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Research Article
Morphophylogenetic evidence reveals four new fungal species within Tetraplosphaeriaceae (Pleosporales, Ascomycota) from tropical and subtropical forest in China
expand article infoXia Tang§, Rajesh Jeewon|, Yong-Zhong Lu#, Abdulwahed Fahad Alrefaei|, Ruvishika S. Jayawardena§, Rong-Ju Xu§, Jian Ma§, Xue-Mei Chen¤, Ji-Chuan Kang
‡ Guizhou University, Guiyang, China
§ Mae Fah Luang University, Chiang Rai, Thailand
| King Saud University, Riyadh, Saudi Arabia
¶ University of Mauritius, Reduit, Mauritius
# Guizhou Institute of Technology, Guiyang, China
¤ Qujing Normal University, Qujing, China
Open Access

Abstract

Tetraplosphaeriaceae (Pleosporales, Ascomycota) is a family with many saprobes recorded from various hosts, especially bamboo and grasses. During a taxonomic investigation of microfungi in tropical and subtropical forest regions of Guizhou, Hainan and Yunnan provinces, China, several plant samples were collected and examined for fungi. Four newly discovered species are described based on morphology and evolutionary relationships with their allies inferred from phylogenetic analyses derived from a combined dataset of LSU, ITS, SSU, and tub2 DNA sequence data. Detailed illustrations, descriptions and taxonomic notes are provided for each species. The four new species of Tetraplosphaeriaceae reported herein are Polyplosphaeria guizhouensis, Polyplosphaeria hainanensis, Pseudotetraploa yunnanensis, and Tetraploa hainanensis. A checklist of Tetraplosphaeriaceae species with available details on their ecology is also provided.

Key words

Anamorphic fungi, checklist, Dothideomycetes, ribosomal genes, species diversity, taxonomy

Introduction

The Southwestern part of China is characterised by a tropical to subtropical climate and several provinces are well known for their high diversity of plants as well as fungi (Feng and Yang 2018; Hyde et al. 2020b; Bao et al. 2021; Yang et al. 2023). Yunnan province, for example, is considered a hotspot for species diversity. Over the last few decades, there has been a number of studies that have reported novel fungal species from this region (Jeewon et al. 2003; Luo et al. 2017, 2018; Huang et al. 2018; Su et al. 2018; Yang et al. 2019; Hyde et al. 2020b; Bao et al. 2021; Mortimer et al. 2021). So far more than 6000 fungal species described alone from Yunnan province (Feng and Yang 2018). Guizhou, as a prominent example of China’s karst landform (also referred to as a ‘karst province’), is also characterised by a geomorphological diversity that can be directly related to species diversity. It boasts a distinctive geographical environment and a special climate that fosters the growth of numerous rare, endangered, and indigenous plant, animal, and fungal species. Over the past few decades, extensive research has focused on fungi, encompassing both macro and microfungi, leading to the identification and documentation of roughly over 2,500 fungal species in Guizhou province (Zhou et al. 2018, 2020a, 2020b, 2022; Dissanayake et al. 2020; Chen et al. 2021; Yang et al. 2023). Hainan Province, the largest island in the Indo-Burma biodiversity hotspot, contains extensive and well-preserved tropical forests (Huang et al. 2023). Recent studies have indicated the presence of diverse fungal species in Hainan, with roughly over 1000 fungal species. Most of these fungi are macrofungi (Zhang et al. 1994; Li et al. 2010; Hapuarachchi et al. 2018; Huang et al. 2023). With our current fungal biodiversity estimates and given that mycologists anticipate many more species remain to be discovered especially in explored habitats, this research study has been undertaken to investigate microfungi in this region, and potentially discover new fungal species.

Tetraplosphaeriaceae was introduced by Tanaka et al. (2009) to accommodate the massarina-like species that produced tetraploa-like anamorphs in culture with Tetraplosphaeria as its type genus. Tetraplosphaeriaceae is known to be widely distributed on various hosts, with most species reported from bamboo or grasses (Poaceae) (Tanaka et al. 2009; Ariyawansa et al. 2015; Li et al. 2016), while Tetraploa species occur on diverse hosts (Hyde et al. 2013). Tetraplosphaeriaceae was first described to accommodate five genera: Polyplosphaeria Kaz. Tanaka & K. Hiray, Pseudotetraploa Kaz. Tanaka & K. Hiray, Quadricrura Kaz. Tanaka & K. Hiray, Tetraplosphaeria Kaz. Tanaka & K. Hiray, and Triplosphaeria Kaz. Tanaka & K. Hiray (Tanaka et al. 2009). Hyde et al. (2013) treated the type genus, Tetraplosphaeria as a synonym of Tetraploa according to nomenclatural priority. Ariyawansa et al. (2015) accepted Shrungabeeja in Tetraplosphaeriaceae based on morphological and molecular evidence from S. longiappendiculata. Later, Ernakulamia was accommodated in Tetraplosphaeriaceae based on morphology and phylogenetic analyses of combined ITS, LSU and tub2 sequence data (Delgado et al. 2017; Hyde et al. 2020a). Pem et al. (2019) transferred Byssolophis from the genera incertae sedis to Tetraplosphaeriaceae based on its massarina-like morphology and phylogenetic analyses based on combined LSU, SSU, tef1-α, and rpb2 sequence data. Hongsanan et al. (2020) provided a taxonomic update on families of Dothideomycetes and eight genera were accepted in Tetraplosphaeriaceae. Recently, Li et al. (2021) discovered a freshwater fungus that had a close phylogenetic affinity with Ernakulamia and Shrungabeeja in Tetraplosphaeriaceae and accommodated it in a new genus Aquatisphaeria based on morphology and phylogeny. To date, Tetraplosphaeriaceae consists of nine genera (Hongsanan et al. 2020; Li et al. 2021; Wijayawardene et al. 2022).

Most members of Tetraplosphaeriaceae contain anamorphic species (Wijayawardene et al. 2022). However, Pseudotetraploa, Tetraploa, and Triplosphaeria exhibit both teleomorphs and anamorphs (Wijayawardene et al. 2022), while Byssolophis is only known in its teleomorphic morph (Tanaka et al. 2009; Ariyawansa et al. 2015; Pem et al. 2019; Hongsanan et al. 2020; Li et al. 2021; Jayawardena et al. 2023). Tetraplosphaeriaceae is characterized by massarina-like teleomorph morphs but can be distinguished from other families by its immersed to superficial, glabrous or brown hyphae at sides of ascomata with flattened bases and cylindrical to clavate, short pedicellate 8-spored asci which are narrowly fusiform to broadly cylindrical, septate, hyaline to pale brown ascospores, usually with a complete sheath or appendage-like sheath (Tanaka et al. 2009; Hyde et al. 2013). The anamorphs are tetraploa-like hyphomycetes having micronematous to macronematous, erect, unbranched, septate, with presence or absence of conidiophores, monoblastic, terminal conidiogenous cells sometimes indistinguishable from creeping hyphae, solitary, cylindrical to obpyriform, comprising 3–8 columns or internal hyphal structure conidia, mostly verrucose at the conidial base and with 2–8-setose appendages (Tanaka et al. 2009; Hyde et al. 2013; Hongsanan et al. 2020).

In this study, the aim is to characterize anamorphic fungal species collected from the southern part of China. The objectives are to 1) to describe novel species collected from Guizhou, Hainan, and Yunnan provinces in China, based on morphological examination of fresh specimens; 2) to document morphological differences and similarities with extant species; 3) to establish four new species within the family Tetraplosphaeriaceae with support from results generated from phylogenetic analyses of LSU, ITS, SSU, and tub2 DNA sequence data; 4) to provide a worldwide checklist of Tetraplosphaeriaceae species with available details on their ecology.

