Three novel species of Aquapteridospora (Distoseptisporales, Aquapteridosporaceae) from freshwater habitats in Tibetan Plateau, China

Rong-Ju Xu; Jun-Fu Li; De-Qun Zhou; Saranyaphat Boonmee; Qi Zhao; Ya-Ya Chen

doi:10.3897/mycokeys.102.112905

Abstract

During an investigation of lignicolous freshwater fungi in the Tibetan Plateau, three Aquapteridospora taxa were collected from freshwater habitats in Xizang, China. The new species possess polyblastic, sympodial, denticles conidiogenous cells and fusiform, septate, with or without sheath conidial, that fit within the generic concept of Aquapteridospora, and multi-gene phylogeny placed these species within Aquapteridospora. Detailed morphological observations clearly demarcate three of these from extant species and are hence described as new taxa. The multi-gene phylogeny of the combined LSU, TEF1-α, and ITS sequence data to infer phylogenetic relationships and discuss phylogenetic affinities with morphologically similar species. Based on morphological characteristics and phylogenetic analyses, three new species viz. A. linzhiensis, A. yadongensis, and A. submersa are introduced. Details of asexual morphs are described, and justifications for establishing these new species are also provided in this study.

Key words

3 new taxa, freshwater fungi, morphology, phylogeny, Sordariomycetes, taxonomy

Introduction

Freshwater ascomycetes are the ecological groups that occur saprobically on submerged or partially submerged plant substrates in aquatic habitats (Shearer 1993). Lignicolous freshwater fungi represent a highly diverse taxonomic group with a substantial population. These fungi play a pivotal role in the transfer of nutrients and the flow of energy between trophic levels in the food chain. They achieve this by breaking down complex organic compounds into simpler inorganic materials derived from dead flora and fauna (Krauss et al. 2011; Sridhar et al. 2013; Wurzbacher et al. 2014; Tsui et al. 2016). Recent research showed that lignicolous freshwater fungi comprise a diverse taxonomic assemblage, with more than 3,870 species listed (Calabon et al. 2022). Among them, most are in the classes Dothideomycetes and Sordariomycetes (Hyde et al. 2016; Maharachchikumbura et al. 2016; Luo et al. 2019; Dong et al. 2020; Calabon et al. 2022; Wijayawardene et al. 2022). Sordariomycetes is a prominent class within Ascomycota, encompassing a wide variety of fungi (Luo et al. 2019; Calabon et al. 2022; Yang et al. 2023). In freshwater environments, Sordariomycetes stands out as a significant fungal group, playing a pivotal role in ecosystems. This class is renowned for its production of bioactive compounds (e.g., penicillins, tetracyclines, macrolides, aminoglycosides, and cephalosporins) (Poch et al. 1992; Jones et al. 2014; Wright et al. 2014; Calabon et al. 2023).

Aquapteridospora was initially introduced and classified within the Diaporthomycetidae genera incertae sedis, based on morphological and phylogenetic analyses by Yang et al. (2015). Aquapteridospora, with A. lignicola as the type species, is characterized by polyblastic, sympodial, denticles conidiogenous cells and fusiform, with pale to dark brown central cells and subhyaline end cells, with or without sheath conidia. Furthermore, Hyde et al. (2021a) introduced the family Aquapteridosporaceae to accommodate Aquapteridospora and placed this family in order Distoseptisporales based on divergence estimates, morphological characters, and phylogenetic analyses.

Aquapteridospora is a hyphomycetous genus that are commonly found in freshwater habitats, but only a few terrestrial species, such as A. bambusinum (≡Pleurophragmium bambusinum) was collected from dead culms of bamboo (Yang et al. 2015; Dai et al. 2017; Luo et al. 2019; Bao et al. 2021; Dong et al. 2021; Ma et al. 2022; Peng et al. 2022). These fungi play an important role in the decomposition of organics and nutrient cycling in aquatic environments (Hyde et al. 2016; Luo et al. 2018). In recent years, an increasing number of species in Aquapteridospora have been described and documented, including A. aquatica, A. bambusinum, A. fusiformis, A. hyalina, A. jiangxiensis and A. lignicola (Yang et al. 2015; Luo et al. 2019; Bao et al. 2021; Dong et al. 2021; Ma et al. 2022; Peng et al. 2022).

