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Research Article
Novel Helicosporium and Neohelicomyces (Tubeufiaceae, Tubeufiales) species from terrestrial habitats in China and Thailand
expand article infoTao Peng‡, Yong-Zhong Lu§, Song Bai|, Jing-Yi Zhang§, Xing-Juan Xiao§, Na Wu#, Jian Ma§|
‡ Department of Brewing Engineering, Moutai Institute, Renhuai, China
§ Guizhou Institute of Technology, Guiyang, China
| Guizhou Industry Polytechnic College, Guiyang, China
¶ Mae Fah Luang University, Chiang Rai, Thailand
# University of Electronic Science and Technology of China, Chengdu, China

Corresponding author: Song Bai ( basonmail@163.com )

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

Academic editor: Samantha C. Karunarathna
© 2025 Tao Peng, Yong-Zhong Lu, Song Bai, Jing-Yi Zhang, Xing-Juan Xiao, Na 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: Peng T, Lu Y-Z, Bai S, Zhang J-Y, Xiao X-J, Wu N, Ma J (2025) Novel Helicosporium and Neohelicomyces (Tubeufiaceae, Tubeufiales) species from terrestrial habitats in China and Thailand. MycoKeys 112: 81-101. https://doi.org/10.3897/mycokeys.112.140211
Open Access

Abstract

During our investigations of saprobic fungi, five fungal collections from terrestrial habitats in China and Thailand were examined using both morphological and multi-gene phylogenetic approaches (LSU, ITS, tef1-α, and rpb2), resulting in the identification of three novel species: Helicosporium rubrum, Neohelicomyces maolanensis, and N. subtropicus. Helicosporium and Neohelicomyces are morphologically similar in their asexual morphs but can be distinguished based on their molecular phylogenetic data. In this study, our new species, Helicosporium rubrum, represents the fourth sexual species within the genus, characterized by yellow-brown ascomata and fusiform ascospores. Detailed descriptions, illustrations, phylogenetic analysis results, and corresponding notes are provided to clarify the distinctions between these new species and related taxa.

Key words

Helicosporous fungi, phylogeny, taxonomy, terrestrial habitats

Introduction

Helicosporium was introduced by Nees (1817), with H. vegetum as the type species. Currently, 108 species are listed in Index Fungorum (2024); however, based on morphological data comparison and molecular data, only 28 are accepted within the genus Helicosporium (Lu et al. 2018b; Xiao et al. 2023; Ma et al. 2024b). Of these, 27 species have been found in freshwater and terrestrial habitats in various regions, including Austria, Belgium, Britain, Canada, China, Cuba, India, Thailand, and the USA (Lu et al. 2017a, 2018b, 2022; Dong et al. 2020; Boonmee et al. 2021; Hsieh et al. 2021; Xiao et al. 2023; Ma et al. 2024b). The remaining species, H. vegetum, has been found widely distributed in terrestrial habitats globally (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; Lu et al. 2018b; Ma et al. 2024b). So far, only H. flavum, H. sexuale, and H. vegetum have been described as sexual morphs in this genus, all of which have been confirmed through DNA sequence data (Boonmee et al. 2014, 2021; Brahmanage et al. 2017). These sexual morphs are characterized by solitary, yellowish-brown to greenish ascomata, cylindric-clavate, eight-spored bitunicate asci, and hyaline to yellowish-brown elongate-fusiform ascospores (Ma et al. 2024b). In contrast, their asexual morphs form pale yellow to yellow-green colonies on natural woody substrates, with erect, setiferous conidiophores and helicoid, hyaline to yellow-green conidia (Xiao et al. 2023; Ma et al. 2024b).

Neohelicomyces was established by Luo et al. (2017), with N. aquaticus as the type species, based on molecular and morphological data. The genus now includes 25 species; all are asexual morphs supported by molecular evidence. No sexual stages have been reported for this genus (Ma et al. 2024a, 2024b). Neohelicomyces species occupy both freshwater and terrestrial habitats and have a worldwide distribution and have been reported from China, Czech Republic, Germany, Italy, Japan, Netherlands, Thailand, and the USA (Luo et al. 2017; Lu et al. 2018b, 2022; Crous et al. 2019a, 2019b; Dong et al. 2020; Hsieh et al. 2021; Yang et al. 2023; Ma et al. 2024a, 2024b). Neohelicomyces species are characterized by macronematous, mononematous, erect, septate conidiophores with holoblastic conidiogenous cells and tightly or loosely coiled helicoid conidia (Hsieh et al. 2021; Lu et al. 2022; Yang et al. 2023; Ma et al. 2024a, 2024b).

Previous studies have highlighted the potential of Helicosporium and Neohelicomyces species to produce secondary metabolites with bioactive properties (Kim et al. 2003; Lee et al. 2006, 2013; Choi et al. 2012; Zheng et al. 2023). For example, Helicosporium nizamabadense has demonstrated inhibitory effects against several agricultural pathogenic fungi, including Botrytis cinerea, Fusarium oxysporum, Phytophthora drechsleri, and Rhizoctonia solani (Kim et al. 2003; Lee et al. 2006, 2013). Additionally, 2-methylresorcinol, isolated from Helicosporium sp., exhibited antimicrobial activities against bacteria and fungi (Choi et al. 2012). More recently, two alkaloid compounds from Neohelicomyces hyalosporus were shown to have moderate cytotoxic effects on human cancer cells (Zheng et al. 2023). Therefore, the metabolites of Helicosporium and Neohelicomyces species could serve as promising sources for developing drugs to prevent and manage human tumors.

