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Research Article
Two novel hyphomycetes associated with ferns from China
expand article infoJing-Yi Zhang‡§, Kevin D. Hyde‡|, Li-Juan Zhang§‡, Song Bai, Dan-Feng Bao#, Fatimah Al-Otibi|, Yong-Zhong Lu#
‡ Mae Fah Luang University, Chiang Rai, Thailand
§ Guizhou Institute of Technology, Guiyang, China
| King Saud University, Riyadh, Saudi Arabia
¶ Guizhou Industry Polytechnic College, Guiyang, China
# Guizhou University, Guiyang, China

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

Corresponding author: Yong-Zhong Lu ( yzlu86@gmail.com )

Academic editor: Danushka Sandaruwan Tennakoon
© 2025 Jing-Yi Zhang, Kevin D. Hyde, Li-Juan Zhang, Song Bai, Dan-Feng Bao, Fatimah Al-Otibi, Yong-Zhong Lu.
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: Zhang J-Y, Hyde KD, Zhang L-J, Bai S, Bao D-F, Al-Otibi F, Lu Y-Z (2025) Two novel hyphomycetes associated with ferns from China. MycoKeys 113: 101-121. https://doi.org/10.3897/mycokeys.113.137678
Open Access

Abstract

During an ongoing investigation of fungi associated with ferns in southwestern China, three hyphomycetes were discovered on the dead rachises of Angiopteris fokiensis and an unidentified fern. Based on morphology and multi-gene phylogenetic analyses, Arthrobotrys angiopteridis and Corynespora septata are introduced as new species. Arthrobotrys angiopteridis is a nematode-trapping fungus characterized by macronematous, mononematous, hyaline conidiophores, conidiogenous cells with polyblastic denticles at each node, and 0–1-septate, clavate to elongate pyriform, hyaline conidia. Corynespora septata features macronematous, mononematous, pale brown to dark brown conidiophores, integrated, monotretic conidiogenous cells and up to 7-distoseptate with one true septum, subcylindrical to obclavate, hyaline to pale brown conidia. Detailed descriptions and illustrations of these two new species are provided, along with morphological comparisons of the new taxa with closely related species.

Key words

Asexual morph, new species, phylogeny, pteridophytes, taxonomy

Introduction

Fungi associated with ferns have historically been overlooked and have received insufficient research attention, despite being an immensely promising and diverse group (Kirschner and Liu 2014; Kirschner et al. 2019; Medel-Ortiz et al. 2020). Recent studies have provided evidence supporting this perspective, with many new species of fern-associated fungi being discovered (Guatimosim et al. 2016a; Guatimosim et al. 2016b; Li et al. 2023a; Wu and Diao 2023; Zhang et al. 2023b). China has the highest fern species, with about 2,130 species, accounting for 19% of the total global fern and fern-allies species (Lin et al. 2013; Zhou et al. 2016). Notably, the Yunnan region houses approximately 1,500 fern species, while Guizhou has about 800, ranking them as the first and third most diverse regions for fern species in China (Li et al. 2015; Zhou et al. 2016). These regions boast abundant and diverse fern resources, offering great potential for the discovery of even more interesting fungi (Zhang et al. 2023b; Hyde et al. 2024a; Phookamsak et al. 2024).

Arthrobotrys was introduced by Corda (1839), with A. superba Corda as the type species and belongs in Orbiliaceae, Orbiliales, Orbiliomycetes (Wijayawardene et al. 2022; Hyde et al. 2024b). Arthrobotrys is characterized by simple or branched conidiophores and obovoid, elliptic, pyriform, 0–3-septate conidia growing asynchronously on the nodes or short denticles of conidiophores (Yu et al. 2014; Zhang et al. 2022a, b, 2023a, 2024; Yang et al. 2023a; Jin et al. 2024). Arthrobotrys is the most complex and largest genus among Orbiliaceae nematode-trapping fungi, comprising 78 accepted species characterized by producing adhesive networks to capture nematodes (Li et al. 2005; Yang et al. 2012; Yu et al. 2014; Zhang and Hyde 2014; Jiang et al. 2017; Zhang et al. 2023a, 2024; Thiyagaraja et al. 2024). These fungal species mainly occur in soil or sediment in various ecosystems such as farmland, forests, mangroves, and freshwater. They have also been recorded in hot springs, animal waste, and tree trunks worldwide (Duddington 1954; Swe et al. 2008; Kim et al. 2001; Kumar et al. 2011; Yu et al. 2014; Jiang et al. 2017; Zhang et al. 2022a, b).