During an investigation of freshwater fungal diversity on the Tibetan Plateau, six collections possessing morphological characteristics that fit within the genus Aquapteridospora were collected. In particular, their morphological characteristics revealed that these collections were morphologically different from the other species in Aquapteridospora. In addition, phylogenetic analyses of a combined LSU, TEF1-α and ITS sequence data show that our new collections belong to distinct clades, which are distinct from other species in Aquapteridospora. Therefore, three new species viz. Aquapteridospora linzhiensis, A. submersa and A. yadongensis are introduced, as well as details of asexual morphs being described, and justifications for establishing these new species are provided in this study.

Materials and methods

Collection, morphological examination and isolation

Submerged decaying wood samples were collected from freshwater habitats in southeast Xizang, China. Fresh specimens were studied following the methods of Senanayake et al. (2020). Microscopic structures were examined by using a stereomicroscope (SteREO Discovery.V12, Carl Zeiss Microscopy GmBH, Germany), photographed by using a Nikon ECLIPSE 80i compound microscope fitted with a Nikon DS-Ri2 digital camera, and measured by using the Tarosoft (R) Image Frame Work program. Illustrated figures were processed by using Adobe Photoshop CS6 version 10.0 software (Adobe Systems, San Jose, CA, USA).

Single spore isolation was performed on potato dextrose agar (PDA) plates following the methods described in Senanayake et al. (2020). Fungal herbarium specimens and axenic living cultures were deposited in the Herbarium of Cryptogams of the Kunming Institute of Botany, Chinese Academy of Sciences (KUN-HKAS) and Kunming Institute of Botany Culture Collection (KUNCC), Kunming, China. Faceoffungi and Index Fungorum numbers of novel species were registered (Jayasiri et al. 2015, http://www.indexfungorum.org/Names/Names.asp).

DNA extraction, PCR amplification, and sequencing

Fresh mycelia were scraped off from colonies on PDA plates and transferred to a 1.5-ml microcentrifuge tube using a sterilized lancet for genomic DNA extraction. The TOLOBIO Plant Genomic DNA Extraction Kit, Shanghai Co. Ltd. P.R. China was used to extract fungal genomic DNA, following the protocols in the manufacturer’s instructions. The DNA polymerase chain reaction (PCR) amplifications were performed by using primer pairs as follows: ITS5/ITS4 for internal transcribed spacer rDNA region and covered 5.8S ribosomal (ITS); LR0R/LR5 for the nuclear ribosomal large subunit 28S rDNA gene (LSU), and TEF1-983F/TEF1-2218R for TEF1-α (Vilgalys and Hester 1990; White et al. 1990). DNA template was carried out in 25 μL reaction volume containing 21 μL of 1 × Power Taq PCR Master Mix, 1 μL of each primer (10 μL stock) and 2 μL of genomic DNA template. Amplifications were carried out by using the BioTeke GT9612 thermocycler (Beijing City, China). The PCR amplification conditions for ITS and LSU consisted of initial denaturation at 98 °C for 3 minutes, followed by 35 cycles of denaturation at 98 °C for 20 seconds, annealing at 53 °C for 10 seconds, extension at 72 °C for 20 seconds, final extension at 72 °C for 5 minutes; the PCR amplification conditions for TEF1-α consisted of initial denaturation at 98 °C for 3 minutes, followed by 35 cycles of denaturation at 98 °C for 20 seconds, annealing at 64 °C for 10 seconds, extension at 72 °C for 20 seconds, final extension at 72 °C for 5 minutes. PCR products were visualized by using 1% agarose gel electrophoresis stained with ethidium bromide and distinct bands were checked in Gel documentation system (Compact Desktop UV Transilluminator analyzer GL-3120). The PCR products were sequenced by Tsingke Company, Beijing, P.R. China.

Phylogenetic analyses

The sequences were uploaded in GenBank database (http://www.ncbi.nlm.nih.gov/blast/) to search for similar taxa. Sequences generated from the LSU, TEF1-α and ITS gene regions were carefully verified before further analyses. The new sequences were submitted to GenBank, and the strain information used in this paper was provided in Table 1. Multiple sequence alignments were aligned with MAFFT v.7 (Katoh and Standley 2016) http://mafft.cbrc.jp/alignment/server/index.html] and dataset was trimmed by TrimAlv.1.3 using the gappyout option (http://phylemon.bioinfo.cipf.es/utilities.html) (Capella-Gutierrez et al. 2009). A combined sequence dataset was performed with the SquenceMatrix v.1.7.8 (Vaidya et al. 2011).

Table 1.

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Strains used for phylogenetic analyses and their corresponding GenBank numbers. The newly generated sequences are in cells with light grey shading and the type strain are in bold font.