This study collected five saprophytic fungi from terrestrial habitats in Chiang Mai, Thailand, and Guizhou Province, China. Based on detailed morphological comparison and multigene phylogenetic analyses (LSU, ITS, tef1-α, and rpb2), three novel species, viz. Helicosporium rubrum, Neohelicomyces maolanensis, and N. subtropicus, were isolated and identified.

Materials and methods

Sample collection, examination, and isolation

Fresh specimens were collected from terrestrial habitats in China and Thailand between September 2020 and April 2022, and important collection information in situ was noted as per Rathnayaka et al. (2024). Fungal colonies and micro-morphological structures on the surface of natural substrates were observed using a stereomicroscope (SMZ-168, Nikon, Japan) and photographed with an ECLIPSE Ni compound microscope (Nikon, Tokyo, Japan) equipped with a Canon 90D digital camera.

Single spore isolations were performed following the method outlined by Chomnunti et al. (2014). Germinating spores were aseptically transferred to fresh potato dextrose agar (PDA) plates, according to the method described by Senanayake et al. (2020). Colony characteristics on PDA, such as shape, color, size, margin, and elevation, were monitored and recorded.

Material deposition

Dried materials were deposited in the Herbarium of Mae Fah Luang University (Herb. MFLU), Chiang Rai, Thailand, and the Herbarium of Guizhou Academy of Agriculture Sciences (Herb. GZAAS), Guiyang, China. Cultures were deposited at Mae Fah Luang University Culture Collection (MFLUCC), Chiang Rai, Thailand, and the Guizhou Culture Collection (GZCC), Guiyang, China. The Faces of Fungi numbers were obtained following the guidelines outlined by Jayasiri et al. (2015). The newly introduced taxa were registered in the MycoBank database (https://www.mycobank.org/).

DNA extraction, PCR amplification, and sequencing

Fresh mycelia were scraped with a sterilized surgical knife and transferred to a sterilized 1.5 mL microcentrifuge tube. Genomic DNA was extracted using the Biospin Fungus Genomic DNA Extraction Kit (BioFlux, China), following the manufacturer’s protocol. LSU, ITS, tef1-α, and rpb2 sequence fragments were amplified using primer pairs LR0R/LR5 (Vilgalys and Hester 1990), ITS5/ITS4 (White et al. 1990), EF1-983F/EF1-2218R (Rehner and Buckley 2005), and fRPB2-5F/fRPB2-7cR (Liu et al. 1999), respectively. The PCR amplification reactions were conducted in a 50 µL reaction volume, consisting of 44 µL of 1.1 × T3 Super PCR Mix (Qingke Biotech, Chongqing, China), 2 µL each of forward and reverse primers, and 2 µL of DNA. The thermal cycling parameters for LSU, ITS, tef1-α, and rpb2 regions followed the method described by Ma et al. (2024a). The PCR products were analyzed using 1% agarose gel electrophoresis, and the sequencing results were obtained by Beijing Qingke Biotechnology Co., Ltd.

Phylogenetic analyses

Newly generated DNA sequence data for each region were checked using BioEdit v 7.0.5.3 (Hall 1999) and assembled with SeqMan v. 7.0.0 (DNASTAR, Madison, WI, USA; Swindell and Plasterer 1997). Classification of the newly introduced taxa was analyzed using the BLASTn tool in GenBank (https://www.ncbi.nlm.nih.gov/). Sequences used in this study were downloaded from GenBank (Table 1). Single-gene datasets were aligned using MAFFT v.7.473 (https://mafft.cbrc.jp/alignment/server/, Katoh et al. 2019) and subsequently trimmed with trimAl.v1.2rev59 software (Capella-Gutiérrez et al. 2009). The trimmed datasets were concatenated (LSU-ITS-tef1-α-rpb2) using SequenceMatrix-Windows-1.7.8 software (Vaidya et al. 2011). The maximum likelihood (ML) tree was constructed using the IQ-TREE web server (http://iqtree.cibiv.univie.ac.at/, Nguyen et al. 2015; Zeng et al. 2023). Bayesian Inference (BI) was conducted in accordance with the methods described by Ma et al. (2022). The trimmed Fasta file for each gene dataset was converted to Nexus format for Bayesian analysis using AliView v. 1.27 (Daniel et al. 2010). The best-fit substitution model for the LSU-ITS-tef1-α-rpb2 matrices was selected using MrModeltest 2.3, based on the Akaike Information Criterion (AIC) (Nylander et al. 2008).