This study will undoubtedly increase our understanding of fungal diversity in China.

Materials and methods

Sample collection, isolation and morphological studies

Fresh samples of unidentified decaying wood and decaying bamboo were collected in Guizhou (Xingyi city, Xianheping National Forest Park), Hainan (Wuzhishan city, Wuzhishan National Nature Reserve), and Yunnan (Puer city, Ailao mountains) provinces respectively. During the collection period, the environmental conditions at the different regions were as follows: Guizhou-average temperature of 26 °C, subtropical climate, humid environment during autumn; Hainan-average temperature of 29 °C, tropical climate, humid environment during autumn; Yunnan-average temperature of 22 °C, subtropical climate, humid environment during spring. The samples were placed in Ziplock bags, labelled with a marker pen, and observed using the stereomicroscope (Motic SMZ-171). The procedure for specimen collection, observation and isolation follows that of Senanayake et al. (2020) and Tang et al. (2022). The morphological measurements were performed by the Tarosoft (R) Image Frame Work tool (IFW 0.97 version), and photoplates were created using the Adobe Photoshop 2019 program (Adobe Systems, USA).

After morphological examination, the specimens were deposited at the herbaria of Kunming Institute of Botany, Chinese Academy of Sciences (HKAS), Kunming, China, and the Guizhou Academy of Agriculture Sciences (GZAAS), Guiyang, China, respectively. The ex-type cultures were deposited at the Guizhou Culture Collection (GZCC) in China and the Kunming Institute of Botany Culture Collection (KUNCC). Faces of Fungi and Index Fungorum numbers are provided as in Jayasiri et al. (2015) and Index Fungorum (2023). Species recognition and justifications for new species establishment were done based on the guide-lines provided by Jeewon and Hyde (2016), Chethana et al. (2021) and Pem et al. (2021).

DNA extraction, PCR amplification and sequencing

Fresh mycelium was scraped from the living culture and transferred to 1.5 mL microcentrifuge tubes and kept in a refrigerator at -20 °C. Total genomic DNA was extracted using the DNA extraction kits (Sangon Biotech (Shanghai) Co. Ltd., China). DNA template amplifications were performed by Polymerase Chain Reaction (PCR) using primer pairs, ITS5/ITS4 for ITS (White et al. 1990), NS1/NS4 for SSU (White et al. 1990), LR0R/LR5 for LSU (Vilgalys and Hester 1990, Cubeta et al. 1991), and BT1/BT2b for tub2 (Glass and Donaldson 1995). For other details pertaining to DNA extraction, PCR amplifications, sequencing, and phylogenetic analyses, see Tang et al. (2022). The polymerase chain reaction was carried out in a volume of 50 μL, and the reagents that were used were as follows: DNA template (2 μL), forward primers (2 μL), reverse primers (2 μL), 2 ×Taq PCR Master Mix (25 μL) and 19 μL of ddH2O (double-distilled water). PCR profiles are as follows: 35 cycles, and the annealing temperatures for each gene are 52 °C for 1 minute and extension at 72 °C for 90 seconds in LSU, ITS and SSU; and 55 °C for 50 s and elongation at 72 °C for 1 minute for tub2. Verification of PCR products was done on 1% agarose gels before being sent to China’s Sangon Biotech (Shanghai) Co., Ltd. for sequencing.

Phylogenetic analyses

The forward and reverse primers of the newly generated sequence were assembled by the Contig Ex-press v3.0.0 application, and the most similar taxa were found by BLASTn (https://blast.ncbi.nlm.nih.gov/Blast.cgi) in NCBI. A combination of different DNA sequence data (LSU, ITS, SSU, and tub2), which are close hits and similar to other Tetraplosphaeriaceae species in GenBank (Table 1), were downloaded to be further analysed along with our new taxa. Each sequence data was aligned by the online version of MAFFT v. 7 (https://mafft.cbrc.jp/alignment/server/index.html) through the “auto” option (Katoh et al. 2017). Multiple genes were assembled by SequenceMatrix (Vaidya et al. 2011). The aligned sequence was trimmed by trimAl v 1.2 with the ‘gappyout’ option (Capella-Gutiérrez et al. 2009). The phylogenetic analyses in this study were based on the maximum likelihood (ML), and Bayesian inference (BI) by using a combined sequence dataset of LSU, ITS, SSU, and tub2.

Table 1.

Taxa used in this study and their GenBank accession numbers for LSU, ITS, SSU and tub2 sequence data.

Taxa name Strain Numbers GenBank Accession Numbers
LSU SSU ITS tub2
Amniculicola immersa CBS 123083T NG_056964 NG_062796
A. parva CBS 123092T NG_056970 NG_016504
Aquatisphaeria thailandica MFLUCC 21–0025T MW890763 MW890967 MW890969
Aq. Thailandica DLUCC B151 MW890764 MW890968
Byssolophis sphaerioides IFRDCC 2053 GU301805 GU296140
Ernakulamia cochinensis MFLUCC 18–1237 MN913716 MT864326 MT627670
E. krabiensis MFLUCC 18–0237T MK347990 MK347880 MK347773
E. tanakae NFCCI 4615T MN937211 MN937229 MN938312
E. xishuangbannaensis KUMCC 17–0187T MH260314 MH260354 MH275080
Polyplosphaeria fusca KT 2124 AB524607 AB524466 AB524791 AB524853
Po. Fusca KT 1616T AB524604 AB524463 AB524789 AB524851
Po. guizhouensis GZCC 23–0598T OR438888 OR427327 OR449118
Po. Hainanensis GZCC 23–0599T OR438889 OR438285 OR427323 OR449115
Po. Hainanensis GZCC 23–0600 OR438890 OR427324
Po. Thailandica MFLUCC 15–0840T KU248767 KU248766
Po. nabanheensis KUMCC 16–0151T MH260312 MH260352 MH275078 MH412745
Po. pandanicola MFLUCC 17–2266T MH260313 MH260353 MH275079
Pseudotetraploa bambusicola CGMCC 3.20939T ON332933 ON332923 ON332915
Ps. bambusicola UESTCC 22.0005 ON332934 ON332924 ON332916
Ps. curviappendiculata JCM 12852T AB524608 AB524467 AB524792 AB524854
Ps. Javanica JCM 12854 AB524611 AB524470 AB524795 AB524857
Ps. Longissima JCM 12853T AB524612 AB524471 AB524796 AB524858
Ps. rajmachiensis NFCCI 4618T MN937204 MN937222
Ps. Yunnanensis KUNCC 10464T OR438891 OR449073
Quadricrura bicornis CBS 125427T AB524613 AB524472 AB524797 AB524859
Q. meridionalis CBS 125684T AB524614 AB524473 AB524798 AB524860
Q. septentrionalis CBS 125429 AB524615 AB524474 AB524799 AB524861
Shrungabeeja aqutica MFLUCC 18–0664T MT627663 MT627722
S. longiappendiculata BCC 76463T KT376472 KT376471 KT376474
S. longiappendiculata BCC 76464 KT376473 KT376475
S. fluviatilis GZCC 20–0505T
S. fluviatilis GZCC 19–0511 MW133853 MW134631
S. vadirajensis MFLUCC 17–2362 MN913685 MT627681
Tetraploa aquatica MFLU 19–0996 MT530453 MT530454 MT530449
T. aquatica MFLUCC 19–0995T MT530452 MT530448
T. aristata CBS 996.70 AB524627 AB524486 AB524805 AB524867
T. bambusae KUMCC 21–0844T ON077067 ON077073 ON077078 ON075065
T. dwibahubeeja NFCCI 4621T MN937207 MN937225 MN938308
T. dwibahubeeja NFCCI 4623 MN937208 MN937226 MN938309
T. endophytica CBS 147114T MW659165 KT270279
T. hainanensis GZCC 23–0601T OR438892 OR438286 OR427325 OR449116
T. hainanensis GZCC 23–0602 OR438893 OR427326 OR449117
T. juncicola CBS 149046 ON603800 ON603780
T. nagasakiensis KUMCC 18–0109 MK079891 MK079888 MK079890
T. nagasakiensis KT 1682T AB524630 AB524489 AB524806 AB524868
T. pseudoaristata NFCCI 4624T MN937214 MN937232 MN938315
T. pseudoaristata NFCCI 4625 MN937212 MN937230 MN938313
T. puzheheiensis MFLUCC 20–0151T MT627655 MT627744
T. sasicola KT 563T AB524631 AB524490 AB524807 AB524869
T. thailandica MFLUCC 21–0030T MZ412530 MZ413274 MZ412518
T. thrayabahubeeja NFCCI 4627T MN937217 MN937235 MN938318
T. thrayabahubeeja NFCCI 4628 MN937215 MN937233 MN938316
T. yunnanensis MFLUCC 19–0319T MN913735 MT864341 MT627743
T. yakushimensis KT 1906T AB524632 AB524491 AB524808 AB524870
T. cylindrica KUMCC 20–0205T MT893204 MT893203 MT893205 MT899417
T. cylindrica ZHKUCC 22–0087 ON555688 ON555690 ON555689 ON564477
T. dashaoensis KUMCC 21–0010T OL473555 OL473556 OL473549 OL505601
T. obpyriformis KUMCC 21–0011T OL473554 OL473557 OL473558 OL505600
Tetraploa sp.1 KT 1684 AB524628 AB524487
Tetraploa sp.2 KT 2578 AB524629 AB524488
T. tetraploa CY112 HQ607964
Triplosphaeria acuta KT 1170T AB524633 AB524492 AB524809 AB524871
Tr. Cylindrica NBRC 106247 AB524636 AB524495 AB524811 AB524873
Tr. cylindrica KT 1800 AB524635 AB524494 AB524810 AB524872
Tr. maxima KT 870T AB524637 AB524496 AB524812 AB524874
Tr. yezoensis KT 1715T AB524638 AB524497 AB524813 AB524875
Tr. yezoensis KT 1732 AB524639 AB524498 AB524814 AB524876
Triplosphaeria sp. HHUF 27481 AB524640 AB524499 AB524815 AB524877
Triplosphaeria sp. KT 2546 AB524641 AB524500 AB524816 AB524878