Species	Voucher number	GenBank accession number			Reference
Species	Voucher number	LSU	ITS	TEF1-α	Reference
*Aquapteridospora aquatica*	MFLUCC 17-2371	MW287767	MW286493	/	Dong et al. (2021)
*A. bambusinum*	MFLUCC 12-0850	KU863149	KU940161	KU940213	Dai et al. (2017)
A. bambusinum	MFLUCC_21_0027	MZ412526	MZ412514	MZ442688	Bao et al. (2021)
*A. hyalina*	GZCC 22-0072	ON527945	ON527937	ON533681	Ma et al. (2022)
A. hyalina	GZCC 22-0073	ON527948	ON527940	ON533684	Ma et al. (2022)
*A. jiangxiensis*	JAUCC 3008	MZ871502	MZ871501	MZ855767	Peng et al. (2022)
*A. fusiformis*	MFLUCC 18-1606	MK849798	MK828652	MN194056	Luo et al. (2019)
*A. lignicola*	MFLUCC 15-0377	KU221018	MZ868774	MZ892980	Yang et al. (2015)
*A. linzhiensis*	KUNCC 10420	OQ970576	OP626343	OR597592	This study
A. linzhiensis	KUNCC 10444	OQ970575	OQ847781	OR597591	This study
*A. submersa*	KUNCC 10446	OQ970579	OQ847783	OR597595	This study
A. submersa	KUNCC 10449	OQ970580	OQ970557	OR597596	This study
*A. yadongensis*	KUNCC 10445	OQ970577	OQ847782	OR597593	This study
A. yadongensis	KUNCC 10448	OQ970578	OQ970556	OR597594	This study
*Distoseptispora atroviridis*	GZCC 20-0511	MZ868763	MZ868772	MZ892978	Yang et al. (2021)
*D. bambusae*	MFLUCC 20-0091	MT232718	MT232713	MT232880	Sun et al. (2020)
D. euseptata	MFLU 20-0568	MW081545	MW081540	MW084994	Li et al. (2021)
*D. fusiformis*	GZCC 20-0512	MZ868764	MZ868773	MZ892979	Yang et al. (2021)
*D. guizhouensis*	GZCC 21-0666	MZ474869	MZ474868	MZ501610	Hyde et al. (2021b)
*D. hyalina*	MFLUCC 17-2128	MZ868760	MZ868769	MZ892976	Yang et al. (2021)
D. multiseptata	MFLU 17-0856	MF077555	MF077544	MF135652	Yang et al. (2018)
*D. rayongensis*	MFLUCC 18-0415	MH457137	MH457172	MH463253	Hyde et al. (2020)
D. rayongensis	MFLUCC 18-0417	MH457138	MH457173	MH463254	Hyde et al. (2020)
*D. rostrata*	MFLUCC 16-0969	MG979766	MG979758	MG988424	Luo et al. (2018)
*D. saprophytica*	MFLUCC 18-1238	MW287780	MW286506	MW396651	Dong et al. (2021)
*D. verrucosa*	GZCC 20-0434	MZ868762	MZ868771	MZ892977	Yang et al. (2021)
*D. xishuangbannaensis*	KUMCC 17-0290	MH260293	MH275061	MH412768	Tibpromma et al. (2018)
*D. yunnansis*	MFLUCC 20-0153	MW081546	MW081541	MW084995	Li et al. (2021)
*Pseudostanjehughesia aquitropica*	MFLUCC 16-0569	MF077559	MF077548	MF135655	Yang et al. (2018)
*P. lignicola*	MFLUCC 15-0352	MK849787	MK828643	MN194047	Luo et al. (2019)
*Sporidesmium dulongense*	MFLUCC 17-0116	MH795817	MH795812	MH801191	Luo et al. (2019)
*S. lageniforme*	DLUCC 0880	MK849782	MK828640	MN194044	Luo et al. (2019)
*S. pyriformatum*	MFLUCC 15-0620	KX710141	KX710146	MF135662	Hyde et al. (2016)
S. thailandense	MFLUCC 15-0617	MF077561	MF077550	MF135657	Yang et al. (2018)
*S. thailandense*	MFLUCC 15-0964	MF374370	MF374361	MF370957	Zhang et al. (2017)
*Myrmecridium aquaticum*	MFLUCC 15-0366	MK849804	/	/	Luo et al. (2019)
M. aquaticum	S-1158	MK849803	MK828656	MN194061	Luo et al. (2019)
*M. banksiae*	CBS 132536	JX069855	JX069871	/	Crous et al. (2012)
M. schulzeri	CBS 100.54	EU041826	EU041769	/	Arzanlou et al. (2007)