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

Taxon Strain GenBank Accessions
LSU ITS tef1-α rpb2
Acanthostigma chiangmaiensis MFLUCC 10-0125T JN865197 JN865209 KF301560 -
Acanthostigma perpusillum UAMH 7237 AY856892 AY916492 - -
Botryosphaeria agaves MFLUCC 10-0051 JX646807 JX646790 - -
Botryosphaeria dothidea CBS 115476 DQ678051 KF766151 DQ767637 DQ677944
Helicosporium acropleurogenum CGMCC 3.25563T PP639430 PP626574 PP596333 PP596460
Helicosporium aquaticum GZCC 22-2120 PP639431 PP626575 PP596334 PP596461
Helicosporium aquaticum MFLUCC 17-2008T MH558859 MH558733 MH550924 MH551049
Helicosporium brunneisporum CGMCC 3.25542T PP639433 PP626577 PP596336 PP596463
Helicosporium changjiangense GZCC 22-2113T PP639434 PP626578 PP596337 PP596464
Helicosporium flavisporum MFLUCC 17-2020T MH558860 MH558734 MH550925 MH551050
Helicosporium flavum GZCC 23-0487 PP639435 PP626579 PP596338 PP596465
Helicosporium flavum MFLUCC 16-1230T KY873621 KY873626 KY873285 -
Helicosporium hainanense GZAAS 22-2006T OP508770 OP508730 OP698081 OP698070
Helicosporium jiangkouense HKAS 128933T PP639436 PP626580 PP596339 PP596466
Helicosporium jiangkouense HKAS 128901 - PP626581 - -
Helicosporium latisporum HKAS 128960T PP639437 PP626582 PP596340 PP596467
Helicosporium liuzhouense GZCC 22-2014T OQ981402 OQ981394 OQ980476 OQ980474
Helicosporium luteosporum MFLUCC 16-0226T KY321327 KY321324 KY792601 MH551056
Helicosporium luteosporum MFLUCC 16-1233 KY873624 - - -
Helicosporium multidentatum GZCC 22-2013T OQ981403 OQ981395 OQ980477 OQ980475
Helicosporium nanningense GZCC 22-2175T OR066425 OR066418 OR058864 OR058857
Helicosporium ramosiphorum CGMCC 3.25541T PP639432 PP626576 PP596335 PP596462
Helicosporium rubrum MFLUCC 24-0090T PQ098514 PQ098477 PQ490681 PQ490675
Helicosporium rubrum GZCC 24-0149 PQ522499 PQ522497 PQ490680 PQ490674
Helicosporium setiferum GZCC 23-0152 PP639438 PP626583 PP596341 PP596468
Helicosporium setiferum BCC 3332 AY856907 AY916490 - -
Helicosporium setiferum BCC 8125 - AY916491 - -
Helicosporium setiferum MFLUCC 17-1994T MH558861 MH558735 MH550926 MH551051
Helicosporium setiferum MFLUCC 17-2006 MH558862 MH558736 MH550927 MH551052
Helicosporium setiferum MFLUCC 17-2007 MH558863 MH558737 MH550928 MH551053
Helicosporium sexuale GZCC 22-2007 OP508771 OP508731 OP698082 OP698071
Helicosporium sexuale MFLUCC 16-1244T MZ538537 MZ538503 MZ567082 MZ567111
Helicosporium sp. NBRC 9014 AY856903 AY916489 - -
Helicosporium thailandense MFLUCC 18-1407T MN913718 MT627698 MT954371 -
Helicosporium vegetum GZCC 23-0060 PP639439 PP626584 PP596342 PP596469
Helicosporium vegetum CBS 941.72 AY856883 AY916488 - -
Helicosporium vegetum NBRC 30345 AY856896 - - -
Helicosporium vegetum CBS 254.75 DQ470982 - DQ471105 -
Helicosporium vegetum CBS 269.52 AY856893 AY916487 - -
Helicosporium vesicarium MFLUCC 17-1795T MH558864 MH558739 MH550930 MH551055
Helicosporium viridiflavum MFLUCC 17-2336T - MH558738 MH550929 MH551054
Helicosporium viridisporum GZCC 23-0044 PP639440 PP626585 - -
Helicosporium viridisporum GZCC 23-0045 PP639441 PP626586 PP596343 PP596470
Helicosporium viridisporum GZCC 22-2008T OP508776 OP508736 OP698087 OP698076
Helicotubeufia hydei MFLUCC 17-1980T MH290026 MH290021 MH290031 MH290036
Helicotubeufia jonesii MFLUCC 17-0043T MH290025 MH290020 MH290030 MH290035
Muripulchra aquatica KUMCC 15-0245 KY320550 KY320533 KY320563 MH551057
Muripulchra aquatica KUMCC 15-0276 KY320551 KY320534 KY320564 MH551058
Muripulchra aquatica DLUCC 0571 KY320548 KY320531 - -
Muripulchra aquatica MFLUCC 15-0249T KY320549 KY320532 - -
Neohelicomyces acropleurogenus CGMCC 3.25549T PP639450 PP626594 PP596351 PP596478
Neohelicomyces aquaticus KUMCC 15-0463 KY320546 KY320529 KY320562 MH551065