Analyses under different criteria such as maximum likelihood (ML) and Bayesian inference (BI) were processed in the CIPRES web portal (Miller et al. 2010) by using the “RAxML-HPC v.8 on XSEDE” tool, and the tool “MrBayes on XSEDE”, respectively (Huelsenbeck and Ronquist 2001; Swofford 2002; Stamatakis et al. 2008; Ronquist et al. 2012).

For BI, MrModeltest v2 was used for the selection of the best-fit model for each gene region. The Markov Chain Monte Carlo (MCMC) algorithm was launched with four chains running concurrently from a random tree topology. The burn-in factor was set at 25%, and the sampling interval for trees was set to every 1000th generation. The Posterior Probabilities (PP) for the remaining trees were computed. Adobe Illustrator and FigTree were used to view trees. Bootstrap support and Bayesian posterior probabilities above 70 and 0.9 were considered as high support respectively.

Results

Phylogenetic analyses

For the phylogenetic analyses, a combined DNA sequence data of 68 taxa on LSU, ITS, SSU, and tub2 was used and analysed under the ML and PP criteria. The data matrix comprised 2995 total characters, including gaps (LSU: 1–848 bp, ITS: 849–1372 bp, SSU: 1373–2363 bp, tub2: 2364–2995 bp). Phylogenetic reconstructions with broadly comparable topologies were recovered from the combined dataset of ML and PP analyses. The top-scoring RAxML tree is shown in Fig. 1, with a final ML optimization likelihood value of -17569.286960 (ln). In the RAxML analysis, the GTRGAMMA+I-Invar model was used and the results showed 969 unique alignment patterns and 25.50% of indeterminate characters. Base frequency estimates were as follows: A = 0.243260, C = 0.247998, G = 0.277213, T = 0.231530; substitution rates were as follows: AC = 3.027135, AG = 4.828263, AT = 2.159193, CG = 1.385950, CT = 10.517436, GT = 1.000000; gamma distribution shape parameter alpha = 0.166037; and tree-length has been as follows: 1.837225. The best-fit models for the BPP analysis were GTR+I+G for LSU, ITS, and tub2 gene regions; HKY+I+G for the SSU gene region. With a final average standard deviation of split frequencies of 0.009909, Bayesian posterior probabilities from MCMC were analysed. The new taxa analysed herein all belong to the Tetraplosphaeriaceae clade based on the results of the combined LSU, ITS, SSU, and tub2 sequence data analysis.

Figure 1. 

Phylogenetic construction of Tetraplosphaeriaceae using RAxML-based maximum likelihood analysis of a combined LSU, ITS, SSU, and tub2 DNA sequence dataset. Bootstrap support values for maximum likelihood (ML) equal to or greater than 70% and Bayesian posterior probabilities (PP) equal to or greater than 0.95 PP are shown above the nodes. The tree is rooted with Amniculicola immersa (CBS 123083) and A. parva (CBS 123092). Newly generated strains are in red, and the type strains are indicated using “T” in superscript.

Taxonomy

Tetraplosphaeriaceae Kaz. Tanaka & K. Hiray, Studies in Mycology 64: 177 (2009)

MycoBank No: 515253

Type genus

Tetraploa Berk. & Broome, Ann. Mag. Nat. Hist. 5: 459, t. 11:6 (1850).

Description

Teleomorph see Tanaka et al. (2009). Anamorph Conidiophores absent. Conidiogenous cells monoblastic. Conidia composed of 3–8 columns or internal hyphal structure, brown, mostly verrucose at the base, with more than 3–8 setose appendages (Tanaka et al. 2009).

Notes

Tetraplosphaeriaceae was described by Tanaka et al. (2009) to accommodate the species which has massarina-like teleomorphic morph and tetraploa-like anamorphs based on a combined SSU and LSU DNA sequence data and established five genera. To date, the members of Tetraplosphaeriaceae are mainly distributed on Poaceae and unidentified decayed wood as saprobes and pathogens from aquatic and terrestrial habitats (Tanaka et al. 2009; Hyde et al. 2013; Hongsanan et al. 2020; Yu et al. 2022; Li et al. 2021). It now contains nine genera and 69 species (Tanaka et al. 2009; Pem et al. 2019; Hongsanan et al. 2020; Li et al. 2021; Liao et al. 2022).

Polyplosphaeria Kaz. Tanaka & K. Hirayama, Studies in Mycology 64: 192 (2009)

MycoBank No: 515256

Type species

Polyplosphaeria fusca Kaz. Tanaka & K. Hirayama, Studies in Mycology 64: 193 (2009).

Description

Teleomorph see Tanaka et al. (2009). Anamorph Conidiophores absent. Conidiogenous cells monoblastic. Conidia globose to subglobose, with thin peel-like outer wall of conidia, composed of numerous internal hyphae at the inside, brown, almost smooth, verrucose at the base. Appendages with 3 to 8 setose appendages, brown, straight (Tanaka et al. 2009).