Maximum likelihood (ML) analysis was performed by RAxML-HPC2 v.8.2.12 (Stamatakis 2014) in the CIPRES Science Gateway web server (http://www.phylo.org/portal2) by using 1,000 rapid bootstrap replicates and the GTRGAMMA+I model. Bootstrap support values for ML equal to or greater than 75% were given above the nodes in the phylogenetic tree (Fig. 1). The model of evolution for the Bayesian inference (BI) analysis was performed by using MrModeltest v2.3 (Nylander 2004). GTR+I+G was selected as the best-fitting model for LSU, TEF1-α and ITS dataset. The Markov chain Monte Carlo sampling (BMCMC) was carried out to assess posterior probabilities (PP) by using MrBayes v.3.2.7 (Ronquist et al. 2012). Six simultaneous Markov chains were run for random trees for 1,000,000 generations, and trees were sampled every 200^th generation. Bayesian posterior probabilities (PP) equal to or greater than 0.95 were given above the nodes in the phylogenetic tree (Fig. 1). Phylograms were visualized by using FigTree v1.4.0 (Rambaut 2012) and rearranged in Adobe Photoshop CS6 software (Adobe Systems, USA). The new sequences were deposited in GenBank (Table 1), and the final alignments and phylogenetic tree were registered in TreeBASE under the submission ID: 30133 (http://www.treebase.org/).

Figure 1.

Maximum likelihood (ML) tree is based on combined LSU, TEF1-α and ITS sequence data. ML bootstrap support values equal to or greater than 70% and Bayesian posterior probabilities (PP) equal to or greater than 0.95 given above the nodes, shown as “ML/PP”. The tree is rooted with Pseudostanjehughesia aquitropica (MFLUCC 16-0569) and P. lignicola (MFLUCC 15-0352). New species are indicated in red and type strains are in bold.

Results

Phylogenetic analyses

The concatenated sequence dataset of LSU, TEF1-α and ITS, comprised 39 strains with Pseudostanjehughesia aquitropica (MFLUCC 16-0569) and P. lignicola (MFLUCC 15-0352) as the outgroup taxa (Fig. 1). The datasets contained 2,168 characters including gaps after alignments (LSU: 1–763 bp, -α = 764–1,660 bp, ITS: 1,661–2,168 bp). The RAxML analysis of the combined datasets yielded a best scoring tree with a final ML optimization likelihood value of -15404.143090. The aligned sequences matrix comprised 849 distinct alignment patterns with 6.45% of undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.229844, C = 0.282249, G = 0.282387, T = 0.205520, with substitution rates AC = 0.921073, AG = 2.039438, AT = 1.172967, CG = 0.817703, CT = 5.518393, GT = 1.000000; gamma distribution shape parameter α = 0.0010000000. The tree topologies of combined sequence data obtained from ML and BI analyses were not significantly different (Fig. 1).

The phylogenetic analyses showed that our six strains nested within the genus Aquapteridospora represent three species. Two strains of A. linzhiensis (KUNCC 10420 and KUNCC 10444) formed a well resolved subclade sister to A. fusiformis (93% ML/1.00 PP support); while strains of A. yadongensis (KUNCC 10445 and KUNCC 10448) formed a distinct subclade sister to A. submersa (KUNCC 10446 and KUNCC 10449) with a high support (100% ML/1.00 PP) and clustered with A. lignicola (MFLUCC 15-10377) with a significant support (75% ML/0.96 PP) (Fig. 1).

Taxonomy

Aquapteridospora linzhiensis R.J. Xu, Q. Zhao & Boonmee, sp. nov.

Index Fungorum: IF901109

Facesoffungi Number: FoF14348

Fig. 2

Etymology

Referring to the location “Linzhi City, China” where the holotype of this fungus was collected.

Holotype

HKAS 128991.