Neohelicomyces aquaticus MFLUCC 16-0993T KY320545 KY320528 KY320561 MH551066
Neohelicomyces aseptatus CGMCC 3.25564T PP639451 PP626595 PP596352 PP596479
Neohelicomyces dehongensis MFLUCC 18-1029T MN913709 NR_171880 MT954393 -
Neohelicomyces denticulatus GZCC 19-0444T MW133855 OP377832 - -
Neohelicomyces denticulatus GZCC 23-0073T PP639452 PP626596 PP596353 PP596480
Neohelicomyces deschampsiae CPC 33686T MK442538 MK442602 - -
Neohelicomyces edgeworthiae CGMCC 3.25565T PP639453 PP626597 PP596354 PP596481
Neohelicomyces grandisporus KUMCC 15-0470T KX454174 KX454173 - MH551067
Neohelicomyces guizhouensis GZCC 23-0725T PP512973 PP512969 PP526727 PP526733
Neohelicomyces guizhouensis GZCC 23-0726 PP512974 PP512970 PP526728 PP526734
Neohelicomyces guttulatus CGMCC 3.25550T PP639454 PP626598 PP596355 -
Neohelicomyces guttulatus GZCC 23-0406 PP639455 PP626599 PP596356 PP596482
Neohelicomyces hainanensis GZCC 22-2009T OP508774 OP508734 OP698085 OP698074
Neohelicomyces hainanensis GZCC 22-2027 OP508775 OP508735 OP698086 OP698075
Neohelicomyces helicosporus GZCC 23-0633T PP512975 PP512971 PP526729 PP526735
Neohelicomyces helicosporus GZCC 23-0634 PP512976 PP512972 PP526730 PP526736
Neohelicomyces hyalosporus GZCC 16-0086T MH558870 MH558745 MH550936 MH551064
Neohelicomyces hydei GZCC 23-0727T PP512977 - PP526731 PP526737
Neohelicomyces hydei GZCC 23-0728 PP512978 - PP526732 PP526738
Neohelicomyces lignicola CGMCC 3.25551T PP639456 PP626600 PP596357 PP596483
Neohelicomyces longisetosus NCYU-106H1-1-1T - MT939303 - -
Neohelicomyces macrosporus CGMCC 3.25552T PP639457 PP626601 PP596358 PP596484
Neohelicomyces maolanensis GZCC 23-0079T PQ098529 - PQ490683 PQ490677
Neohelicomyces maolanensis GZCC 24-0148 PQ522500 - PQ490682 PQ490676
Neohelicomyces melaleucae CPC 38042T MN567661 MN562154 MN556835 -
Neohelicomyces melaleucae KUNCC 23-14314 PP664112 PP664108 PP680211 -
Neohelicomyces pallidus CBS 245.49 - MH856510 - -
Neohelicomyces pallidus CBS 271.52 AY856887 AY916461 - -
Neohelicomyces pallidus CBS 962.69 AY856886 AY916460 - -
Neohelicomyces denticulatus UAMH 10535 AY856913 AY916462 - -
Neohelicomyces pandanicola KUMCC 16-0143T MH260307 MH275073 MH412779 -
Neohelicomyces qixingyaensis CGMCC 3.25569T PP639458 PP626602 PP596359 PP596485
Neohelicomyces submersus MFLUCC 16-1106T KY320547 KY320530 - MH551068
Neohelicomyces subtropicus GZCC 23-0076T PQ098530 PQ098492 PQ490685 PQ490679
Neohelicomyces subtropicus GZCC 24-0147 PQ522501 PQ522498 PQ490684 PQ490678
Neohelicomyces thailandicus MFLUCC 11-0005T MN913696 NR_171882 -
Neohelicomyces thailandicus GZCC 23-0400 PP639459 PP626603 PP596360 PP596486
Neohelicomyces xiayadongensis CGMCC 3.25568T PP639460 PP626604 PP596361 PP596487
Neohelicomyces yunnanensis GZCC 23-0735T PP664113 PP664109 – –
Tubeufiaceae sp. ATCC 42524 AY856911 AY916458 - -
Tubeufia guttulata GZCC 23-0404T OR030834 OR030841 OR046678 OR046684
Tubeufia hainanensis GZCC 22-2015T OR030835 OR030842 OR046679 OR046685
Neohelicomyces xiayadongensis MUCL 15702 AY856873 AY916459 - -
Tubeufia javanica MFLUCC 12-0545T KJ880036 KJ880034 KJ880037 -
Tubeufia krabiensis MFLUCC 16-0228T MH558917 MH558792 MH550985 MH551118
Tubeufia latispora MFLUCC 16-0027T KY092412 KY092417 KY117033 MH551119
Tubeufia laxispora MFLUCC 16-0232T KY092408 KY092413 KY117029 MF535287
Tubeufia machaerinae MFLUCC 17-0055 MH558920 MH558795 MH550988 MH551122
Tubeufia mackenziei MFLUCC 16-0222T KY092410 KY092415 KY117031 MF535288
Tubeufia muriformis GZCC 22-2039T OR030836 OR030843 OR046680 OR046686
Tubeufia nigroseptum CGMCC 3.20430T MZ853187 MZ092716 OM022002 OM022001
Tubeufia pandanicola MFLUCC 16-0321T MH260325 MH275091 - -

