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. 2025 Aug 29;121:237–251. doi: 10.3897/mycokeys.121.158721

Morphological and phylogenetic analyses reveal two new species of Neohelicomyces (Tubeufiales, Tubeufiaceae) from China

Xiao-Yan Ma 1,2, Dan-Dan Song 3,✉, Jian Ma 1,4,✉
PMCID: PMC12413619  PMID: 40917441

Abstract

Neohelicomyces is a genus of helicosporous hyphomycetes with the potential to produce bioactive secondary metabolites. During a survey of helicosporous fungi in Guizhou and Hainan provinces, southern China, four isolates were obtained from both freshwater and terrestrial habitats. Based on combined analyses of multigene phylogenetic data (ITS, LSU, tef1-α, and rpb2) and morphological characteristics, two novel species, Neohelicomyces aquisubtropicus and N. wuzhishanensis, are proposed. Detailed descriptions, illustrations, and phylogenetic analyses of the new taxa are presented. Additionally, a checklist of currently accepted Neohelicomyces species supported by molecular data is provided.

Key words: 2 new species, asexual morph, Dothideomycetes , hyphomycetes, taxonomy

Introduction

Luo et al. (2017) introduced the genus Neohelicomyces and designated N. aquaticus as the type species based on phylogenetic analysis of a combined dataset (ITS, LSU, and tef1-α) and morphological features. The asexual morph of Neohelicomyces is characterized by gregarious colonies that are white, grayish-brown, yellowish-green, or pinkish; macronematous, mononematous, erect, septate, pale brown, branched and/or unbranched conidiophores; mono- to polyblastic, denticulate, integrated, terminal or intercalary conidiogenous cells; and acropleurogenous or pleurogenous, aseptate or septate, guttulate, hyaline, helicoid conidia (Lu et al. 2018, 2022; Tibpromma et al. 2018; Crous et al. 2019a,b; Dong et al. 2020; Hsieh et al. 2021; Yang et al. 2023; Ma et al. 2024a, b; Peng et al. 2025).

Currently, Neohelicomyces comprises 28 species (Yang et al. 2023; Ma et al. 2024b; Peng et al. 2025; Sun et al. 2025). Among them, 12 species are found in freshwater habitats, 12 in terrestrial habitats, and four in both freshwater and terrestrial habitats (Table 1). Neohelicomyces species are distributed in China, the Czech Republic, Germany, Italy, Japan, the Netherlands, Thailand, and the USA (Hsieh et al. 2021; Yang et al. 2023; Ma et al. 2024b; Peng et al. 2025). They occur as saprobes on bamboo culms, Deschampsia cespitosa, Fraxinus excelsior, Melaleuca styphelioides, Miscanthus floridulus, Pandanus sp., Quercus robur, and decaying wood in both freshwater and terrestrial habitats (Linder 1929; Goos 1985, 1986, 1989; Tsui et al. 2006; Zhao et al. 2007; Ruibal et al. 2009; Luo et al. 2017; Lu et al. 2018, 2022; Tibpromma et al. 2018; Crous et al. 2019a, b; Dong et al. 2020; Hsieh et al. 2021; Yang et al. 2023; Ma et al. 2024b; Peng et al. 2025; Sun et al. 2025).

Table 1.

A checklist of accepted Neohelicomyces species with molecular data.