Description

Saprobic on decaying wood submerged in freshwater. Sexual morph: Undetermined. Asexual morph: Colonies on the natural substrate effuse, hairy, pale brown to brown, scattered or in small groups. Mycelium mostly superficial, consisting of branched, septate, smooth, pale brown to brown hyphae. Conidiophores 113–210 × 4–6 μm (x̄ = 162 × 4 μm, n = 15), macronematous, mononematous, solitary or 2–3 group, erect, straight or slightly flexuous, simple, unbranched, smooth, cylindrical, 6–12-septate, brown at the base, pale brown towards apex. Conidiogenous cells polyblastic, monoblastic, terminal, becoming intercalary, cylindrical, pale brown, integrated, with several sympodial proliferations, conspicuous denticles, bearing tiny, protuberant, circular scars. Conidia 10–14 × 5–6 μm (x̄ = 12 × 6 μm, n = 25), solitary or acropleurogenous, fusiform or elliptical, smooth, 2-septate, truncate at base, dark brown in central cells and subhyaline at end cells, guttulate. Conidial secession schizolytic.

Figure 2.

Aquapteridospora linzhiensis (HKAS 128991, holotype) a colonies on the substratum b–e conidiophores, conidiogenous cells with conidia f, g conidiogenous cells with developmental conidia h–k conidia l, m culture on PDA. Scale bars: 50 μm (b–e); 20 μm (f, g); 10 μm (h–k).

Culture characteristics

Conidia were germinated on PDA within 48 hours. Germ tubes produced from each end. Colonies grown on PDA, circular, flat, superficial, dark brown from above, reverse-side brown in the centre, with greyish white near the edge.

Material examined

China, Xizang, Linzhi City, Motuo County, on submerged decaying wood, 1675 msl, 29°10'56"N, 95°8'53"E, 11 July 2022, R.J. Xu, XK-33–3 (HKAS 128991, holotype), ex-type living culture (KUNCC 10420). Xizang, Linzhi City, Motuo County, Gelin Village, on submerged decaying wood, 1143 msl, 29°1'43"N, 94°48'5.7"E, 12 July 2022, R.J. Xu, XK-32, (HKAS 128990), living culture (KUNCC 10444).

Notes

Phylogenetic analyses show that Aquapteridospora linzhiensis (KUNCC 10420 and KUNCC 10444) clustered into a distinct subclade and sister to A. fusiformis (MFLUCC 18-1606) with bootstrap support (93% ML/1.00 PP, Fig. 1). However, A. linzhiensis differs from A. fusiformis in having obvious, guttulate conidia and less septate on maturity (2-septate vs. 3–4-septate) (Luo et al. 2019). Additionally, comparisons of ITS sequences demonstrate a 6.7% (39/586 bp, excluding gaps) difference between A. linzhiensis and A. fusiformis Jeewon and Hyde (2016). Therefore, A. linzhiensis was identified as a new species supported with both morphological and phylogenetic evidences.

Aquapteridospora yadongensis R.J. Xu, Q. Zhao & Boonmee, sp. nov.

Index Fungorum: IF901110

Facesoffungi Number: FoF14349

Fig. 3

Etymology

Referring to the location “Yadong County, China” where the holotype of this fungus was collected.

Holotype

HKAS 128992.

Description

Saprobic on decaying wood submerged in freshwater. Sexual morph: Undetermined. Asexual morph: Colonies on the natural substrate effuse, hairy, pale brown to brown, scattered or in small groups, usually retiform. Mycelium mostly superficial, consisting of branched, septate, smooth, pale brown to brown hyphae. Conidiophores 440–856 × 4–6 μm (x̄ = 581 × 5 μm, n = 20), macronematous, mononematous, solitary, erect, straight or slightly flexuous, unbranched, smooth, cylindrical, multi-septate, tapering towards apex, brown to pale brown, slightly constricted at some septa. Conidiogenous cells polyblastic, monoblastic, terminal, becoming intercalary, cylindrical, pale brown, integrated, denticles, bearing tiny, protuberant, circular scars. Conidia 14–20 × 4–7 μm (x̄ = 17 × 5 μm, n = 30), acropleurogenous, fusiform, smooth, 3-septate, rounded at apex, truncate at base, dark brown in central cells and light at end cells. Conidial secession schizolytic.

Figure 3.

Aquapteridospora yadongensis (HKAS 128992, holotype) a colonies on the substratum b, c conidiophore and conidiogenous cell d-g conidiogenous cells with developmental conidia h–k conidia l germinating conidium m culture on PDA. Scale bars: 100 μm (b, c); 20 μm (d, g); 10 μm (h–l).

Culture characteristics

Conidia were germinated on PDA within 48 hours. Germ tubes produced from each end. Colonies grown on PDA, regular concentric circles, flat, superficial, with dense mycelium at around, grey brown from above, dark brown from below.