Note: “T” indicates ex-type strains. The newly generated sequences are highlighted in bold black. “-” indicates the unavailable data in GenBank.

The ML and BI trees were visualized using FigTree v. 1.4.4 and edited with Adobe Illustrator CC 2019 (v. 23.1.0; Adobe Systems, USA). Photoplates were created using Adobe Photoshop CC 2019 (Adobe Systems, USA) and the Tarosoft (R) Image Frame Work program.

Phylogenetic results

The phylogenetic positions of the newly introduced species were elucidated through a multi-gene phylogenetic analysis incorporating LSU, ITS, tef1-α, and rpb2 sequences. The concatenated sequence matrix consisted of 3,418 characters: LSU (1–851), ITS (852–1,444), tef1-α (1,445–2,356), and rpb2 (2,357–3,418), encompassing 102 ingroup taxa and two outgroup taxa, Botryosphaeria agaves and B. dothidea. Both ML and BI analyses yielded similar tree topologies. Fig. 1 presents the highest-scoring ML tree, which achieved a final likelihood value of -31,594.957.

Figure 1. 

Phylogram generated from the best scoring of the RAxML tree based on the combined LSU, ITS, tef1-α, and rpb2 sequence dataset, indicating Helicosporium and Neohelicomyces species. Botryosphaeria agaves (MFLUCC 10–0051) and B. dothidea (CBS 115476) were selected as outgroup taxa. Bootstrap support values from maximum likelihood (ML) equal to or greater than 75% and Bayesian posterior probabilities (PP) equal to or greater than 0.95 are shown near the nodes as ML/PP, respectively. “T” denotes ex-holotype strains. New species are in bold blue.

Based on the phylogenetic analysis (Fig. 1), our collections belong to Helicosporium and Neohelicomyces within Tubeufiaceae (Tubeufiales, Dothideomycetes). Two isolates (MFLUCC 24–0090 and GZCC 24–0149) formed a sister clade with Helicosporium flavum (MFLUCC 16–1230 and GZCC 23–0487), supported by 100% ML and 1.00 PP. Additionally, our new isolates (GZCC 23–0079 and GZCC 24–0148) cluster together with a clade comprising N. helicosporus, N. hyalosporus, N. qixingyaensis, and N. subtropicus. Furthermore, GZCC 23–0076 and GZCC 24–0147 grouped together and clustered sister to N. helicosporus (GZCC 23–0633 and GZCC 23–0634) and N. hyalosporous (GZCC 16–0086), supported by 84% ML and 1.00 PP.

Helicosporium rubrum J. Ma & Y.Z. Lu, sp. nov.

MycoBank No: 902923
Facesoffungi Number: FoF16908
Fig. 2

Holotype

MFLU 24–0035.

Etymology

‘‘rubrum’’ refers to the red-brown colonies on the woody substrate.

Description

Saprobic on decaying wood in a terrestrial habitat. Asexual morph Undetermined. Sexual morph: Ascomata 151–185.5 µm high, 138–157 µm diam., superficial, seated on a subiculum, solitary, scattered, globose to subglobose, bright reddish yellow to brown yellow, with central narrow ostiole; setae were not observed, comprising short projections of setae-like, 10–35 × 4.5–8 µm. Peridium 17.5–22 µm wide, composed of several layers of hyaline to bright yellow cells of textura angularis, outer layer yellow cells, and inner layer pale yellow to hyaline cells. Hamathecium comprising numerous, 1.5–2.5 µm wide, filiform, branched, septate, hyaline pseudoparaphyses. Asci 51–77 × 8–12.5 µm (x̄ = 64 × 10 μm, n = 20), 8-spored, bitunicate, fissitunicate, cylindrical to clavate or saccate, short-pedicellate, apically rounded, basally flexious. Ascospores 27–35 × 3–4.5 µm (x̄ = 31.5 × 4 μm, n = 20), overlapping 2–3-seriate, fusiform, tapering towards the ends, widest at the central part, straight to slightly curved, multi-septate, hyaline, smooth-walled.

Culture characteristics

Conidia germinated on PDA, producing germ tubes within 10 hours. Colonies on PDA reached a diameter of 29 mm after 49 days of incubation at 25 °C, exhibiting an irregular shape with radially furrowed at the centre and velvety surface, white to pale brown in PDA medium.

Material examined

Thailand • Chiangmai, Mushroom Research Center (MRC), on rotting wood in a terrestrial habitat, 11 September 2020, Jing-Yi Zhang, Y251 (MFLU 24–0035, holotype), ex-type living culture MFLUCC 24–0090 = GZCC 24–0149.

Figure 2. 

Helicosporium rubrum (MFLU 24–0035, holotype) a, b ascomata on the host surface c vertical sections of ascomata d peridium e hamathecium pseudoparaphyses f, g short projections of setae-like h–j asci l–o ascospores k germinating ascospore p, q surface and reverse colonies on PDA after 49 days of incubation at 25 °C. Scale bars: 50 μm (c); 20 μm (g–j); 10 μm (d–f, k–o).