No. Species Distribution Habitat Reference
1 Neohelicomyces acropleurogenus China Terrestrial Ma et al. (2024b)
2 Neohelicomyces aquaticus China Freshwater Luo et al. (2017)
3 Neohelicomyces aquisubtropicus China Terrestrial This study
4 Neohelicomyces aseptatus China Terrestrial Ma et al. (2024b)
5 Neohelicomyces dehongensis China Freshwater Dong et al. (2020)
6 Neohelicomyces denticulatus China Freshwater Yang et al. (2023)
7 Neohelicomyces deschampsiae Germany Terrestrial Crous et al. (2019a)
8 Neohelicomyces edgeworthiae China Terrestrial Ma et al. (2024b)
9 Neohelicomyces guizhouensis China Freshwater Ma et al. (2024a)
10 Neohelicomyces guttulatus China Freshwater/Terrestrial Ma et al. (2024b)
11 Neohelicomyces grandisporus China Freshwater Luo et al. (2017)
12 Neohelicomyces hainanensis China Terrestrial Lu et al. (2022)
13 Neohelicomyces helicosporus China Terrestrial Ma et al. (2024a)
14 Neohelicomyces hyalosporus China Freshwater Lu et al. (2018)
15 Neohelicomyces hydei China Freshwater Ma et al. (2024a)
16 Neohelicomyces lignicola China Freshwater Ma et al. (2024b)
17 Neohelicomyces longisetosus China Freshwater Hsieh et al. (2021)
18 Neohelicomyces macrosporus China Freshwater Ma et al. (2024b)
19 Neohelicomyces maolanensis China Terrestrial Peng et al. (2025)
20 Neohelicomyces melaleucae China, USA Freshwater/Terrestrial Crous et al. (2019b)
21 Neohelicomyces pallidus China, Czech Republic, Italy, Japan, Netherlands, USA Freshwater/Terrestrial Linder (1929); Goos (1989); Tsui et al. (2001); Zhao et al. (2007); Lu et al. (2018); Ma et al. (2024b)
22 Neohelicomyces pandanicola China Terrestrial Tibpromma et al. (2018)
23 Neohelicomyces qixingyansis China Terrestrial Ma et al. (2024b)
Neohelicomyces sexualis China Terrestrial Sun et al. (2025)
24 Neohelicomyces submersus China Freshwater Luo et al. (2017)
25 Neohelicomyces subtropicus China Terrestrial Peng et al. (2025)
26 Neohelicomyces thailandicus China, Thailand Freshwater/Terrestrial Dong et al. (2020); Ma et al. (2024b)
27 Neohelicomyces wuzhishanensis China Freshwater This study
28 Neohelicomyces xiayadongus China Terrestrial Ma et al. (2024b)
29 Neohelicomyces yunnanensis China Freshwater Ma et al. (2024b)
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Note: The newly isolated species in this study are highlighted in bold.

Neohelicomyces has the potential to produce secondary metabolites (Zheng et al. 2023). For example, two alkaloid compounds isolated from Neohelicomyces hyalosporus exhibited moderate cytotoxic effects on human cancer cells (Zheng et al. 2023).

In this study, four helicosporous hyphomycete isolates, representing two distinct taxa, were obtained from both freshwater and terrestrial habitats in Guizhou and Hainan provinces, China. Based on morphological characteristics, illustrations, and multigene phylogenetic analyses, two novel species are introduced: Neohelicomyces aquisubtropicus and N. wuzhishanensis.

Materials and methods

Sample collection, examination, and isolation

Decaying wood samples were collected from freshwater and terrestrial habitats in Guizhou and Hainan provinces, China, from August 2021 to April 2022. After the collection information was recorded (Rathnayaka et al. 2024), the fungal specimens were taken to the mycology laboratory at the Guizhou Institute of Technology for examination. Fresh specimens from freshwater habitats were cultured at room temperature, with moisture maintained for 1–2 weeks. Morphological characteristics were observed using a stereomicroscope (SMZ-168, Nikon, Japan) and photographed using an ECLIPSE Ni compound microscope (Nikon, Tokyo, Japan) equipped with a Canon 90D digital camera.

Single-spore isolations were conducted following the procedure outlined in Senanayake et al. (2020). Subsequently, the germinating spores were aseptically transferred to fresh potato dextrose agar (PDA). Dried specimens were deposited in the Herbarium of Kunming Institute of Botany, Chinese Academy of Sciences (Herb. HKAS), Kunming, China, and the Herbarium of Guizhou Academy of Agriculture Sciences (Herb. GZAAS), Guiyang, China. Pure cultures were deposited at the Guizhou Culture Collection (GZCC), Guiyang, China. The MycoBank numbers were obtained as described at https://www.mycobank.org/.