Material examined

China, Xizang, Shigatse City, Yadong County, on submerged decaying wood, 3061 msl, 27°21'11"N, 88°58'10"E, 01 July 2022, R.J. Xu, LTS-20 (HKAS 128992, holotype), ex-type living culture (KUNCC 10445). Xizang, Shigatse City, Dingjie County, on submerged decaying wood, 3042 msl, 27°53'8.7"N, 87°27'36"E, 05 July 2022, L.T. Shun, LTS-20–1, (HKAS 128993), living culture (KUNCC 10448).

Notes

Aquapteridospora yadongensis possess its conidial characteristics that fit with Aquapteridospora (Yang et al. 2015). In phylogenetic analyses, A. yadongensis formed a distinct lineage close to A. submersa with high bootstrap support (100% ML/1.00 PP, Fig. 1). A comparison of ITS nucleotide shows that A. yadongensis (KUNCC 10445) differs from A. submersa (KUNCC 10446) in 10/572 bp (1.8%, excluding gap), a comparison of TEF1-α nucleotide shows that A. yadongensis (KUNCC 10445) differs from A. submersa (KUNCC 10446) in 8/821 bp (0.8%, excluding gap) (Jeewon and Hyde 2016). In addition, A. yadongensis differs from A. submersa in having narrower conidiophores (4–6 vs. 5–12 μm), while conidia of A. submersa have slightly constricted septa; the culture of A. yadongensis have regular concentric circles differing from A. submersa having pale mycelium in the centre. Furthermore, A. yadongensis differs from A. lignicola in having long conidiophores (440–856 vs. 70–200 μm) and conidia without a conspicuous sheath (Yang et al. 2015).

Aquapteridospora submersa R.J. Xu, Q. Zhao & Boonmee, sp. nov.

Index Fungorum: IF901111

Facesoffungi Number: FoF14350

Fig. 4

Etymology

Referring to the fungus’s habitat “decaying wood submerged in freshwater habitats”.

Holotype

HKAS 128980.

Description

Saprobic on decaying wood submerged in freshwater. Sexual morph: Undetermined. Asexual morph: Colonies on the natural substrate effuse, glistening, pale brown to brown, scattered or in small groups. Mycelium mostly superficial, consisting of branched, septate, smooth, pale brown to brown hyphae. Conidiophores 376–708 × 5–12 μm (x̄ = 451 × 7 μm, n = 20), macronematous, mononematous, solitary, erect, straight or slightly flexuous, unbranched, smooth, cylindrical, multi-septate, tapering towards apex, brown to pale brown. Conidiogenous cells polyblastic, monoblastic, terminal, becoming intercalary, cylindrical, pale brown, integrated, with several sympodial proliferations, conspicuous denticles, bearing tiny, protuberant, circular scars. Conidia 19–22 × 6–8 μm (x̄ = 21 × 7 μm, n = 20), solitary or acropleurogenous, fusiform, smooth, 2–3-septate, rounded at apex, truncate at base, slightly constricted at septa, hyaline when young, sub-hyaline to pale brown when mature, two big guttulate when young. Conidial secession schizolytic.

Figure 4.

Aquapteridospora submersa (HKAS 128980, holotype) a colonies on the substratum b–d conidiophores, conidiogenous cells with conidia e–g conidiogenous cells with developmental conidia h–k conidia l germinating conidium m, n culture on PDA. Scale bars: 50 μm (b–d); 20 μm (e–g); 10 μm (h–l).

Culture characteristics

Conidia were germinated on PDA within 48 hours. Germ tubes produced from each end. Colonies grown on PDA, circular, flat, superficial, raised, with dense, pale mycelium in the centre. Grey brown from above, dark brown from below.

Material examined

China, Xizang, Linzhi City, Motuo County, on submerged decaying wood, 677 msl, 29°19'43"N, 95°21'19"E, 13 July 2022, R.J. Xu, LJN-15 (HKAS 128980, holotype), ex-type living culture (KUNCC 10446). Xizang, Linzhi City, Motuo County, Gelin Village, on submerged decaying wood, 677 msl, 29°19'43"N, 95°21'19"E, 12 July 2022, R.J. Xu, LJN-15–5, (HKAS 128981), living culture (KUNCC 10444).