Notes

Our newly isolated strains (MFLUCC 24–0090 and GZCC 24–0149) formed a sister relationship with H. flavum (MFLUCC 16–1230 and GZCC 24–0487), supported by 100% ML/1.00 PP support (Fig. 1). A comparison of the ITS, LSU, and tef1-α sequence data between our strain (MFLUCC 24–0090) and H. flavum (MFLUCC 16–1230) revealed nucleotide base differences of 19/506 bp (3.8%, including four gaps), 10/757 bp (1.3%, without gap), and 26/904 bp (2.9%, including four gaps), respectively. Morphologically, Helicosporium rubrum resembles H. flavum (MFLU 17–0704) in having solitary, scattered, globose to subglobose, bright reddish-yellow to brown-yellow ascomata; bitunicate, fissitunicate, cylindrical to clavate asci; and fusiform, straight to slightly curved, multi-septate, hyaline ascospores (Brahmanage et al. 2017). However, H. rubrum differs from H. flavum by having smaller asci (51–77 × 8–12.5 µm vs. 70–130 × 12–16 µm) and shorter ascospores (27–35 × 3–4.5 µm vs. 40–60 × 8–12 µm) (Brahmanage et al. 2017). Additionally, H. flavum exhibits brown to black-brown setae, absent in H. rubrum (Brahmanage et al. 2017). Therefore, based on morphological and molecular data, we propose Helicosporium rubrum as a new species (Chethana et al. 2021).

Neohelicomyces maolanensis J. Ma & Y.Z. Lu, sp. nov.

MycoBank No: 902921
Facesoffungi Number: FoF16909
Fig. 3

Holotype

HKAS 128855.

Etymology

‘‘maolanensis’’ refers to its collection site, where the fungus was collected.

Description

Saprobic on decaying wood in a forest. Sexual morph Undetermined. Asexual morph Hyphomycetous, helicosporous. Colonies on natural substrate superficial, effuse, solitary, scattered or gregarious, white to pale brown. Mycelium partly immersed, partly superficial, composed of pale brown to brown, branched, septate, guttulate, smooth. Conidiophores 201–230 μm long, 3–4.5 μm wide (x̄ = 220 × 3.5 μm, n = 20), macronematous, mononematous, procumbent, solitary, cylindrical, tapering at tip, flexuous, unbranched, septate, slightly constricted at septa, hyaline to pale brown, smooth-walled, thick-walled. Conidiogenous cells 13.5–18.5 μm long, 2.5–4 μm wide (x̄ = 16 × 3.5 μm, n = 30), holoblastic, monoblastic to polyblastic, integrated, intercalary or terminal, cylindrical or subcylindrical, with a denticulate protrusion, truncate at apex after conidial secession, hyaline to pale brown, smooth-walled. Conidia solitary, acropleurogenous, helicoid, tapering towards the rounded ends, developing on tooth-like protrusions, 13.5–19 μm diam. and conidial filament 2.5–3 μm wide (x̄ = 16 × 2.8 μm, n = 30), 105–134 μm long (x̄ = 116.5 μm, n = 30), aseptate, tightly coiled 3–33/4 times, becoming loosely coiled when the conidia are young in water and not becoming loose when the conidia mature in water, guttulate, hyaline, smooth-walled.

Culture characteristics

Conidia germinated on PDA, producing germ tubes within 8 hours. Colonies on PDA reached a diameter of 24 mm after 37 days of incubation at 25 °C, exhibiting an irregular shape with a flat surface and undulate margin, pale brown to brown in PDA medium.

Figure 3. 

Neohelicomyces maolanensis (HKAS 128855, holotype) a colonies on the host surface b–d conidiophores, conidiogenous cells with conidia e, f conidiogenous cells h, i, l–n conidia g germinated conidium j, k surface and reverse colonies on PDA after 37 days of incubation at 25 °C. Scale bars: 50 μm (b, c); 20 μm (d–f); 10 μm (g); 5 μm (h, i, l–n).

Material examined

China • Guizhou Province, Qiannan Buyi and Miao Autonomous Prefecture, Libo County, on rotting wood in a terrestrial habitat, 10 April 2022, Jian Ma, MN5 (HKAS 128855, holotype), ex-type living culture GZCC 23–0079; • Ibid., MN5.1 (GZAAS 23–0634, paratype), ex-paratype living culture GZCC 23–0148.

Notes

Neohelicomyces maolanensis (HKAS 128855) is morphologically similar to N. deschampsiae (CBS H–23590) in having erect, flexuous, multi-septate, brown conidiophores; monoblastic to polyblastic, intercalary, pale brown conidiogenous cells; and solitary, hyaline conidia (Crous et al. 2019a). However, Neohelicomyces maolanensis can be distinguished from N. deschampsiae by its greater number of coils (3–33/4 times vs. 2–3 times), smaller conidial diameter (13.5–19 μm vs. 19–22 μm), and wider conidial filaments (2.5–3 μm vs. 2–2.5 μm) (Crous et al. 2019a). According to the phylogenetic analysis (Fig. 1), our new isolates formed a distinct lineage within the clade, which comprises N. helicosporous (GZCC 23–0633 and GZCC 23–0634), N. hyalosporous (GZCC 16–0086), N. qixingyaensis (CGMCC 3.25569), and N. subtropicus (GZCC 23–0076 and GZCC 24–0147), indicating that GZCC 23–0079 and GZCC 23–0148 represent a distinct species. Therefore, we propose Neohelicomyces maolanensis (GZCC 23–0079 and GZCC 23–0148) as a novel species based on molecular and morphological evidence.