DNA extraction, PCR amplification, and sequencing

Fresh fungal mycelia grown on PDA media for 39–45 days were scraped with a sterilized toothpick and transferred to a 1.5 ml microcentrifuge tube for genomic DNA extraction. Genomic DNA was extracted using the Biospin Fungus Genomic DNA Extraction Kit (BioFlux, China), following the manufacturer’s protocol. Primer pairs ITS5/ITS4 (White et al. 1990), LR0R/LR5 (Vilgalys and Hester 1990), EF1-983F/EF1-2218R (Rehner and Buckley 2005), and fRPB2-5F/fRPB2-7cR (Liu et al. 1999) were used to amplify ITS, LSU, tef1-α, and rpb2 sequence fragments, respectively. The PCR amplification reactions were carried out in a 25 µL reaction volume, including 1 µL of DNA, 1 µL each of the forward and reverse primers, and 22 µL of 1.1× T3 Super PCR Mix (Qingke Biotech, Chongqing, China). The polymerase chain reaction (PCR) followed the protocol reported by Ma et al. (2024a). The PCR products were detected by 1% agarose gel electrophoresis, and sequencing was performed at Beijing Tsingke Biotechnology Co., Ltd.

Phylogenetic analyses

The forward and reverse sequence data of the new taxa were 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), respectively. The sequences used in this study were downloaded from GenBank (Table 2; https://www.ncbi.nlm.nih.gov/). The single-gene datasets (ITS, LSU, tef1-α, and rpb2) were aligned using MAFFT v. 7.473 (https://mafft.cbrc.jp/alignment/server/, Katoh et al. 2019) and trimmed using trimAl v.1.2rev59 software (Capella-Gutiérrez et al. 2009). The aligned datasets (LSU–ITS–tef1-α–rpb2) were concatenated using SequenceMatrix-Windows v. 1.7.8 software (Vaidya et al. 2011). The maximum likelihood (ML) tree was constructed using the IQ-TREE webserver (http://iqtree.cibiv.univie.ac.at/, Nguyen et al. 2015).

Table 2.

Taxa used in this study, along with their corresponding GenBank accession numbers.