Notes

Tanaka et al. (2009) established Polyplosphaeria and typified with Po. fusca based on a combined SSU and LSU DNA sequence data. All the members of Polyplosphaeria were reported as saprobes from various plant hosts, such as Pleioblastus chino, Phyllostachys bambusoides and Pandanaceae (Tanaka et al. 2009; Li et al. 2016; Tibpromma et al. 2018). Polyplosphaeria is distributed in Japan, China and Thailand in terrestrial habitats (Tanaka et al. 2009; Li et al. 2016; Tibpromma et al. 2018). Dong et al. (2020) transferred Po. xishuangbannaensis into Ernakulamia based on phylogenetic analyses and differences in morphology. In this study, two new Polyplosphaeria species are introduced from unidentified decaying wood from China. The genus contains six species viz. Polyplosphaeria guizhouensis, Po. hainanensis, Po. fusca, Po. nabanheensis, Po. pandanicola and Po. thailandica (Tanaka et al 2009; Li et al. 2016; Tibpromma et al. 2018; This study; Table 2).

Table 2.

Tetraplosphaeriaceae species and their country, life cycle, habitat, host and reference.

Species name Country Life cycle Habitat Host Reference
Aquatisphaeria thailandica Thailand saprobic freshwater decaying wood Li et al. (2021)
Byssolophis byssiseda France saprobic terrestrial branch of Carpinus, decaying wood Zhang et al. (2012)
B. sphaerioides Finland, UK saprobic terrestrial decaying stemp of Rubus, decaying wood birch Berkeley and Broome (1854); Karsten (1870)
Ernakulamia cochinensis Argentina, Cuba, India, Japan, Malaysia, Mexico, Panama, Thailand saprobic freshwater, terrestrial Astrocaryum standleyanum, Benthamidia japonica, dead leaves, dead spathes of Cocos nucifera, decomposing leaves of Satakentia liukivensis, Freycinetia multi, palm tree, Ilex sp., Ocotea leucoxylon, Pandanus tectorius, P. monticola, submerged wood, Syagrus romanzoffiana, Stewartia monadelpha, Vitex sp. Ellis (1976); Holubová-Jechová and Mercado (1986); Holubová-Jechová (1989); Mercado et al. (1997, 2005); Taylor and Hyde (2003); Delgado and Mena (2004); Capdet and Romero (2010); Whitton et al. (2012); Delgado et al. (2017); Dong et al. (2020); Farr and Rossma (2023)
E. krabiensis Thailand saprobic terrestrial Acacia sp. Jayasiri et al. (2019)
E. tanakae India saprobic terrestrial dead spathes of Cocos nucifera Hyde et al. (2020a)
E. xishuangbannaensis China saprobic terrestrial dead leaves of Pandanus sp. Tibpromma et al. (2018); Dong et al. (2020)
Polyplosphaeria guizhouensis China saprobic terrestrial unidentified decaying wood This study
Po. Hainanensis China saprobic terrestrial unidentified decaying wood This study
Po. Fusca Japan saprobic terrestrial culms of Chimonobambusa marmorea, culms of Phyllostachys bambusoides, culms of Pleioblastus chino, culms of Sasa kurilensis Tanaka et al. (2009)
Po. thailandica Thailand saprobic terrestrial decaying bamboo Li et al. (2016)
Po. nabanheensis China saprobic terrestrial decaying leaves of Pandanus sp. Tibpromma et al. (2018)
Po. pandanicola China saprobic terrestrial decaying leaves of Pandanus sp. Tibpromma et al. (2018)
Pseudotetraploa bambusicola China saprobic terrestrial dead branches of Bamboo Yu et al. (2022)
Ps. curviappendiculata Japan saprobic terrestrial culms of Sasa kurilensis Tanaka et al. (2009)
Ps. yunnanensis China saprobic freshwater bamboo This study
Ps. Javanica Indonesia, Japan saprobic terrestrial culms of decaying Bambusa glaucescens, culms of Phyllostachys bambusoides, culms of Pleioblastus chino, culms of Sasa sp., dead bark of broad-leaved tree, dead stems of an unidentified herbaceous plant Hatakeyama et al. (2005); Tanaka et al. (2009); Rifai et al. (2014)
Ps. longissima Japan saprobic terrestrial culms of Pleioblastus chino Tanaka et al. (2009)
Ps. rajmachiensis India saprobic terrestrial decaying bamboo culms, Dendrocalamus stocksii (Poaceae) Hyde et al. (2020a)
Quadricrura bicornis Japan saprobic terrestrial culms of Sasa kurilensis, leaf litter of a conifer Tanaka et al. (2009)
Q. meridionalis Japan saprobic terrestrial bamboo Tanaka et al. (2009)
Q. septentrionalis Japan saprobic terrestrial culms of Sasa kurilensis Tanaka et al. (2009)
Shrungabeeja aquatica Thailand saprobic freshwater submerged wood Dong et al. (2020)
S. longiappendiculata Thailand saprobic terrestrial dead culm of Bambusa sp. (Poaceae) Ariyawansa et al. (2015)
S. vadirajensis Brazil, China, India saprobic terrestrial dead branches of unidentified plant Rao and Reddy (1981); Zhang et al. (2009)
S. begoniae China saprobic terrestrial dead branches of Begonia semperflorens Zhang et al. (2009)
S. melicopes China saprobic terrestrial dead branches of Melicope triphylla Zhang et al. (2009)
S. piepenbringiana Panama saprobic terrestrial dead Poaceae Kirschner et al. (2017)
S. fluviatilis China saprobic freshwater submerged decaying twig Yang et al. (2023)
Tetraploa abortiva Argentina saprobic freshwater N/A Arambarri et al. (1987)
T. aquatica China saprobic freshwater submerged decaying wood Li et al. (2020)
T. aristata Africa, Barbados, Bolivia, China, Cuba, Denmark, Eire, Europe, Fiji, Germany, Ghana, Hong Kong (China), India, Italy, Japan, Jamaica, Malaysia, Nepal, New Caledonia, Pakistan, Papua New Guinea (New Britain), Philippines, Puerto Rico, Sierra Leone, Thailand, The Dominican Republic, The Netherlands, Uganda, Venezuela, USA(Alabama) pathogenic (human), saprobic terrestrial Alpinia formosa, Ammophila arenaria, Anadelphia leptocoma, Andropogon, Angelica sylvestris, Avena pralensis, Axonopus, Bambusa, Carex paniculata, Cladium mariscus, Cladium selloana, Cocos, Cortaderia, Cymbopogon afronardus, Cyperus longus, Dactylis, Deschampsia, Erianthus, Euchlaena, Festuca, Gynerium argenteum, Gynerium, Heracleum sphondylium, Heteropogon, Hevea brasiliensis, Juncus, Musa, Phalaris arundinacea, Phaseolus, Phoenix, Phormium, Phragmites communis , Poa pratensis, Pteridium aquilinum, Saccharum officinarum, Sorghum, straw, Triticum, unnamed host, wheat stubble, Zea Ellis (1949, 1971); Markham et al. (1990); Tanaka et al. (2009); Senwanna et al. (2021)
T. bambusae China saprobic terrestrial dead twigs of bamboo Phookamsak et al. (2022)
T. biformis Japan saprobic terrestrial dead bark of broad-leaved tree Matsushima and Matsushima (1996)
T. circinata India saprobic terrestrial decaying bamboo twig Pratibha and Bhat (2008)
T. conata F India N/A N/A N/A Saxena and Sarkar (1986); Gupta (2002)
T. cylindrica China saprobic terrestrial decaying stems of Saccharum arundinaceum (Poaceae) Liao et al. (2022)
T. dashaoensis China saprobic terrestrial dead stem of Saccharum arundinaceum Jayawardena et al. (2023)
T. divergens USA (Mississippi) saprobic terrestrial leaves of Panicum agrostidiforme Tracy and Earle (1895)
T. dwibahubeeja India saprobic terrestrial decaying spathes of Cocos nucifera Hyde et al. (2020a)
T. ellisii Argentina, USA (New Jersey), Zimbabwe saprobic terrestrial Chloris, Dactylis, Hevea brasiliensis, stalks of Zea mays Cooke and Ellis (1879); Ellis (1971); Senwanna et al. (2021)
T. endophytica Germany endophytic terrestrial roots of Microthlaspi perfoliatum Crous et al. (2021)
T. hainanensis China Saprobic terrestrial unidentified decaying wood This study
T. indica F India N/A N/A N/A Saxena and Khare (1991)
T. josettae F France N/A N/A N/A Nuñez Otaño et al. (2022)
T. juncicola The Netherlands saprobic terrestrial dead culm of Juncus inflexus (Juncaceae) Crous et al. (2022)
T. muscicola Spain N/A N/A fronds of Aneura multifida, Lophozia quinquedentata González Fragoso (1916)
T. nagasakiensis Japan, China saprobic terrestrial culms of bamboo Hyde et al. (2013, 2019)
T. obpyriformis China saprobic terrestrial dead grass under Saccharum arundinaceum (Gramineae) Unpublished
T. opaca China saprobic terrestrial dead culms of bamboo, decaying branches of unidentified tree Zhao et al. (2009)
T. pseudoaristata India saprobic terrestrial decaying spathes of Cocos nucifera (Arecacceae) Hyde et al. (2020a)
T. puzheheiensis China saprobic freshwater submerged wood Dong et al. (2020)
T. sasicola China, Japan saprobic terrestrial culms of Sasa senanensis, dead leaves of Pennisetum purpureum (Poaceae) Tanaka et al. (2009); Hyde et al. (2020a)
T. scabra USA N/A terrestrial Scirpus sp. Harkness (1885)
T. scheueri UK saprobic freshwater, terrestrial leaves of Carex acutiformis, rotten leaves Scheuer (1991); Hyde et al. (2013)
T. setifera Hungary saprobic terrestrial rotten wood Révay (1993)
T. siwalika F N/A N/A N/A N/A Saxena et al. (1987)
T. taugourdeaui F India N/A N/A N/A Saxena and Sarkar (1986)
T. thailandica Thailand saprobic freshwater Submerged decaying wood Bao et al. (2021)
T. thrayabahubeeja India saprobic terrestrial decaying spathes of Cocos nucifera (Arecacceae) Hyde et al. (2020a)
T. yakushimensis Japan saprobic terrestrial culms of Arundo donax Tanaka et al. (2009); Hyde et al. (2013)
T. yunnanensis China, Thailand saprobic freshwater submerged wood Dong et al. (2020)
Tetraploa sp. 1 Japan saprobic terrestrial culms of bamboo Tanaka et al. (2009)
Tetraploa sp. 2 Japan saprobic terrestrial culms of Gramineae Tanaka et al. (2009)
Triplosphaeria cylindrica Japan saprobic terrestrial culms of Sasa kurilensis Tanaka et al. (2009)
Tr. maxima Japan saprobic terrestrial culms of Sasa kurilensis Tanaka et al. (2009)
Tr. yezoensis Japan saprobic terrestrial culms of Sasa palmata Tanaka et al. (2009)
Tr. acuta Japan saprobic freshwater submerged culms of bamboo Tanaka et al. (2009)
Triplosphaeria sp. Japan saprobic terrestrial culms of Sasa kurilensis Tanaka et al. (2009)