Neohelicomyces subtropicus J. Ma & Y.Z. Lu, sp. nov.

MycoBank No: 902922
Facesoffungi Number: FoF16910
Fig. 4

Holotype

HKAS 128847.

Etymology

‘‘subtropicus’’ named after the climate from which the holotype was found.

Description

Saprobic on decaying wood in a forest. Sexual morph Undetermined. Asexual morph Hyphomycetous, helicosporous. Colonies on natural substrate superficial, effuse, solitary, gregarious, white to pale brown. Mycelium partly immersed, partly superficial, composed of pale brown to brown, branched, septate, guttulate, smooth. Conidiophores up to 420 μm long, 2.5–5.5 μm wide (x̄ = 3.5 μm, n = 30), macronematous, mononematous, erect, solitary or in a group, cylindrical, long or short, tapering at tip, flexuous, mostly branched, septate, slightly constricted at septa, hyaline to pale brown at base, becoming hyaline toward apex, smooth-walled, thick-walled. Conidiogenous cells 10.5–19.5 μm long, 2–5.5 μm wide (x̄ = 15.5 × 3.5 μm, n = 35), holoblastic, monoblastic to polyblastic, integrated, intercalary or terminal, cylindrical or subcylindrical, with a denticulate protrusion, truncate at apex after conidial secession, hyaline to pale brown, smooth-walled. Conidia solitary, acropleurogenous, helicoid, dry, tapering towards the rounded ends, developing on tooth-like protrusions, 14.5–16.5 μm diam. and conidial filament 1.5–3 μm wide (x̄ = 15.5 × 2 μm, n = 25), 87–132 μm long (x̄ = 110.5 μm, n = 25), aseptate, tightly coiled 2–23/4 times, becoming loosely coiled when the conidia are young in water and not becoming loose when the conidia mature in water, guttulate, hyaline, smooth-walled.

Figure 4. 

Neohelicomyces subtropicus (HKAS 128847, holotype) a colonies on the host surface b–e conidiophores, conidiogenous cells with conidia f, g, i conidiogenous cells j–n conidia h germinated conidium o, p surface and reverse colonies on PDA after 46 days of incubation at 25 °C. Scale bars: 100 μm (b); 20 μm (c–h, j–m); 10 μm (i, n).

Culture characteristics

Conidia germinated on PDA, producing germ tubes within 12 hours. Colonies on PDA reached a diameter of 44 mm after 46 days of incubation at 25 °C, exhibiting an irregular shape with a flat surface and undulate margin, pale brown to black brown in PDA medium.

Material examined

China • Guizhou Province, Qiannan Buyi and Miao Autonomous Prefecture, Libo County, on rotting wood in a terrestrial habitat, 10 April 2022, Jian Ma, MN2 (HKAS 128847, holotype), ex-type living culture GZCC 23–0076; • Ibid., MN2.1 (GZAAS 23–0632, paratype), ex-paratype living culture GZCC 23–0147.

Notes

Phylogenetically, our new isolates (GZCC 23–0076 and GZCC 23–0147) formed a sister clade with N. helicosporus (GZCC 23–0633 and GZCC 23–0634) and N. hyalosporus (GZCC 16–0086) (Fig. 1). A comparison of the ITS, LSU, tef1-α, and rpb2 sequence data between our strain (GZCC 23–0076) and N. hyalosporus (GZCC 16–0086) revealed nucleotide base differences of 29/515 bp (5.6%, including 12 gaps), 1/842 bp (0.1%, without gap), 21/912 bp (2.3%, including one gap), and 22/1045 bp (2.1%, without gap), respectively. Morphologically, our newly collected specimen (HKAS 128847) differs from N. helicosporus (HKAS 134923) and N. hyalosporus (HKAS 97441) in having longer conidiophores (up to 420 μm vs. 105–199 μm and 210–290 μm, respectively) (Lu et al. 2018b; Ma et al. 2024a). Additionally, Neohelicomyces subtropicus exhibits branched conidiophores, which are absent in N. helicosporus and N. hyalosporus (Lu et al. 2018b; Ma et al. 2024a). Therefore, we propose Neohelicomyces subtropicus as a new species based on morphological comparison and multi-gene phylogenetic analysis.

Discussion

This study collected five saprobic taxa from terrestrial habitats in China and Thailand. Based on a multi-gene phylogenetic analysis (using LSU, ITS, tef1-α, and rpb2), combined with morphological descriptions, the sexual species, Helicosporium rubrum, and two new helicosporous Neohelicomyces species, N. maolanensis and N. subtropicus, are proposed.