Taxon Strain GenBank Accessions
ITS LSU tef1-α rpb2
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 - -
Neohelicomyces acropleurogenus CGMCC 3.25549T PP626594 PP639450 PP596351 PP596478
Neohelicomyces aquisubtropicus GZCC 23-0080T PQ098499 PQ098537 PV768327 PV768336
Neohelicomyces aquisubtropicus GZCC 24-0163 PV730410 PV730414 PV768328 PV768337
Neohelicomyces aquaticus MFLUCC 16-0993T KY320528 KY320545 KY320561 MH551066
Neohelicomyces aseptatus CGMCC 3.25564T PP626595 PP639451 PP596352 PP596479
Neohelicomyces dehongensis MFLUCC 18-1029T NR_171880 MN913709 MT954393 -
Neohelicomyces denticulatus GZCC 19-0444T OP377832 MW133855 - -
Neohelicomyces deschampsiae CPC 33686T MK442602 MK442538 - -
Neohelicomyces edgeworthiae CGMCC 3.25565T PP626597 PP639453 PP596354 PP596481
Neohelicomyces grandisporus KUMCC 15-0470T KX454173 KX454174 - MH551067
Neohelicomyces guizhouensis GZCC 23-0725T PP512969 PP512973 PP526727 PP526733
Neohelicomyces guttulatus CGMCC 3.25550T PP626598 PP639454 PP596355 -
Neohelicomyces hainanensis GZCC 22-2009T OP508734 OP508774 OP698085 OP698074
Neohelicomyces helicosporus GZCC 23-0633T PP512971 PP512975 PP526729 PP526735
Neohelicomyces hyalosporus GZCC 16-0086T MH558745 MH558870 MH550936 MH551064
Neohelicomyces hydei GZCC 23-0727T - PP512977 PP526731 PP526737
Neohelicomyces lignicola CGMCC 3.25551T PP626600 PP639456 PP596357 PP596483
Neohelicomyces longisetosus NCYU-106H1-1-1T MT939303 - - -
Neohelicomyces macrosporus CGMCC 3.25552T PP626601 PP639457 PP596358 PP596484
Neohelicomyces maolanensis GZCC 23-0079T - PQ098529 PQ490683 PQ490677
Neohelicomyces melaleucae CPC 38042T MN562154 MN567661 MN556835 -
Neohelicomyces pallidus CBS 271.52 AY916461 AY856887 - -
Neohelicomyces pallidus CBS 962.69 AY916460 AY856886 - -
Neohelicomyces pandanicola KUMCC 16-0143T MH275073 MH260307 MH412779 -
Neohelicomyces qixingyaensis CGMCC 3.25569T PP626602 PP639458 PP596359 PP596485
Neohelicomyces submersus MFLUCC 16-1106T KY320530 KY320547 - MH551068
Neohelicomyces subtropicus GZCC 23-0076T PQ098492 PQ098530 PQ490685 PQ490679
Neohelicomyces thailandicus MFLUCC 11-0005T NR_171882 MN913696 - -
Neohelicomyces wuzhishanensis GZCC 23-0410T PQ098494 PQ098532 PV768325 PV768334
Neohelicomyces wuzhishanensis GZCC 24-0164 PV730409 PV730413 PV768326 PV768335
Neohelicomyces xiayadongensis CGMCC 3.25568T PP626604 PP639460 PP596361 PP596487
Neohelicomyces yunnanensis GZCC 23-0735T PP664109 PP664113 - -
Tubeufia guttulata GZCC 23-0404T OR030841 OR030834 OR046678 OR046684
Tubeufia hainanensis GZCC 22-2015T OR030842 OR030835 OR046679 OR046685
Tubeufia javanica MFLUCC 12-0545T KJ880034 KJ880036 KJ880037 -
Tubeufia krabiensis MFLUCC 16-0228T MH558792 MH558917 MH550985 MH551118
Tubeufia latispora MFLUCC 16-0027T KY092417 KY092412 KY117033 MH551119
Tubeufia laxispora MFLUCC 16-0232T KY092413 KY092408 KY117029 MF535287
Tubeufia mackenziei MFLUCC 16-0222T KY092415 KY092410 KY117031 MF535288
Tubeufia muriformis GZCC 22-2039T OR030843 OR030836 OR046680 OR046686
Tubeufia nigroseptum CGMCC 3.20430T MZ092716 MZ853187 OM022002 OM022001
Tubeufia pandanicola MFLUCC 16-0321T MH275091 MH260325 - -
Tubeufiaceae sp. ATCC 42524 AY916458 AY856911 - -
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Note: “T” indicates ex-type strains. Newly generated sequences are in bold. “-” indicates the unavailable data in GenBank.

Bayesian analyses were carried out using MrBayes v. 3.2.7a, and the best nucleotide substitution model for each data partition was selected using MrModeltest v. 2.3 under the Akaike Information Criterion (AIC) (Nylander et al. 2008). The aligned FASTA file was converted to a NEXUS format for Bayesian analysis using AliView v. 1.27 (Larsson 2014).

Phylogenetic trees were visualized using FigTree v. 1.4.4 and subsequently edited in Adobe Illustrator CC 2019 (v. 23.1.0; Adobe Systems, USA). Photo plates and scale bars were prepared using Adobe Photoshop CC 2019 (Adobe Systems, USA) and the Tarosoft Image Framework program, respectively.