Polyplosphaeria guizhouensis X. Tang, Jayaward., R. Jeewon & J.C. Kang, sp. nov.

MycoBank No: 900950
Fig. 2

Etymology

The specific epithet ‘guizhouensis’ refers to the place where the fungus was collected, Guizhou Province, China.

Holotype

GZAAS 23–0600.

Description

Saprobic on unidentified decaying wood in the forest. Teleomorph not observed. Anamorph Hyphomycetous. Colonies effuse, gregarious on host substrate, brown to dark brown. Mycelium semi-immersed or immersed, pale brown, branched, septate. Conidiophores absent. Conidiogenous cells forming directly on creeping hyphae, integrated, monoblastic,determinate. Conidia 34–61 × 41–63 μm (x̅ = 51 × 51 μm, n = 20), globose to subglobose to turbinate, solitary, olivaceous-green to brown, verrucose and darker at base, with setose appendages on surface. Appendages with two forms, solitary, cylindrical, unbranched, septate, smooth, brown at base and paler towards to apex, long appendages 51–152 × 3–5 μm (x̅ = 89 × 4.0 μm, n = 20), wide at the base, 2–6-septate, arising from apical part of conidia; short appendages 13–38 × 2.5–6 μm (x̅ = 25 × 4 μm, n = 20), wide at the base, 0–3-septate, arising randomly from conidial apex.

Figure 2. 

Polyplosphaeria guizhouensis (GZAAS 23–0600, holotype) a colonies on decaying wood b, c colonies on natural substrates d–n conidia bearing appendages o germinating conidium p colony on PDA (front at right, reverse at left). Scale bars: 50 μm (d–o).

Culture characteristics

Conidia germinated on PDA and incubate at room temperature (25 ˚C). Colonies circular, cottony, flat, slightly grey with an undulate margin, forming three concentric zonation, margin regular, brownish grey. The reverse side is greenish grey in the centre, with a dark brown margin and pigment.

Material examined

China, Guizhou Province, Xingyi City, Xianheping National Forest Park, on unidentified decaying wood, 25 September 2021, Xia Tang, xhp08 (GZAAS 23–0600, holotype), ex-type culture GZCC 23–0598.

Notes

The phylogenetic results (Fig. 1) showed that Polyplosphaeria guizhouensis is sister to Po. pandanicola within Polyplosphaeria with high support (ML = 100, BPP = 1). The comparison of pairwise nucleotides showed that Polyplosphaeria guizhouensis is different from Po. pandanicola in 2/801 bp (0.2%) in LSU and 11/460 (2.5%) in ITS. Thus, we describe Polyplosphaeria guizhouensis herein as a novel species in Polyplosphaeria following recommendations proposed by Jeewon and Hyde (2016) and Chethana et al. (2021).

Polyplosphaeria hainanensis X. Tang, Jayaward., R. Jeewon & J.C. Kang, sp. nov.

MycoBank No: 900951
Figs 3, 4

Etymology

The specific epithet ‘hainanensis’ refers to the place where the fungus was collected, Hainan Province, China.

Holotype

GZAAS 23–0601

Description

Saprobic on unidentified decaying wood in the forest. Teleomorph not observed. Anamorph Hyphomycetous. Colonies effuse, gregarious on host substrate, brown to blackish brown. Mycelium semi-immersed or immersed, dark brown, branched, septate. Conidiophores absent. Conidiogenous cells indistinguishable from creeping hyphae, integrated, monoblastic, determinate. Conidia 49–134.5 × 52–90.5 μm (x̅ = 86 × 71 μm, n = 20), globose, subglobose, obconical, broadly ellipsoidal to broadly pyriform, variable in shape, sometimes with thin peel on the outer wall of conidia, internally filled with a mass of hyaline, solitary, brown to dark brown, smooth. Appendages 36–58 × 3–5.5 μm (x̅ = 44.5 × 4 μm, n = 20), cylindrical, solitary, straight or flexuous, unbranched and almost hyaline at the apex, 0–4-septate, smooth, round at apex, pervasive.

Figure 3. 