To date, 286 helicosporous species, including the two new species described in this study, are classified within the family Tubeufiaceae (Tubeufiales, Dothideomycetes, Ascomycota) (Ma et al. 2024b). The family Tubeufiaceae comprises 31 helicosporous genera, viz. Acanthohelicospora, Acanthostigmina, Acrohelicosporium, Berkleasmium, Camporesiomyces, Chlamydotubeufia, Dematiohelicoma, Dematiohelicomyces, Dematiohelicosporum, Helicangiospora, Helicoarctatus, Helicodochium, Helicohyalinum, Helicoma, Helicomyces, Helicosporium, Helicotruncatum, Helicotubeufia, Hyalohelicoon, Hyalohelisphora, Hyalotubeufia, Neoacanthostigma, Neohelicomyces, Neohelicosporium, Parahelicomyces, Pleurohelicosporium, Pseudohelicoon, Pseudohelicosporium, Pseudotubeufia, Thaxteriella, and Tubeufia (Liu et al. 2018; Lu et al. 2018a, 2018b, 2023a, 2023b; Lu and Kang 2020; Ma et al. 2023a, 2023b, 2024b; Sruthi et al. 2024). Of which, only 11 genera (Acanthohelicospora, Acanthostigmina, Berkleasmium, Helicangiospora, Helicoma, Helicosporium, Helicotubeufia, Neoacanthostigma, Neohelicosporium, Thaxteriella, and Tubeufia) have been confirmed to have asexual-sexual links based on molecular data and/or morphological characteristics (Samuels et al. 1979; Boonmee et al. 2011; Hyde et al. 2016; Lu et al. 2017b, 2017c, 2018b, 2023a, 2023b; Liu et al. 2018; Lu and Kang 2020; Ma et al. 2023a, 2023b, 2024b; Sruthi et al. 2024). Moreover, three genera (Camporesiomyces, Chlamydotubeufia, and Parahelicomyces) have documented sexual morphs, although their asexual morphs remain undescribed (Hyde et al. 2017, 2020; Hsieh et al. 2021).

Previous multi-gene phylogenetic analyses have demonstrated that the sexual and asexual morphs of certain helicosporous genera exhibit significant diversity (Lu et al. 2017b, 2018b; Boonmee et al. 2014; Ma et al. 2023b, 2024b). For instance, the sexual morph of Tubeufia latispora is characterized by black ascomata with attached unbranched, dark brown setae, sessile asci, and fusiform to cylindrical ascospores, while T. javanica possesses cream-white ascomata, short pedicellate asci, and filiform ascospores. Conidial morphology among Tubeufia species also shows diversity. Most Tubeufia species have hyaline, coiled conidia (Lu et al. 2017b, 2018b; Ma et al. 2024b), but T. africana has ellipsoid to ovoid, spherical to obclavate conidia (Kuo and Goh 2021; Ma et al. 2024b), T. dictyospora has globose to subglobose, ovoid to irregular conidia (Lu et al. 2018b), and T. muriformis features dorsoventrally curved, muriform conidia (Ma et al. 2023b). Additionally, T. sessilis has muriform, curved, coiled conidia (Ma et al. 2024b), and T. subrenispora produces dorsoventrally curved to subreniform, multicelled muriform dark brown conidia (Zhao et al. 2006, 2007; Ma et al. 2024b). Some Tubeufia species also bear small, globose secondary conidia (Zhao et al. 2006, 2007; Ma et al. 2024b). Therefore, multi-gene phylogenetic analyses are essential for correctly identifying these taxa.

Morphological similarities between helicosporous genera can sometimes be misleading, but molecular data often reveal distinct taxonomic positions (Ma et al. 2024b). For example, the asexual morph of Helicosporium species closely resembles Acanthohelicospora scopula and Pseudohelicosporium laxisporum in their pale yellow to yellow-green colonies, setiferous, brown to dark brown conidiophores, and helicoid, hyaline to yellow-green conidia (Linder 1929; Zhao et al. 2007; Boonmee et al. 2014; Brahmanage et al. 2017; Lu et al. 2017a, 2018b, 2022; Xiao et al. 2023; Ma et al. 2024b). However, multi-gene phylogenetic analysis shows that Acanthohelicospora, Helicosporium, and Pseudohelicosporium are distributed across different genera within the family Tubeufiaceae (Ma et al. 2024b).

Acknowledgments

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

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was funded by the Youth Guidance Project of Guizhou Province Basic Research Program (Natural Sciences) in 2024 (Qiankehe Jichu [2024] Qingnian 202).

Author contributions

Morphological data, photo-plates, and phylogenetic analyzes were completed by Jian Ma and Tao Peng. The original draft was written by Jian Ma and Tao Peng. Yong-Zhong Lu, Song Bai, Jing-Yi Zhang, and Na Wu revised the paper.

Author ORCIDs

Tao Peng https://orcid.org/0009-0005-4690-3277

Yong-Zhong Lu https://orcid.org/0000-0002-1033-5782

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

Jing-Yi Zhang https://orcid.org/0000-0003-0606-6169

Xing-Juan Xiao https://orcid.org/0009-0003-8769-4534

Na Wu https://orcid.org/0000-0002-4837-9019

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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