Phylogenetic analysis results

The phylogenetic placements of the four new strains were determined by multilocus phylogenetic analysis. The concatenated sequence matrix comprised 3,410 characters (ITS: 1–573, LSU: 574–1,430, tef1-α: 1,431–2,341, and rpb2: 2,342–3,410) across 46 taxa. Both ML and BI analyses produced congruent topologies. Fig. 1 presents the best-scoring ML tree, which had a final log-likelihood value of –19,563.192. Species delimitation and the introduction of new taxa were conducted following the taxonomic framework proposed by Chethana et al. (2021).

Figure 1.

Figure 1.

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Phylogenetic tree generated from ML analysis based on the combined ITS, LSU, tef1-α, and rpb2 sequence data. Bootstrap support values for ML (≥ 75%) and BI (≥ 0.95) are indicated near their respective nodes. The tree is rooted with Helicotubeufia hydei (MFLUCC 17-1980) and H. jonesii (MFLUCC 17-0043). Ex-type strains are denoted with “T,” and newly obtained strains are in bold black fonts.

Based on the multigene phylogenetic tree (Fig. 1), our collections represent two distinct species of Neohelicomyces within the family Tubeufiaceae. Isolates GZCC 23-0080 and GZCC 24-0163 formed a sister clade to the clade comprising Neohelicomyces denticulatus (GZCC 19-0444), N. edgeworthiae (CGMCC 3.25565), and N. pandanicola (KUMCC 16-0143). Isolates GZCC 23-0410 and GZCC 24-0164 formed a clade that is sister to Neohelicomyces guizhouensis (GZCC 23-0725) with 92% ML bootstrap support.

Taxonomy

. Neohelicomyces aquisubtropicus

X.Y. Ma, Y.Z. Lu & J. Ma sp. nov.

6CA725B7-D38A-5694-9E7F-411C2D3CC073

904050

Fig. 2

Figure 2.

Figure 2.

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Neohelicomyces aquisubtropicus (HKAS 128947, holotype). a, b. Colonies on the host surface; c–e. Conidiophores, conidiogenous cells, and conidia; f, g. Conidiogenous cells; h. Germinated conidium; i–m. Conidia; n, o. Colonies on PDA from above and below after 45 days of incubation at room temperature. Scale bars: 20 μm (c–e); 10 μm (f–m).

Etymology.

“aqui-’’ refers to the aquatic habitat of this fungus, and ‘‘-subtropicus’’ means the climate type where the fungus was collected.

Holotype.

HKAS 128947

Description.

Saprobic on decaying wood in a terrestrial habitat. Sexual morph Undetermined. Asexual morph Hyphomycetous, helicosporous. Colonies on natural substrate superficial, white, effuse, gregarious, with massive glistening conidia. Mycelium partly superficial, composed of hyaline to pale brown, branched, septate, guttulate, smooth hyphae. Conidiophores 144–193.5 × 3.5–6.5 μm (x–¯ = 167.5 × 5 μm, n = 25), macronematous, mononematous, erect, cylindrical, straight or slightly flexuous, typically curved at the apex, unbranched, septate, subhyaline to pale brown, thick-walled. Conidiogenous cells 11.5–15 × 3.5–5 μm (x–¯ = 13.5 × 4 μm, n = 30), holoblastic, monoblastic, or polyblastic, integrated, intercalary, cylindrical, with denticles, subhyaline to pale brown, smooth-walled. Conidia solitary, pleurogenous, helicoid, tapering towards the rounded ends, developing on tooth-like protrusions, 14.5–17 μm diam., and conidial filament 2–4 μm wide (x–¯ = 15.5 × 3 μm, n = 25), 82.5–126.5 μm long (x–¯ = 105.5 μm, n = 30), tightly coiled up to 31/2 times, becoming loosely coiled in water, aseptate, guttulate, hyaline, smooth-walled.

Culture characteristics.

Conidia germinate on PDA within 10 hours, producing germ tubes from the conidial body. Colonies on PDA are circular with a raised surface and entire margin, reaching 5 cm in diameter after 45 days at room temperature (approximately 25 °C), and are pale brown to dark brown on both the surface and reverse sides.