Polyplosphaeria hainanensis (GZAAS 23–0601, holotype) a colonies on decay wood b colonies on natural substrates c–m conidia bearing appendages n germinating conidium o colony on PDA (from front) p colony on PDA (from reverse). Scale bars: 50 μm (c–n).

Culture characteristics

Conidia germinated from both ends on PDA and incubated at room temperature (25 ˚C). Colonies circular, cottony, flat, olivaceous with a slightly grey entire margin. The reverse side is an olive drab, which gradually extends outwards to form a deep colour ring in the centre with a pale grey margin and no pigment.

Figure 4. 

Polyplosphaeria hainanensis (GZAAS 23–0602, paratype) a colonies on decay wood b, c colonies on natural substrates d–o conidia bearing appendages p colony on PDA (from front) q colony on PDA (from reverse). Scale bars: 100 μm (d–i, k–m); 50 μm (j, n, o).

Material examined

China, Hainan Province, Wuzhishan City, Wuzhishan National Nature Reserve, on unidentified decaying wood, 25 September 2021, Zili Li, WZS27 (GZAAS 23–0601, holotype), ex-type culture GZCC 23–0599; WZS31 (GZAAS 23–0602, paratype), culture GZCC 23–0600.

Notes

Based on the phylogenetic analysis (Fig. 1), two of our Polyplosphaeria collections share similar morphology and clustered together with high support (ML = 100, and BPP = 1). The base pair differences between the two strains (GZCC 23–0599 and GZCC 23–0600) were: LSU = 0.2% (2/834), ITS = 0.1% (1/840), respectively, therefore, we considered them as the same species according to the guidelines for species delineation proposed by Jeewon and Hyde (2016). The phylogenetic result (Fig. 1) showed that Polyplosphaeria hainanensis is sister to Po. nabanheensis within Polyplosphaeria. Based on the comparison of the morphological characters with other species in Polyplosphaeria, our collection can be distinct in having obconical, broadly ellipsoidal to broadly pyriform, variable conidial shape (without verrucose at the base) and pervasive appendages. The comparison of pairwise nucleotides showed that Polyplosphaeria hainanensis is different from Po. nabanheensis in 24/826 bp (3%) in LSU, 20/758 (2.6%) in SSU, 17/472 (3.6%) in ITS and 16/344 (5%) in tub2. Thus, we describe Polyplosphaeria hainanensis herein as a novel species in Polyplosphaeria according to the guidelines of Jeewon and Hyde (2016) and Chethana et al. (2021).

Pseudotetraploa Kaz. Tanaka & K. Hirayama, Studies in Mycology 64: 193 (2009)

MycoBank No: 515257

Type species

Pseudotetraploa curviappendiculata (Sat. Hatak., Kaz. Tanaka & Y. Harada) Kaz. Tanaka & K. Hirayama, Studies in Mycology 64: 195 (2009).

Description

Teleomorph morph not observed. Anamorph Mycelium superficial. Conidiophores absent. Conidiogenous cells monoblastic, indistinguishable from creeping hyphae. Conidia composed of 4 to 8 columns, obpyriform to long obpyriform, brown to dark brown, almost smooth, verrucose at the base, pseudoseptate, with setose appendages at the apical part. Appendages mostly 4, rarely 6 to 8, curved or straight (Tanaka et al. 2009).

Notes

Tanaka et al. (2009) established Pseudotetraploa (Ps.) with three species, which were previously described in Tetraploa and typified by Ps. curviappendiculata based on a combined SSU and LSU DNA sequence data. Pseudotetraploa species are reported as saprobes on bamboo, dead bark of the broad-leaved tree, and unidentified herbaceous plants in Japan, China, and India (Hatakeyama et al. 2005; Tanaka et al. 2009; Hyde et al. 2020a; Yu et al. 2022). Pseudotetraploa is only known in its anamorphic state and dwells in terrestrial habitats and contains six species viz. Ps. bambusicola, Ps. curviappendiculata, Ps. javanica, Ps. longissima, Ps. Rajmachiensis, and Ps. yunnanensis (Hatakeyama et al. 2005; Tanaka et al. 2009; Hyde et al. 2020a; Yu et al. 2022; This study; Table 2). In this study, a new Pseudotetraploa species isolated from bamboo is introduced.

Pseudotetraploa yunnanensis X. Tang, Jayaward., R. Jeewon & J.C. Kang, sp. nov.

MycoBank No: 900963
Fig. 5

Etymology

The specific epithet ‘yunnanensis’ refers to the place where the fungus was collected, Yunnan Province, China.

Holotype

HKAS 129442.

Description

Saprobic on bamboo. Teleomorph not observed. Anamorph Hyphomycetous. Colonies effuse, gregarious on host substrate, brown to dark brown. Mycelium superficial, hyaline to pale brown. Conidiophores absent. Conidiogenous cells micronematous, mononematous, monoblastic, integrated, usually undistinguishable from superficial hyphae. Conidia 67–120 × 16.5–35 μm (x̅ = 95 × 24 μm, n = 20), solitary, septate, brown to dark brown, ovoid to obclavate or narrowly obpyriform, consisting of 3–6 columns of cells, rounded at the base 19–36 μm wide (x̅ = 26 μm, n = 20), slightly constricted at septa, rarely branched and make V-shaped conidia; setose appendages at the apical part 15–87 × 3.5–7 μm (x̅ = 37 × 5 μm, n = 20), appendages 3–6 in number, 1–8-septate, brown at the base and almost hyaline at the apex, smooth, unbranched, shorter appendage is straight and longer appendage is curved.

Figure 5. 

Pseudotetraploa yunnanensis (HKAS 129442, holotype) a, b colonies on natural substrates c–n conidia. Scale bars: 20 μm (c); 50 μm (d–n).

Culture characteristics

Conidia germinated from both ends on PDA and incubated at room temperature (25 ˚C). Colonies circular, cottony, flat, slightly grey with an entire margin, containing a circular white mycelium in the centre. The reverse side is a pale brown in the centre that gradually extends outwards while the colour changes to pale grey, with a brown margin and no pigment.

Material examined

China, Yunnan Province, Puer City, Ailao mountains, on bamboo, May 23, 2022, Rong-Ju Xu, ALS 29 (HKAS 129442, holotype), ex-type culture KUNCC 10464.

Notes

Pseudotetraploa yunnanensis is similar to Ps. curviappendiculata and Ps. longissima. However, Pseudotetraploa yunnanensis differs from Ps. curviappendiculata in having branched and V-shaped conidia, consisting of 3–6 columns of cells with 3–6 apical appendages, larger conidia [67–120 μm vs. 52–67(–75) μm] in length and [16–35 μm vs. 15–22 μm] in width, while Ps. curviappendiculata consists of 4–5 columns of cells with 4 apical appendages; Pseudotetraploa yunnanensis differs from Ps. longissima in having smaller conidia [67–120 μm vs. (98–)110–148(–155) μm] in length and [16–35 μm vs. 18–25 μm] in width, without verrucose at the base. The phylogenetic analysis showed that Pseudotetraploa yunnanensis is sister to Ps. rajmachiensis and Ps. javanica. The comparison of pairwise nucleotides showed that Pseudotetraploa yunnanensis is different from Ps. rajmachiensis in 27/1021 bp (2.6%) in LSU and 30/560 (6%) in ITS; Pseudotetraploa yunnanensis is different from Ps. javanica in 11/1020 bp (1.1%) in LSU and 17/538 (3.2%) in ITS. Thus, we describe Pseudotetraploa yunnanensis herein as a novel species in Pseudotetraploa according to the guidelines Jeewon and Hyde (2016) and Chethana et al. (2021).