Notes

Phylogenetic analyses show that Aquapteridospora submersa (KUNCC 10446, KUNCC 10444), formed a sister grouped with A. yadongensis (KUNCC 10445 and KUNCC 10488) and was close to A. lignicola (MFLUCC 15-0377) with 75% ML/0.96 PP, Fig. 1. However, the comparison of conidial characteristics and nucleotides shows that A. submersa differs from A. yadongensis (see the notes of A. yadongensis). Indeed, A. submersa differs from A. lignicola in having long conidiophores (376–708 vs. 70–200 μm) and conidia without a conspicuous sheath (Yang et al. 2015). Aquapteridospora submersa is introduced here as a new species.

Discussion

Species of Aquapteridospora are morphologically unique in the taxonomic characteristics, especially in the features of the conidiophores and conidia (Table 2). In most species, the conidia are fusiform and pigmented, featuring brown to dark brown central cells and subhyaline end cells. However, some species exhibit conidia with a distinct sheath, such as A. aquatica, A. jiangxiensis and A. lignicola (Yang et al. 2015; Dong et al. 2021; Peng et al. 2022). Additionally, a few species are characterized by hyaline to sub-hyaline conidia, as observed in A. hyalina (Ma et al. 2022). In addition, the length of conidiophores in species of Aquapteridospora varies significantly. Most species have conidiophores ranging in length from 70 to 305 μm, as observed in species like A. aquatica, A. fusiformis, A. hyalina, A. jiangxiensis, A. lignicola and A. linzhiensis (Yang et al. 2015; Luo et al. 2019; Dong et al. 2021; Ma et al. 2022; Peng et al. 2022), a few species exhibit conidiophores exceeding 400 μm in length, with the longest reaching 856 μm. This is the case for species such as A. bambusinum, A. yadongensis and A. submersa (Bao et al. 2021, this study).

Table 2.

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Synopsis of known species in Aquapteridospora.

Species	Conidiophores (μm)	Conidiogenous cells (μm)	Conidia (μm)	Host	Habitat	Distribution	Reference
Aquapteridospora aquatica	125–215 × 3–5	10–85 × 4–5.5, Polyblastic, terminal, intercalary, denticles	19–27.5 × 5–7.5, acropleurogenous, solitary, olivaceous or brown in the middle cells, fusiform, 3-septate, gelatinous, thin sheath	Unidentified, submerged wood	Freshwater	Thailand	Dong et al. (2021)
A. bambusinum	615–715 × 9–13	Polyblastic, sympodial, denticulate, integrated, terminal	15–18 × 5.5–7, acrogenous, solitary, pale brown to dark brown, ellipsoid to fusiform, 3-septate, straight	Unidentified, submerged wood	Freshwater	Thailand	Bao et al. (2021)
A. fusiformis	(88–) 134–188 × 5–7	Polyblastic, terminal, intercalary, sympodial proliferations	14–18 × 5–7, solitary, brown to dark brown in central cells and subhyaline at end cells, fusiform, 3–4-septate,	Unidentified, submerged wood	Freshwater	China	Luo et al. (2019)
A. hyalina	68–130 × 4.5–6.5	25–62 × 4–6.5, polyblastic, monoblastic, denticles	17–28 × 4–6, acropleurogenous, solitary, sub-hyalina to pale brown, fusiform, 1–3-septate,	Unidentified, submerged wood	Freshwater	China	Ma et al. (2022)
A. jiangxiensis	78–305 × 4–7	20–68 × 4–6, integrated, terminal, intercalary	20–25 × 6–7.5, acrogenous or lateral, dark brown to black, fusiform to subclavate, 3-septate, sometimes with a sheath	Unidentified, submerged wood	Freshwater	China	Peng et al. (2022)
A. lignicola	70–200 × 4–7	14.5–30 × 4.5–7.5, polyblastic, terminal, intercalary	15–24 × 6–8, solitary, acropleurogenous, with pale to dark brown central cells and subhyaline end cells, fusiform, 3-septate, with a conspicuous sheath	Unidentified, submerged wood	Freshwater	Thailand	Yang et al. (2015)
A. linzhiensis	113–210 × 4–6	Polyblastic, terminal, intercalary, denticles	10–14 × 5–6, solitary or acropleurogenous, dark brown in central cells and subhyaline at end cells, fusiform or elliptical, 2-septate, guttulate	Unidentified, submerged wood	Freshwater	China	This study
A. yadongensis	440–856 × 4–6	Polyblastic, monoblastic, terminal, intercalary, denticles	14–20 × 4–7, acropleurogenous, dark brown in central cells and subhyaline at end cells, fusiform, 3-septate	Unidentified, submerged wood	Freshwater	China	This study
A. submersa	376–708 × 5–12	Polyblastic, monoblastic, terminal, intercalary, denticles	19–22 × 6–8, solitary or acropleurogenous, hyaline when young, sub-hyaline to pale brown when mature, fusiform, 2–3-septate, two big guttulate when young	Unidentified, submerged wood	Freshwater	China	This study