Material examined.

China • Guizhou Province, Qiannan Buyi and Miao Autonomous Prefecture, Libo County, on decaying wood in a terrestrial habitat, 10 April 2022, Jian Ma, MN6 (HKAS 128947, holotype), ex-type living cultures GZCC 23-0080; • Ibid., MN6.1 (GZAAS 24-0077, paratype), living culture GZCC 24-0163.

Notes.

Based on phylogenetic analyses, our isolates (GZCC 23-0080 and GZCC 24-0163) clustered with Neohelicomyces denticulatus (GZCC 19-0444), N. edgeworthiae (CGMCC 3.25565), and N. pandanicola (KUMCC 16-0143) (Fig. 1). Neohelicomyces aquisubtropicus (HKAS 128947) differs from N. edgeworthiae (HKAS 128877) in having smaller conidia (14.5–17 μm diam. and 82.5–126.5 μm long vs. 21.5–34 μm diam. and 121–177 μm long) (Ma et al. 2024b). Additionally, N. denticulatus (GZAAS 20-0339) and N. pandanicola (HKAS 96202) can be distinguished from N. aquisubtropicus (HKAS 128947) by their wider conidial diameters (16–22 μm and 28–44 μm vs. 14.5–17 μm) (Tibpromma et al. 2018; Yang et al. 2023). Moreover, base pair comparisons between N. aquisubtropicus (GZCC 23-0080) and related species reveal the following differences. Compared to N. denticulatus (GZCC 19-0444), there are 23/487 bp differences in ITS (4.7%, with 13 gaps). Compared to N. edgeworthiae (CGMCC 3.25565), there are 26/521 bp differences in ITS (5.0%, with 14 gaps), 21/929 bp differences in tef1-α (2.3%, with 7 gaps), and 27/811 bp differences in rpb2 (3.3%, with 10 gaps). In comparison with N. pandanicola (KUMCC 16-0143), there are 27/509 bp differences in ITS (5.3%, with 13 gaps) and 15/827 bp differences in tef1-α (1.8%, with 7 gaps). Therefore, we introduce N. aquisubtropicus as a new species based on morphology and multigene phylogenetic analysis.

. Neohelicomyces wuzhishanensis

X.Y. Ma, Y.Z. Lu & J. Ma sp. nov.

80BE7AFF-2F46-59B0-99AC-881ADA99853E

904051

Fig. 3

Figure 3.

Figure 3.

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Neohelicomyces wuzhishanensis (HKAS 128942, holotype). a, b. Colonies on the host surface; c–h. Conidiophores, conidiogenous cells, and conidia; i–l. Conidia; m. A germinated conidium; n, o. Colonies on PDA from above and below after 39 days of incubation at room temperature. Scale bars: 30 μm (c–f); 20 μm (h, m); 10 μm (g, i–l).

Etymology.

“wuzhishanensis” refers to the type location “Wuzhishan National Nature Reserve.”

Holotype.

HKAS 128942

Description.

Saprobic on submerged decaying wood in a freshwater habitat. Sexual morph Undetermined. Asexual morph Hyphomycetous, helicosporous. Colonies on natural substrate superficial, effuse, gregarious, white. Mycelium partly immersed, composed of hyaline to pale brown, branched, septate, guttulate, smooth hyphae. Conidiophores 92–190 × 3.5–5 μm (x–¯ = 140 × 4.5 μm, n = 25), macronematous, mononematous, erect, cylindrical, widest at the base, tapering towards narrow apex, straight or slightly flexuous, occasionally branched, septate, subhyaline to pale brown, thick-walled. Conidiogenous cells 9.5–16.5 × 2.5–5 μm (x–¯ = 14 × 4 μm, n = 20), holoblastic, monoblastic, or polyblastic, integrated, intercalary or terminal, cylindrical, with tiny tooth-like or bladder-like protrusions, subhyaline to pale brown, smooth-walled. Conidia solitary, acropleurogenous, helicoid, tapering towards the rounded ends, developing on tooth-like protrusions, 23–26 μm diam., and conidial filament 2.3–3.5 μm wide (x–¯ = 24.5 × 2.8 μm, n = 20), 118–143.5 μm long (x–¯ = 129 μm, n = 20), tightly coiled 1.5–2 times, becoming loosely coiled in water, aseptate, guttulate, hyaline, smooth-walled.

