

Author contributions

Morphological data, photo-plates and phylogenetic analyzes were completed by Song Bai and Jian Ma. The original draft was written by Song Bai and Jian Ma, and Fang Wang, Su-Ran Wan , Xiao-Kang Lv, Li-Jun Chen , Rong Wu revised the paper.

Author ORCIDs

Song Bai https://orcid.org/0000-0002-1972-2834

Fang Wang https://orcid.org/0000-0002-3341-6788

Su-Ran Wan https://orcid.org/0009-0000-1164-6921

Xiao-Kang Lv https://orcid.org/0009-0006-7254-8747

Li-Jun Chen https://orcid.org/0009-0004-8562-875X

Rong Wu https://orcid.org/0000-0002-4946-8806

Jian Ma https://orcid.org/0009-0008-1291-640X

Data availability

All of the data that support the findings of this study are available in the main text.

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