Corynespora was established by Güssow (1905) with C. mazei Güssow as the type species. Corynespora was placed in Corynesporascaceae as the asexual morph associated with Corynesporasca, based on cultural studies (Sivanesan 1996), although the latter is still accepted as a distinct genus (Wijayawardene et al. 2022; Hyde et al. 2024b; Pem et al. 2024). Phylogenetic analyses demonstrated that Corynespora belongs to Corynesporascaceae, Pleosporales (Voglmayr and Jaklitsch 2017; Hyde et al. 2024b; Thiyagaraja et al. 2024). The genus is characterized by distinct conidiophores; integrated, terminal, monotretic, determinate, or percurrently extending conidiogenous cells; and acrogenous, solitary or catenate, distoseptate conidia (Sharma and Chaudhary 2002; Pal et al. 2007; Voglmayr and Jaklitsch 2017; Capital and Lao 2020; Xu et al. 2020; Pem et al. 2024). Synopses of Corynespora species have been provided by Siboe et al. (1999), Kumar et al. (2021) and Xu et al. (2020). Subsequently, Liu et al. (2023) provided the latest list of identified and accepted species of Corynespora with major morphological features, host information, and locality data. Corynespora species have a wide distribution and can be found as saprobes, pathogens, and endophytes on living leaves, or from decaying woody material of various plants, as well as on other fungi, nematodes, and human skin (Furukawa et al. 2008; Dixon et al. 2009; Kumar and Singh 2016; Capital and Lao 2020; Xu et al. 2020; Liu et al. 2022; Liu et al. 2023). Li et al. (2023b) then introduced a new species, on branches of Idesia polycarpa from Sichuan Province, China. A total of 213 epithets were listed under Corynespora (http://www.indexfungorum.org, accessed 20, September 2024), with 129 species being accepted (Liu et al. 2023).

In this study, collections representing two new species (Arthrobotrys angiopteridis and Corynespora septata) associated with ferns were made in Yunnan and Guizhou provinces in southwestern China. The identification and establishment of these taxa were based on morphological characteristics and phylogenetic evidence, a polyphasic approach, following the guidelines of Maharachchikumbura et al. (2021).

Material and methods

Collections, isolation and conservation

Samples of dead fern tissues were collected from Yunnan and Guizhou Provinces, China. The samples were packed in plastic bags for transportation to the laboratory, and subsequently examined using the methods described in Senanayake et al. (2020). A stereomicroscope (Leica EZ4 Microsystems (Schweiz) AG, Singapore) was used to examine and observe fungal colonies on the host surface. Morphological characteristics were documented using a Nikon DS-Ri2 digital camera fitted to a Nikon ECLIPSE Ni compound microscope (Nikon, Japan). Measurements of fungal structure were made using the Tarosoft (R) Image Frame Work, and the images used for figures were processed and combined in Adobe Illustrator CS6 (Adobe Systems, San Jose, CA, USA). Single spores were isolated following the method described by Chomnunti et al. (2014) to obtain pure cultures. Dried specimens were deposited in the Herbarium of Cryptogams, Kunming Institute of Botany, Academia Sinica (HKAS), Kunming, China, and the Herbarium of Guizhou Academy of Agricultural Sciences (GZAAS), Guiyang, China. Pure cultures were deposited in Kunming Institute of Botany Culture Collection (KUNCC), Kunming, China, and Guizhou Culture Collection, China (GZCC). Index Fungorum numbers (https://www.indexfungorum.org/Names/Names.asp) and Facesoffungi numbers (Jayasiri et al. 2015) are provided.

DNA Extraction, PCR amplification and sequencing

Fresh fungal mycelia were scraped from the surface of colonies grown on PDA, which had been incubated at 25 °C–28 °C for one month. Fungal genomic DNA was then extracted using the Biospin Fungus Genomic DNA Extraction Kit (BioFlux®, Shanghai, China). Four partial gene regions, the nuclear ribosomal internal transcribed spacer region (ITS: ITS1-5.8S-ITS2), the partial nuclear ribosomal large subunit rRNA gene (LSU) and the partial second‐largest subunit of the RNA polymerase II gene (rpb2), were amplified using polymerase chain reaction (PCR). The primers used were ITS5/ITS4 for ITS (White et al. 1990), LR0R/LR5 for LSU (Vilgalys and Hester 1990) and fRPB2-5F/fRPB2-7cR for rpb2 (Liu et al. 1999). The quality of the PCR products was checked on 1% agarose electrophoresis gels stained with ethidium bromide. Purification and sequencing of PCR products were performed by Beijing Qingke Biotechnology Co., Ltd.