Culture characteristics.

Conidia germinate on PDA within 14 hours, producing germ tubes from the conidial body. Colonies on PDA are irregular with a raised surface and undulate margin, reaching 3 cm in diameter after 39 days at room temperature (approximately 25 °C), and are brown to dark brown on both the surface and reverse sides.

Material examined.

China • Hainan Province, Wuzhishan City, Shuimanhe tropical rainforest scenic area in Wuzhishan, 18°92'N, 109°63'E, on rotting wood in a freshwater habitat, 15 August 2021, Jian Ma, WZS8.2 (HKAS 128942, holotype), ex-type living cultures GZCC 23-0410; • Ibid., WZS8.5 (GZAAS 24-0078, paratype), living culture GZCC 24-0164.

Notes.

In our phylogenetic tree (Fig. 1), our isolates (GZCC 23-0410 and GZCC 24-0164) formed a sister clade to N. guizhouensis (GZCC 23-0725) with 92% ML bootstrap support. Neohelicomyces wuzhishanensis (HKAS 128942) can be distinguished from N. guizhouensis (KAS 134924) by its wider conidial diameters (23–26 μm vs. 18–21.5 μm) (Ma et al. 2024a). Moreover, base pair comparison of N. wuzhishanensis (GZCC 23-0410) and N. guizhouensis (GZCC 23-0725) shows 31/539 bp differences in ITS (5.8%, gaps 13 bp), 4/530 bp differences in LSU (0.8%, gaps 3 bp), 13/877 bp differences in tef1-α (1.5%), and 23/939 bp differences in rpb2 (2.4%). Therefore, based on the multigene phylogenetic analysis and morphological differences, we introduce N. wuzhishanensis as a novel species.

Discussion

Neohelicomyces currently comprises 30 species, including the two newly described species, N. aquisubtropicus and N. wuzhishanensis (Hsieh et al. 2021; Yang et al. 2023; Ma et al. 2024a, b; Peng et al. 2025; Sun et al. 2025).

According to previously published studies, six genera—Helicodendron, Helicoma, Helicosporium, Neohelicomyces, Neohelicosporium, and Tubeufia—represent the most species-rich genera among helicosporous hyphomycetes (Lu and Kang 2020, 2022; Ma et al. 2024a, b; Lu et al. 2025; Peng et al. 2025). The asexual morph of most Neohelicomyces species closely resembles that of Helicomyces, Pseudotubeufia, and Tubeufia in having mononematous, septate, pale brown conidiophores; mono- to polyblastic conidiogenous cells; and acropleurogenous or pleurogenous, aseptate or septate, hyaline, helicoid conidia (Yang et al. 2023; Ma et al. 2023, 2024a, b; Peng et al. 2025; Sun et al. 2025). For these morphologically similar helicosporous genera, accurate identification requires a combination of multigene phylogenetic analyses and detailed morphological examination (Linder 1929; Goos 1985, 1986, 1989; Tsui et al. 2006; Zhao et al. 2007; Ruibal et al. 2009; Hyde et al. 2016; Luo et al. 2017; Lu et al. 2018, 2022; Tibpromma et al. 2018; Crous et al. 2019a, b; Dong et al. 2020; Boonmee et al. 2011, 2014, 2021; Hsieh et al. 2021; Yang et al. 2023; Ma et al. 2024a, b; Peng et al. 2025). It is important to note that comparisons of conidial diameter and number of coils among helicosporous species should be based on tightly coiled conidia to ensure consistency and accuracy (Lu et al. 2022, 2023a, b; Xiao et al. 2023; Yang et al. 2023; Ma et al. 2024a, b; Peng et al. 2025).