Tetraploa Berk. & Broome, Ann. Mag. Nat. Hist. 5: 459, t. 11:6 (1850)

MycoBank No: 10199

= Tetraplosphaeria Kaz. Tanaka & K. Hiray., in Tanaka et al., Stud. Mycol. 64: 177 (2009).

Type species

Tetraploa aristata Berk. & Broome, Ann. Mag. Nat. Hist. 5: 459 (1850).

Description

Teleomorph see Tanaka et al. (2009). Anamorph Tetraploa sensu stricto Conidiophores absent. Conidiogenous cells monoblastic. Conidia composed of 4 columns, short-cylindrical, brown, verrucose at the base, euseptate, with 4 setose appendages at the apex (Tanaka et al. 2009).

Notes

Tanaka et al. (2009) established Tetraplosphaeria to accommodate pleosporalean species that have massarina/lophiostoma-like teleomorph and anamorphs belonging to Tetraploa sensu stricto based on a combined SSU and LSU DNA sequence data. Later, Hyde et al. (2013) treated Tetraploa as a synonym of Tetraplosphaeria, which has been applied previously to anamorphic species and used Tetraploa instead of Tetraplosphaeria. Species of Tetraploa are mainly reported as saprobes, distributed in freshwater and terrestrial habitats, and only T. aristata has been reported as a pathogen on various plants and human pathogen that cause cysts (Markham et al. 1990; Tanaka et al. 2009; Hyde et al. 2013; Liao et al. 2022). Tetraploa has been recovered from more than 80 plants, such as bamboo culms, submerged wood, palms, and Poaceae, on the leaves of Acer and liverworts (Ellis 1949; Ando 1992; Hyde et al. 2013; Liao et al. 2022). Saxena et al. (2021) mentioned that Frasnacritetrus is probably a fossil of Tetraploa. Nuñez Otaño et al. (2022) considered Frasnacritetrus as a synonym of Tetraploa and transferred five Frasnacritetrus fossil species into Tetraploa viz. Tetraploa conata, T. indica, T. josettae, T. siwalika and T. taugourdeaui based on the observation that the spores of both fossil and contemporary species exhibit identical morphological characteristics. To date, there are 35 species accepted in Tetraploa (Wijayawardene et al. 2022; Jayawardena et al. 2023; this study; Table 2). In this study, a new Tetraploa species is introduced.

Tetraploa hainanensis X. Tang, Jayaward., R. Jeewon & J.C. Kang, sp. nov.

MycoBank No: 900952
Figs 6, 7

Etymology

The specific epithet ‘hainanensis’ refers to the place where the fungus was collected, Hainan Province, China.

Holotype

GZAAS 23–0603.

Description

Saprobic on unidentified decaying wood in forest. Teleomorph morph Not observed. Anamorph Hyphomycetous. Colonies effuse, gregarious on host substrate, brown to dark brown. Mycelium semi-immersed or immersed, pale brown, branched, septate. Conidiophores absent. Conidiogenous cells integrated, monoblastic, determinate. Conidia 30–46 × 18–36 μm (x̅ = 38 × 27 μm, n = 20), cylindrical with obtuse ends, pale brown to brown, verrucose, composed of four columns of cells, sometimes five columns of cells, 4–5-septate in each column, smooth, mostly with four apical appendages, some with one or two or five appendages. Appendages 52–209 × 3–6 μm (x̅ = 140 × 4 μm, n = 20) cylindrical, solitary, unbranched, guttulate, septate, wide at the base, divergent, pale brown to brown, 5–16-septate, straight or slightly flexuous, smooth-walled.

Figure 6. 

Tetraploa hainanensis (GZAAS 23–0603, holotype) a, b colonies on natural substrates c–n conidia bearing 1–5 appendages o germinating conidium p colony on PDA (from reverse) q colony on PDA (from front). Scale bars 20 μm (c); 50 μm (d–g); 100 μm (h–o).

Culture characteristics

Conidia germinated from both ends on PDA and incubated at room temperature (25 ˚C). Colonies circular, cottony, flat, slightly grey with an entire margin, contain a circular white mycelium in the centre. The reverse side is a pale brown in the centre that gradually extends outwards while the colour changes to pale grey, with a brown margin and no pigment.

Figure 7. 

Tetraploa hainanensis (GZAAS 23–0604, paratype) a colonies on decay wood b, c colonies on natural substrates d–p conidia bearing 1–4 appendages q germinating conidium r colony on PDA (from front) s colony on PDA (from reverse). Scale bars: 20 μm (d–g); 100 μm (h–l, o, q); 50 μm (m, n, p).

Material examined

China, Hainan Province, Wuzhishan City, Wuzhishan National Nature Reserve, on unidentified decaying wood, 25 September 2021, Zili Li, WZS59 (GZAAS 23–0603, holotype), ex-type culture GZCC 23–0601; WZS66.2 (GZAAS 23–0604, paratype), culture GZCC 23–0602.

Notes

Tetraploa hainanensis is morphologically similar to T. pseudoaristata. However, Tetraploa hainanensis can be distinguished from T. pseudoaristata in having larger conidia (30.5–46 × 18–36 μm vs. 22–31 × 15–20 µm) with four columns of cells, sometimes five columns of cells, and longer appendages (52–209 × 3–6 μm vs. 23–107 × 2–5 μm), commonly four in number, sometimes five. Based on the phylogenetic analysis, two of our Tetraploa collections which share similar morphology clustered together with high support (ML = 100, and BPP = 1 (Fig. 1)). The base pair differences between the two strains were: LSU = 0.1% (1/806), ITS = 0% (0/516), and tub2 = 0% (1/633), respectively. Therefore, we considered them as the same species according to the guidelines for species delineation proposed by Jeewon and Hyde (2016). Tetraploa hainanensis forms a distinct lineage but close to T. yakushimensis and T. tetraploa. However, Tetraploa hainanensis differs from T. yakushimensis by having four or five columns and appendages, while T. yakushimensis has only four columns and appendages; Tetraploa hainanensis differs from T. tetraploa in having four or five columns and shorter appendages (52–209 × 3–6 μm vs. 263–350 × 2–3 μm), while T. tetraploa has only four columns and slender appendages. The comparison of pairwise nucleotide showed that Tetraploa hainanensis is different from T. yakushimensis in 31/620 bp (3%) in LSU, 7/814 (0.98%) in ITS, and 87/450 (19%) in tub2 and Tetraploa hainanensis is different from T. tetraploa in 31/620 bp (3%) in LSU, 7/814 (0.98%) in ITS, and 87/450 (19%) in tub2. Based on the combination of morphological characters and multigene phylogeny, we describe Tetraploa hainanensis herein as a distinct species according to the guidelines of Jeewon and Hyde (2016) and Chethana et al. (2021).