Phylogenetic analyses

Original sequences were checked using BioEdit v. 7.1.3.0 (Hall 1999) and assembled using SeqMan v. 7.0.0 (DNASTAR, Madison, WI, USA). The newly assembled sequences were subjected to BLAST searches in NCBI-GenBank to preliminarily determine related taxa. The sequence data obtained from the BLAST search results and the latest publications (Li et al. 2023b; Liu et al. 2023; Jin et al. 2024; Zhang et al. 2024) were used for phylogenetic analyses. Alignments for sequences of each locus were performed with the online multiple alignment program MAFFT version 7 (https://mafft.cbrc.jp/alignment/server/, accessed on September 2024; Katoh et al. 2019), and the alignment files were further trimmed in trimAl version 1.2 (Capella-Gutiérrez et al. 2009) with the option “-gt 0.6”. Multi-gene alignments were combined using Sequence Matrix 1.7.8 (Vaidya et al. 2011). Sequences generated in this study were deposited in GenBank (Table 1 and Table 2).

Table 1.
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Taxa used in the phylogenetic analyses for Arthrobotrys genus, and their GenBank accession numbers.

Taxa Strain Number ITS tef1-α rpb2
Arthrobotrys amerospora CBS 268.83 NR_159625 N/A N/A
Arthrobotrys angiopteridis KUNCC 23-14121 PQ346307 N/A PQ356383
Arthrobotrys angiopteridis KUNCC 23-14119 PQ346306 N/A N/A
Arthrobotrys anomala YNWS02-5-1 AY773451 AY773393 AY773422
Arthrobotrys arthrobotryoides AOAC MF926580 N/A N/A
Arthrobotrys blastospora CGMCC 3.20940 OQ332405 OQ341651 OQ341649
Arthrobotrys botryospora CBS 321.83 NR_159626 N/A N/A
Arthrobotrys cibiensis DLUCC 109 OR880379 OR882792 OR882797
Arthrobotrys cibiensis EY10 OR902195 OR882787 OR882802
Arthrobotrys cladodes 1.03514 MH179793 MH179616 MH179893
Arthrobotrys clavispora CBS 545.63 MH858353 N/A N/A
Arthrobotrys conoides 670 AY773455 AY773397 AY773426
Arthrobotrys cookedickinson YMF 1.00024 MF948393 MF948550 MF948474
Arthrobotrys cystosporia CBS 439.54 MH857384 N/A N/A
Arthrobotrys dendroides YMF 1.00010 MF948388 MF948545 MF948469
Arthrobotrys dianchiensis 1.00571 MH179720 N/A MH179826
Arthrobotrys elegans 1.00027 MH179688 N/A MH179797
Arthrobotrys eryuanensis CGMCC 3.19715 MT612105 OM850307 OM850301
Arthrobotrys eudermata SDT24 AY773465 AY773407 AY773436
Arthrobotrys flagrans 1.01471 MH179741 MH179583 MH179845
Arthrobotrys gampsospora CBS 127.83 U51960 N/A N/A
Arthrobotrys globospora 1.00537 MH179706 MH179562 MH179814
Arthrobotrys gongshanensis CGMCC 3.23753 OM801277 OM809162 OM809163
Arthrobotrys guizhouensis YMF 1.00014 MF948390 MF948547 MF948471
Arthrobotrys heihuiensis DLUCC 108-1 OR880378 OR882791 OR882796
Arthrobotrys heihuiensis Y710 OR902194 OR882786 OR882801
Arthrobotrys hengjiangensis CGMCC 3.24983 OQ946587 OQ989312 OQ989302
Arthrobotrys hyrcanus IRAN 3650C MH367058 OP351540 N/A
Arthrobotrys indica YMF 1.01845 KT932086 N/A N/A
Arthrobotrys iridis 521 AY773452 AY773394 AY773423
Arthrobotrys janus Jan-85 AY773459 AY773401 AY773430