Some Neohelicomyces species exhibit morphological variations across different habitats and geographical regions (Linder 1929; Goos 1989; Tsui et al. 2001; Zhao et al. 2007; Lu et al. 2018; Yang et al. 2023; Ma et al. 2024b). For example, two collections—HMAS 98776 from a terrestrial habitat in Hebei Province and GZAAS 20-0339 from a freshwater habitat in Guizhou Province, China—both identified as Neohelicomyces pallidus, show distinct morphological characters (Zhao et al. 2007; Yang et al. 2023). HMAS 98776 differs from GZAAS 20-0339 by possessing smaller conidia (10–16 µm vs. 16–22 µm) and exclusively intercalary conidiogenous cells, whereas GZAAS 20-0339 exhibits both terminal and intercalary conidiogenous cells (Zhao et al. 2007; Yang et al. 2023). These morphological differences are speculated to result from environmental variation across habitats and geographic locations. Therefore, species identification within this genus primarily relies on molecular data, which serve as the principal criterion in taxonomic decision-making. This study enriches our understanding of fungal diversity in subtropical and tropical ecosystems and provides cultures of fungal strains for subsequent research on the secondary metabolites of Neohelicomyces.

Supplementary Material

XML Treatment for Neohelicomyces aquisubtropicus
XML Treatment for Neohelicomyces wuzhishanensis

Acknowledgments

We would like to thank Shaun Pennycook (Manaaki Whenua Landcare Research, New Zealand) for his valuable suggestions on the Latin names of the new species.

Citation

Ma X-Y, Song D-D, Ma J (2025) Morphological and phylogenetic analyses reveal two new species of Neohelicomyces (Tubeufiales, Tubeufiaceae) from China. MycoKeys 121: 237–251. https://doi.org/10.3897/mycokeys.121.158721

Funding Statement

This work was funded by the Guizhou Institute of Technology High-level Talent Scientific Research Start-up Fund, grant number 2023GCC066, Guizhou Provincial Higher Education Undergraduate Teaching Content and Curriculum System Reform Project, grant number GZGJ2024209, and Zunyi Technology and Big data Bureau Moutai institute Joint Science and Technology Research and Development Project, grant number ZunShiJiaoHe HZ zi[2023]110, and the Science and Technology Foundation of Guizhou Province (Qian Ke He Pingtai ZSYS[2025]029).

Contributor Information

Dan-Dan Song, Email: 2645729@qq.com.

Jian Ma, Email: yanmajian@163.com.

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 funded by the Guizhou Institute of Technology High-Level Talent Scientific Research Start-Up Fund (grant number 2023GCC066), the Guizhou Provincial Higher Education Undergraduate Teaching Content and Curriculum System Reform Project (grant number GZJG2024209), the Zunyi Technology and Big Data Bureau–Moutai Institute Joint Science and Technology Research and Development Project (grant number ZunShiJiaoHe HZ zi[2023]110), and the Science and Technology Foundation of Guizhou Province (Qian Ke He Pingtai ZSYS[2025]029).

Author contributions

Morphological data, photo-plates and phylogenetic analyzes were completed by Jian Ma and Xiao-Yan Ma. The original draft was written by Jian Ma, and Xiao-Yan Ma and Dan-Dan Song revised the paper.

Author ORCIDs

Xiao-Yan Ma https://orcid.org/0000-0001-5874-9979

Dan-Dan Song https://orcid.org/0009-0005-3175-6438

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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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Neohelicomyces aquisubtropicus
mycokeys.121.158721-treatment1.xml (22KB, xml)
XML Treatment for Neohelicomyces wuzhishanensis
mycokeys.121.158721-treatment2.xml (15.3KB, xml)

Data Availability Statement

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

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