Discussion

Hyde et al. (2018) reported that more than 95% of fungi collected in northern Thailand could be new to science and there is a dire need to collect more samples from a wide variety of hosts to better understand fungal diversity estimates. In the same way, fungal diversity in Yunnan, Guizhou and Hainan is expected to be rather high. In this study, collections of decayed wood samples and bamboo were done to assess which fungal species are potentially colonising them. Our study reveals four anamorphic fungal species that belong to the family Tetraplosphaeriaceae. In this study, we characterise two new anamorphic species collected from unidentified decayed wood samples that belong to Polyplosphaeria. In Polyplosphaeria this study brings the number of species to six. The first new species is described as Po. guizhouensis and our multigene phylogeny depict a close relationship to Po. pandanicola. The latter was collected from fallen dead and decaying leaves of Pandanus sp. in China (Tibpromma et al. 2018) and characterized by micronematous conidiophores; monoblastic, incomplete globose connected to base of conidia conidiogenous cell with guttules, hyaline; globose to subglobose, solitary, verrucose at base conidia with almost hyaline at apex, unbranched setose appendages on surface. However, Po. guizhouensis differs in having turbinate conidia which are verrucose and darker at the base. Furthermore, it possesses a longer conidium base and two types of appendages originating from the apical part of the conidia. With regard to DNA sequence data comparison, Po. guizhouensis differs from Po. pandanicola (MFLUCC 17–2266) in having 11 out of 460 (2.5%) and 2 out of 801 (0.2%) different base pairs (bp) in the ITS alignments and LSU gene respectively. Our second new species, named Polyplosphaeria hainanensis, forms a strongly supported subclade with Po. nabanheensis. The latter was collected from fallen dead and decaying leaves of Pandanus sp. in China (Tibpromma et al. 2018). It is characterised by monoblastic, hyaline conidiogenous cells with guttules; oval to ellipsoid conidia, made up of 2–3 cells, and verrucose at base, rough-walled, with apical setose appendages. However, Po. hainanensis differs with regards to having an obconical, broadly ellipsoidal to broadly pyriform, variable shaped conidia (no verrucose at the base) and pervasive appendages. Comparison of available LSU, ITS, SSU and tub2 sequences also reveal differences in base pairs that support species distinctiveness. For instance, Po. hainanensis differs from Po. nabanheensis in having 24/826 bp (3% difference) in LSU, 20/758 (2.6% difference) in SSU, 17/472 (3.6% difference) in ITS and 16/344 (5% difference) in tub2. Another peculiar finding when we analysed the relationships of Polyplosphaeria species, we found that two strains of Polyplosphaeria thailandica (MFLU 15–3273) and Aquatisphaeria thailandica (MFLUCC 21–0025 and DLUCC B151) clustered together with 75% ML/0.99 BPP support and sister to species of Shrungabeeja. However, it’s important to note that this relationship lacked significant statistical support, a pattern observed in various previous studies as well (Li et al. 2021; Liao et al. 2022). Aquatisphaeria thailandica has been reported as a saprobe on submerged decaying wood in China. It is characterised by macronematous, mononematous, solitary, unbranched conidiophores with 3–4 septa; monoblastic, integrated, terminal, subcylindrical conidiogenous cells; and acrogenous, solitary, subglobose or turbinate, muriform, dictyoseptate conidia with 3–4 (mostly 4) cylindrical, upward appendages with 1–2-septa. At the same time, Po. thailandica is recognised as a saprobe found in decaying bamboo in Thailand. It is characterised by monoblastic conidiogenous cells; acrogenous, solitary, globose, obovoid, pyriform, ellipsoidal, obconical, muriform, verrucose conidia with 2–5-septate appendages, occasionally, two conidia are associated together at the basal cell (Li et al. 2016; Li et al. 2020). Based on the phylogenetic analyses, it seems that Po. thailandica is a member of Aquatisphaeria. There is not much taxonomic data available for Aquatisphaeria, hence we recommend that further collections of this genus are required to elucidate its relationships to Po. thailandica. Alternatively, there might be a need to relook into the taxonomy of Po. thailandica and verify whether the DNA sequences submitted are reliable.

The third anamorphic species was collected from bamboo in Yunnan, China and subsequently assigned to Pseudotetraploa. To date, five species have been reported, and this study extends the known species count to six. The new species is described as Ps. yunnanensis and our multigene phylogeny depict a close relationship to Ps. rajmachiensis. The latter was collected from decaying bamboo culms in India (Hyde et al. 2020a) and characterised by the absence of conidiophores; micronematous, mononematous, monoblastic conidiogenous cells; ovoid to obclavate or obpyriform conidia, minutely verrucose at the base; unbranched, septate, setose appendages at the apical part, consisting of two appendages with one straight and one curved. However, Ps. yunnanensis differs in having larger conidia that rarely separate, consisting of 3–6 columns of cells, forming a V-shape conidia, 3–6 apical appendages. In terms of DNA sequence data comparison, Ps. yunnanensis differs from Ps. rajmachiensis (NFCCI 4618) in LSU by 27/1021 bp (2.6% difference) and in ITS by 30/560 bp (6% difference).

The last anamorphic species was collected from unidentified decaying wood in Hainan, China, and was assigned to Tetraploa. With the addition of this species, the genus now comprises a total of 35 species. The new species is described as T. hainanensis, and the multigene phylogeny depicts a close relationship to T. yakushimensis. The latter was collected on culms of Arundo donax in Japan (Tanaka et al. 2009) and characterised by the absence of conidiophores; monoblastic conidiogenous cells; solitary, short cylindrical, brown, verruculose conidia, composed of 4 columns with 4 apical setose appendages. However, T. hainanensis differs in having four or five culms and appendages. In terms of DNA sequence data comparison, T. hainanensis differs from T. yakushimensis (KT 1906) in 31/620 bp (3% difference) in LSU, 7/814 (0.98% difference) in ITS, and 87/450 (19% difference) in tub2.

Tetraplosphaeriaceae is a well-known family in terrestrial habitats with mostly saprobes being reported so far, and previous and recent studies have shown that Tetraplosphaeriaceae is widely associated with many plants in different countries. In this work, we describe four new Tetraplosphaeriaceae species based on phylogenetic and morphological comparisons with allied taxa, update the phylogeny of the Tetraplosphaeriaceae family and also provide a checklist of species with other details (Table 2). To date, there are 69 species in Tetraplosphaeriaceae, of which 23 species (including this study) are from China. This study enriches the diversity of fungi in China of Tetraplosphaeriaceae species.

Acknowledgements

The authors are grateful to Shaun Pennycook for his suggestions on naming the new fungus. In addition, the authors also would like to thank Mae Fah Luang University for its support in the tuition fee scholarship. This work was supported by the Distinguished Scientist Fellowship Program (DSFP) at King Saud University, Riyadh, Saudi Arabia.

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 grants from the National Natural Science Foundation of China (NSFC Grants Nos. 32170019 & 31460011) and the Open Fund Program of Engineering Research Centre of Southwest Bio-Pharmaceutical Resources, Ministry of Education, Guizhou University No. GZUKEY20160702. The authors are grateful to the Thailand Research Fund grant “Impact of climate change on fungal diversity and biogeography in the Greater Mekong Sub-region” (RDG6130001).

Author contributions

Xia Tang conducted the experiments, analysed the data, and wrote the first draft of the manuscript. Rajesh Jeewon, Yong-Zhong Lu, Ruvishika S. Jayawardena and Ji-Chuan Kang planned the experiments. Xia Tang, Jian Ma and Rong-Ju Xu analysed the data. Xia Tang and Xue-Mei Chen conducted the experiments. Rajesh Jeewon, Yong-Zhong Lu, Ruvishika S. Jayawardena, Abdulwahed Fahad Alrefaei and Ji-Chuan Kang corrected and revised the manuscript. Yong-Zhong Lu and Ji-Chuan Kang funded the experiments. All authors revised and agreed to the published version of the manuscript.

Author ORCIDs

Xia Tang https://orcid.org/0000-0003-2705-604X

Rajesh Jeewon https://orcid.org/0000-0002-8563-957X

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

Abdulwahed Fahad Alrefaei https://orcid.org/0000-0002-3761-6656

Ruvishika S. Jayawardena https://orcid.org/0000-0001-7702-4885

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

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

Xue-Mei Chen https://orcid.org/0009-0004-8631-0735

Ji-Chuan Kang https://orcid.org/0000-0002-6294-5793

Data availability

All of the data that support the findings of this study are available in the main text. DNA sequences generated have been submitted to Genbank.

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