Arthrobotrys javanica 105 EU977514 N/A N/A
Arthrobotrys jindingensis CGMCC 3.20895 OP236810 OP272511 OP272515
Arthrobotrys jinpingensis CGMCC 3.20896 OM855569 OM850311 OM850305
Arthrobotrys jinshaensis DLUCC 133 OR880381 OR882794 OR882799
Arthrobotrys jinshaensis MA142 OR902197 OR882789 OR882804
Arthrobotrys koreensis C45 JF304780 N/A N/A
Arthrobotrys lanpingensis CGMCC 3.20998 OM855566 OM850308 OM850302
Arthrobotrys latispora H.B. 8952 MK493125 N/A N/A
Arthrobotrys longiphora 1.00538 MH179707 N/A MH179815
Arthrobotrys lunzhangensis CGMCC 3.20941 OK643973 OM621809 OM621810
Arthrobotrys luquanensis CGMCC 3.20894 OM855567 OM850309 OM850303
Arthrobotrys mangrovispora MGDW17 EU573354 N/A N/A
Arthrobotrys megalospora TWF800 MN013995 N/A N/A
Arthrobotrys microscaphoides YMF 1.00028 MF948395 MF948552 MF948476
Arthrobotrys multiformis CBS 773.84 MH861834 N/A N/A
Arthrobotrys musiformis SQ77-1 AY773469 AY773411 AY773440
Arthrobotrys musiformis 1.03481 MH179783 MH179607 MH179883
Arthrobotrys nonseptata YMF 1.01852 FJ185261 N/A N/A
Arthrobotrys obovata YMF 1.00011 MF948389 MF948546 MF948470
Arthrobotrys oligospora 920 AY773462 AY773404 AY773433
Arthrobotrys paucispora ATCC 96704 EF445991 N/A N/A
Arthrobotrys polycephala 1.01888 MH179760 MH179592 MH179862
Arthrobotrys pseudoclavata 1130 AY773446 AY773388 AY773417
Arthrobotrys psychrophila 1.01412 MH179727 MH179578 MH179832
Arthrobotrys pyriformis YNWS02-3-1 AY773450 AY773392 AY773421
Arthrobotrys reticulata CBS 550.63 MH858355 N/A N/A
Arthrobotrys robusta nefuA4 MZ326655 N/A N/A
Arthrobotrys salina SF 0459 KP036623 N/A N/A
Arthrobotrys scaphoides 1.01442 MH179732 MH179580 MH179836
Arthrobotrys shizishanna YMF 1.00022 MF948392 MF948549 MF948473
Arthrobotrys shuifuensis CGMCC 3.19716 MT612334 OM850306 OM850300
Arthrobotrys sinensis 105-1 AY773445 AY773387 AY773416
Arthrobotrys sphaeroides 1.0141 MH179726 MH179577 MH179831
Arthrobotrys superba 127 EU977558 N/A N/A
Arthrobotrys thaumasia 917 AY773461 AY773403 AY773432
Arthrobotrys tongdianensis CGMCC 3.20942 OP236809 OP272509 OP272513
Arthrobotrys vermicola 629 AY773454 AY773396 AY773425
Arthrobotrys weixiensis CGMCC 3.24984 OQ946585 OQ989310 OQ989300
Arthrobotrys xiangyunensis YXY10-1 MK537299 N/A N/A
Arthrobotrys yangbiensis DLUCC 36-1 OR880382 OR882795 OR882800
Arthrobotrys yangbiensis Y678 OR902198 OR882790 OR882805
Arthrobotrys yangjiangensis DLUCC 124 OR880380 OR882793 OR882798
Arthrobotrys yangjiangensis YB19 OR902196 OR882788 OR882803
Arthrobotrys yunnanensis YMF 1.00593 AY50993 N/A N/A
Arthrobotrys zhaoyangensis CGMCC 3.20944 OM855568 OM850310 OM850304
Dactylellina cangshanensis CGMCC 3.19714 MK372062 MN915115 MN915114
Dactylellina copepodii CBS 487.90 U51964 DQ999835 DQ999816

Note: “N/A” indicates no data are available in GenBank. The newly generated sequences are indicated in blue.
Table 2.
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Taxa used in the phylogenetic analyses for Corynespora genus, and their GenBank accession numbers.