Molecular phylogenetic analyses play a crucial role in elucidating the classification of hyphomycetous fungi (Dhanasekaran et al. 2006; Tekpinar and Kalmer 2019). Pleurophragmium bambusinum was initially described by Dai et al. (2017), and was previously assigned to Sordariomycetes incertae sedis based on its morphological characteristics. According to the phylogenetic analysis conducted by Dong et al. (2021), P. bambusinum was found to cluster within the Aquapteridospora clade with (100% ML/1.00 PP) support. However, their studies did not synonymize P. bambusinum under Aquapteridospora due to the ellipsoidal and conidia without a sheath, which indicate that it does not fit within the characteristics of Aquapteridospora species. Subsequently, Bao et al. (2021) transferred P. bambusinum to Aquapteridospora and synonymized A. bambusinum instead of P. bambusinum, based on both phylogeny and morphology.

The Tibetan Plateau is renowned for its distinctive biological diversity and extensive array of aquatic habitats, encompassing lakes, rivers, and wetlands, which provide sustenance for various fungal communities (Yao et al. 2019). While freshwater fungi play a crucial role in the ecosystem, they have remained understudied in this region, primarily due to the limited number of researchers focusing on freshwater fungi in the Tibetan Plateau. During our investigation into freshwater fungal diversity on the Tibetan Plateau, we introduced three new species within the genus Aquapteridospora, supported by both phylogenetic analysis and morphology. The discovery of these new species revealed the abundant fungal diversity in Tibetan Plateau and more scientific studies in this region are expected in the future.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This study is supported by the Survey of Wildlife Resources in Key Areas of Tibet (ZL202203601), the Second Tibetan Plateau Scientific Expedition and Research (STEP) Program (Grant No. 2019QZKK0503); Major science and technology projects and key R&D plans/programs, Yunnan Province (202202AE090001), the Biodiversity Survey and Assessment Project of the Ministry of Ecology and Environment, P.R. China (2019HJ2096001006). The authors appreciate the support given by Thesis Writing Grant of Mae Fah Luang University, Thailand, to Rong Ju Xu.

Author contributions

Funding acquisition: QZ. Writing - original draft: RJX. Writing - review and editing: SB, JFL, DQZ.

Author ORCIDs

Rong-Ju Xu https://orcid.org/0000-0002-3968-8442

Jun-Fu Li https://orcid.org/0009-0008-6088-2072

De-Qun Zhou https://orcid.org/0009-0009-3459-3186

Saranyaphat Boonmee https://orcid.org/0000-0001-5202-2955

Qi Zhao https://orcid.org/0000-0001-8169-0573

Ya-Ya Chen https://orcid.org/0000-0002-8293-168X

Data availability

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

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﻿Abstract

Key words

﻿Introduction

﻿Materials and methods

﻿Collection, morphological examination and isolation

﻿DNA extraction, PCR amplification, and sequencing

﻿Phylogenetic analyses

﻿Results

﻿Phylogenetic analyses

﻿Taxonomy

﻿ Aquapteridospora linzhiensis R.J. Xu, Q. Zhao & Boonmee, sp. nov.

Etymology

Holotype

Description

Culture characteristics

Material examined

Notes

﻿ Aquapteridospora yadongensis R.J. Xu, Q. Zhao & Boonmee, sp. nov.

Etymology

Holotype

Description

Culture characteristics

Material examined

Notes

﻿ Aquapteridospora submersa R.J. Xu, Q. Zhao & Boonmee, sp. nov.

Etymology

Holotype

Description

Culture characteristics

Material examined

Notes

﻿Discussion

﻿Additional information

Conflict of interest

Ethical statement

Funding

Author contributions

Author ORCIDs

Data availability

﻿References

Abstract

Introduction

Materials and methods

Collection, morphological examination and isolation

DNA extraction, PCR amplification, and sequencing

Phylogenetic analyses

Results

Phylogenetic analyses

Taxonomy

Aquapteridospora linzhiensis R.J. Xu, Q. Zhao & Boonmee, sp. nov.

Aquapteridospora yadongensis R.J. Xu, Q. Zhao & Boonmee, sp. nov.

Aquapteridospora submersa R.J. Xu, Q. Zhao & Boonmee, sp. nov.

Discussion

Additional information

References