Taxa Strain Number ITS LSU
Corynespora cassiicola CBS 100822 N/A GU301808
Corynespora citricola CBS 169.77 FJ852594 N/A
Corynespora doipuiensis MFLUCC 14-0022 MN648322 MN648326
Corynespora encephalarti CBS 145555 MK876383 MK876424
Corynespora lignicola MFLUCC 16–1301 MN860549 MN860554
Corynespora mengsongensis HJAUP C2000T OQ060574 OQ060578
Corynespora nabanheensis HJAUP C2048T OQ060577 OQ060580
Corynespora pseudocassiicola CPC 31708 MH327794 MH327830
Corynespora septata GZCC 23-0741 PQ346308 PQ346311
Corynespora smithii L120 KY984297 KY984297
Corynespora smithii L130 KY984298 KY984298
Corynespora smithii CABI 5649b FJ852597 GU323201
Corynespora smithii CBS 139925 KY984299 KY984299
Corynespora submersa MFLUCC 16–1101 MN860548 MN860553
Corynespora torulosa CBS 136419 MH866095 MH877634
Corynespora thailandica CBS 145089 MK047455 MK047505
Corynespora yunnanensis HJAUP C2132T OQ060579 OQ060583
Periconia byssoides H 4600 LC014581 AB807570
Periconia digitata CBS 510.77 LC014584 AB807561
Periconia pseudodigitata KT 1395 NR_153490 NG_059396
Periconia pseudodigitata UESTCC 23.0022 OR253146 OR253305
Periconia pseudodigitata UESTCC 23.0023 OR253147 OR253306

Note: “N/A” indicates no data are available in GenBank. The newly generated sequences are indicated in blue.

The fasta files were converted to the formats required for the AliView program (Larsson 2014), PHYLIP for maximum likelihood analysis (ML), and NEXUS for Bayesian analysis (BYPP). Maximum likelihood (ML) analyses were performed using RAxML-HPC Blackbox (8.2.10) tool on the XSEDE Teragrid at the CIPRES Science Gateway (https://www.phylo.org; accessed on 10 September 2024), with rapid bootstrap analysis followed by 1,000 bootstrap replicates (Miller et al. 2010; Stamatakis 2014). The final tree was selected from the suboptimal trees of each run by comparing likelihood scores under the GTRGAMMA substitution model. Bayesian analyses were performed by MrBayes 3.2.7a on XSED via CIPRES (Miller et al. 2010). MrModeltest v.2.3 was used to determine the best nucleotide substitution model for each data partition (Nylander 2004). Posterior probabilities (PP) (Rannala and Yang 1996) were calculated using the Bayesian Markov Chain Monte Carlo (BMCMC) sampling method (Huelsenbeck 2001; Zhaxybayeva and Gogarten 2002). Four simultaneous Markov chains were run for 1 million generations, with trees sampled every 100th generations, yielding 10,000 trees. Phylogenetic trees were visualized using FigTree v. 1.4.4 (Rambaut 2014), and the layouts were created using Adobe Illustrator CS5 software (Adobe Systems, San Jose, CA, USA). The newly obtained sequences in this study were deposited in GenBank.

Taxonomy

Arthrobotrys angiopteridis J.Y. Zhang, Y.Z. Lu & K.D. Hyde, sp. nov.

MycoBank No: 902682
Facesoffungi number: FoF16618
Fig. 2

Etymology

Named after the fungal host genus Angiopteris.

Holotype

HKAS 129855.

Description

Saprobic on dead rachis of Angiopteris fokiensis in terrestrial habitats. Sexual morph Undetermined. Asexual morph Colonies on natural substrate superficial, effuse, hyaline, with white and glistening masses of conidia on the apex of conidiophores. Mycelium partly superficial, partly immersed, composed of septate, branched, smooth hyphae. Conidiophores 345–502 µm long, 6–8.5 µm wide at the base (x̄ = 418 × 6.9 µm, n = 20), macronematous, mononematous, solitary, erect, straight or slightly flexuous, unbranched, cylindrical, septate, smooth-walled, hyaline. Conidiogenous cells 95–176 × 2–4.5 µm (x̄ = 129 × 3.5 µm, n = 20), polyblastic, producing 1–5 separate nodes by the repeated elongation, with multi polyblastic denticles at each node, hyaline. Conidia 25–35 × 8–11 µm (x̄ = 28.8 × 9 µm, n = 25), aseptate, or 1-septate at the median to submedian, not constricted or slightly constricted at the septum, clavate to elongate pyriform, broadly rounded at apex, pointed or sometimes truncate at the base, sometimes with a bud-like projection at base, straight or slightly curved, smooth-walled or rough walled, guttulate, hyaline.

Culture characteristics

Conidia germinating on WA within 15 h and germ tube produced from conidia. Colonies growing on PDA, reaching 60 mm diameter in 10 days at 26 °C, circular, cottony, white, and not producing pigmentation in culture.

Material examined

China • Guizhou Province, Zunyi City, Xishui County (28°22'19"N, 106°0'35"E), on dead rachis of Angiopteris fokiensis (Marattiaceae) in a disturbed forest nearby the roadside, 13 April 2023, J.Y. Zhang, ZY06 (HKAS 129855, holotype; GZAAS 23–0758, isotype), ex-type living culture, KUNCC 23–14121; • ibid., ZY02 (HKAS 129854, paratype), ex-paratype living culture, KUNCC 23–14119. Additional sequence: KUNCC 23–14121: PQ346313 (SSU) and PQ346310 (LSU); KUNCC 23–14119: PQ346312 (SSU) and PQ346309 (LSU).

Notes

Phylogenetically, the new isolates KUNCC 23–14121 and KUNCC 23–14119 of Arthrobotrys angiopteridis clustered together formed a separate clade with 100% ML/1.00 PP bootstrap support and are sister to A. pyriformis (Fig. 1). A comparison of nucleotide base pairs between them reveals differences of 30/459 (6.5%, including 15 gaps) and 82/730 bp (11%, no gap) in the ITS and rpb2 sequences, respectively. This indicates that they are distinct species. Morphologically, A. angiopteridis aligns well with the generic concept and resembles A. oligospora in having hyaline conidiophores with the successive production of additional denticle nodes (Yu et al. 2014). However, A. angiopteridis can be easily distinguished from A. oligospora by its longer conidiophores (345–502 µm vs. 110–440 μm) and clavate to elongate pyriform conidia, with 0–1 septate near the middle, whereas A. oligospora has pyriform or obovoid conidia with 1-septate near the base. Therefore, we introduce A. angiopteridis as a novel species based on its distinct morphological and phylogenetic evidence following the guidelines of Maharachchikumbura et al. (2021).

Figure 1. 

Phylogram generated from maximum likelihood analysis based on combined ITS, tef1-α and rpb2 sequence data. Seventy-eight taxa were included in the combined analyses, which comprised 1920 characters (ITS = 583 bp, tef1-α = 512 bp and rpb2 = 825 bp) after alignment. The best scoring RAxML tree with a final likelihood value of -22800.405782 is presented. The matrix had 983 distinct alignment patterns, with 25.05% of undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.260557, C = 0.263561, G = 0.230377, T = 0.245504; substitution rates: AC = 1.414531, AG = 3.978691, AT = 1.319991, CG = 0.945884, CT = 6.618473, GT = 1.000000; gamma distribution shape parameter α = 0.262034. Bootstrap support values for ML equal to or greater than 60% and prior probabilities (PPs) equal to or greater than 0.95 are given above the nodes as ML/PP. The tree was rooted to Dactylellina copepodii (CBS 487.90) and D. cangshanensis (CGMCC 3.19714). The strain numbers are noted after the species names with ex-type strains indicated by T. The newly generated sequences are indicated in blue.

Corynespora septata J.Y. Zhang, Y.Z. Lu & K.D. Hyde, sp. nov.

MycoBank No: 902683
Facesoffungi number: FoF16619
Fig. 4

Etymology

Named after the presence of eu-septate conidia.

Holotype

HKAS 129839.

Description

Saprobic on dead rachis of an unidentified fern in terrestrial habitats. Sexual morph undetermined. Asexual morph Colonies on natural substrate superficial, effuse, gregarious, hairy, brown to black. Mycelium partly superficial, partly immersed, composed of branched, septate, pale brown to brown, smooth-walled hyphae. Conidiophores 490–671 µm long, 3.5–6.5 µm wide at the base (x̄ = 600 × 5 µm, n = 15), macronematous, mononematous, erect, straight or flexible, unbranched, or occasionally branched, septate, smooth, dark brown at the base, pale towards the apex. Conidiogenous cells 21–60 × 3–5.5 µm (x̄ = 36.3 × 3.8 µm, n = 15), integrated, terminal, monotretic, cylindrical, smooth, pale brown to brown. Conidia 42–74 × 4.5–7.5 µm (x̄ = 54 × 5.7 µm, n = 25), acrogenous, solitary, up to 7-distoseptate with one true septum, straight or slightly curved, subcylindrical to obclavate, rounded at the apex, base short obconically truncate, somewhat thickened and darkened, sometimes with percurrent proliferation which forms another conidium from the conidial apex, hyaline to pale brown.

Culture characteristics

Conidia germinating on WA within 15 h and germ tube produced from conidia. Colonies growing on PDA, reaching 55 mm diameter in 10 days at 26 °C, circular, flat with entire margin, velvety, fluffy, white from above, reverse dark brown at center, paler to light yellow towards margin, and not producing pigmentation in culture.

Material examined

China • Yunnan Province, Xishuangbanna Dai Autonomous Prefecture, Mengla County, Menglun Town, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences (21°55'39"N, 101°15'15"E), on dead rachis of an unidentified fern, 16 November 2019, J.Y. Zhang, Y159 (HKAS 129839, holotype; GZAAS 23–0769, isotype), ex-type living culture, GZCC 23–0741.

Notes

A BLASTn search in NCBI-GenBank revealed that the LSU and ITS sequences of our newly collected strain of Corynespora septata exhibited 99% similarity to C. encephalarti (NG_067878) and 95.62% similarity to C. cassiicola (MN648322), respectively. Phylogenetic analysis confirmed that C. septata formed a distinct clade within Corynespora and shared a sister relationship with C. pseudocassiicola Crous & M.J. Wingf. (Fig. 3). There are 10 bp (10/841 bp with 0 gap, 1%) and 38 bp (38/527 bp with 13 gaps, 7%) differences between the C. pseudocassiicola and C. septata in the LSU and ITS gene regions, respectively. Morphologically, C. septata has longer conidiophores (490–671 µm vs. 200–400 µm), and smaller conidia (42–74 × 4.5–7.5 µm vs. 95–160 × 9–10 µm) compared to C. pseudocassiicola (Crous et al. 2018). Similarly, C. septata is most similar to C. lignicola Z.L. Luo, H.Y. Su & K.D. Hyde in the shapes of conidiophores, conidiogenous cells, and conidia (Capital and Lao 2020). However, C. septata differs from C. lignicola in having narrower conidiophores (3.5–6.5 µm vs. 9–13 µm) and notably smaller conidia (42–74 × 4.5–7.5 µm vs. 110–156 × 7–9 µm).

Discussion

During a survey of bracken (Pteridiurn aquilinum (L.) Kuhn) petiole decomposition in the United Kingdom, Arthrobotrys megalosporus (Drechsler) M. Scholler, Hagedorn & A. Rubner (Synonym: Dactylella megalospora Drechsler) was found to be a member of the common fungi (Frankland 1976). This is also the only record of Arthrobotrys species being associated with ferns. Corynespora cassiicola (Berk. & M.A. Curtis) C.T. Wei and an unidentified Corynespora species have been discovered on ferns in United States (Alfieri et al. 1984; Smith 2008; Farr et al. 2021). Specifically, C. cassiicola has been found on six fern species including Arachniodes aristata (Davalliaceae), Athyrium niponicum (Dryopteridaceae), Adiantum cuneatum (Adiantaceae), Adiantum tenerum (Adiantaceae), Davallia repens (Davalliaceae), and Platycerium spp. (Pteridaceae) (Alfieri et al. 1984; Smith 2008). Additionally, an unidentified Corynespora species was collected from Nephrolepis exaltata (Davalliaceae) (Alfieri et al. 1984; Farr et al. 2021). Based on phylogenetic and morphological evidence, Arthrobotrys angiopteridis and Corynespora septata, isolated from ferns are reported as new species in this study from Yunnan and Guizhou provinces. These findings contribute to a better understanding of fern-related fungi and aim to enhance attention and awareness of fungal communities associated with ferns.

Most Corynespora species were introduced based on morphology (Sivanesan 1996; Siboe et al. 1999; Sharma and Chaudhary 2002; Kumar and Singh 2016; Liu et al. 2023). However, distinguishing between Corynespora and some similar genera, especially Helminthosporium, based solely on morphology has proven to be challenging (Castañeda Ruiz et al. 2004; Voglmayr and Jaklitsch 2017). The application of molecular data has confirmed this difficulty. For example, Corynespora caespitosa, C. endiandrae, C. leucadendri and C. olivacea were transferred to Helminthosporium based on phylogenetic evidence (Voglmayr and Jaklitsch 2017). Currently, sequence data is available in GenBank for only 15 species. Therefore, it is essential to obtain more collections with sequence data for verification of Corynespora species.

Arthrobotrys angiopteridis sp. nov., isolated from Angiopteris fokiensis, is a member of nematode-trapping fungi with trapping device of adhesive networks (Fig. 2l). Nematode-trapping fungi are crucial for preserving ecological balance and possess the potential for biologically controlling harmful nematodes (Jiang et al. 2017; Zhang et al. 2024). Arthrobotrys angiopteridis is a valuable fungus that is expected to contribute to the exploration of ecological protection in the future.

Figure 2. 

Arthrobotrys angiopteridis (HKAS 129855, holotype) a the host b colonies on the host c, d conidiophores with conidiogenous cells e–h conidiogenous cells i–k conidia l trapping mycelia: adhesive networks m pure culture from front and reverse. Scale bar: 100 µm (c, d); 20 µm (e–l).

Figure 3. 

Phylogram generated from maximum likelihood analysis based on combined LSU and ITS sequence data. Twenty-two taxa were included in the combined analyses, which comprised 1393 characters (LSU = 844 bp and ITS = 549 bp) after alignment. The best scoring RAxML tree with a final likelihood value of -4677.993509 is presented. The matrix had 336 distinct alignment patterns, with 9.65% of undetermined characters or gaps. Estimated base frequencies were as follows: A = 0.243499, C = 0.247400, G = 0.289966, T = 0.219136; substitution rates: AC = 3.067252, AG = 2.397685, AT = 1.551875, CG = 1.069828, CT = 6.624253, GT = 1.000000; gamma distribution shape parameter α = 0.240919. Bootstrap support values for ML equal to or greater than 60% and prior probabilities (PPs) equal to or greater than 0.95 are given above the nodes as ML/PP. The tree was rooted to Periconia byssoides (H 4600), P. digitata (CBS 510.77) and P. pseudodigitata (KT 1395). The strain numbers are noted after the species names with ex-type strains indicated by T. The newly generated sequences are indicated in blue.

Figure 4. 

Corynespora septata (HKAS 129839, holotype) a, b colonies on the host c–e conidiophores with conidiogenous cells f–k conidia l pure culture from front and reverse. Scale bar: 200 µm (b); 100 µm (c–e); 20 µm (f–k).

Yunnan and Guizhou provinces are not only the most abundant areas for fern plants in China (Li et al. 2015; Zhou et al. 2016), but also hotspots for the discovery of new fungal species (Wang et al. 2021; Yang et al. 2023b; Zhang et al. 2023b; Dissanayake et al. 2024). The introduction of A. angiopteridis and C. septata adds to the growing evidence of high fungal diversity in Guizhou and Yunnan province, China (Dissanayake et al. 2024; Dong et al. 2024; Li et al. 2024).

Acknowledgements

We would like to thank Shaun Pennycook (Manaaki Whenua Landcare Research, New Zealand) for advising on the fungal names and Yu Yang for helping in experiment. Jing-Yi Zhang thanks the Mae Fah Luang University for granting her the tuition scholarship and the dissertation writing grant (Grant number: 7702(6)/842 (no.0320). Dan-Feng Bao would like to thank the Postdoctoral Fellowship Program of CPSF under Grant Number GZC20240346.

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 National Natural Science Foundation of China (NSFC 32060013) and the Youth Science and Technology Talent Development Project from Guizhou Provincial Department of Education (QJHKYZ[2022]345). The authors also extend their appreciation to the Researchers Supporting Project number (RSP2024R114), King Saud University, Riyadh, Saudi Arabia.

Author contributions

Conceptualization: KDH, JYZ. Data curation: JYZ. Formal analysis: DFB, JYZ, LJZ. Funding acquisition: YZL, LJZ. Investigation: JYZ. Methodology: JYZ, FAO. Project administration: YZL. Supervision: KDH. Writing - original draft: JYZ. Writing - review and editing: SB, DFB, FAO, KDH.

Author ORCIDs

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

Kevin D. Hyde https://orcid.org/0000-0002-2191-0762

Li-Juan Zhang https://orcid.org/0000-0002-3234-6757

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

Dan-Feng Bao https://orcid.org/0000-0002-5697-4280

Fatimah Al-Otibi https://orcid.org/0000-0003-3629-5755

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

Data availability

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

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