<8) MycoKeys MycoKeys 100: 171-204 (2023) DOI: 10.3897/mycokeys.100.113141 Research Article Morphophylogenetic evidence reveals four new fungal species within Tetraplosphaeriaceae (Pleosporales, Ascomycota) from tropical and subtropical forest in China Xia Tang'?3®, Rajesh Jeewon**®©, Yong-Zhong Lu'®®, Abdulwahed Fahad Alrefaei™®, Ruvishika S. Jayawardena2*©, Rong-Ju Xu2*®, Jian Ma?2®, Xue-Mei Chen”®, Ji-Chuan Kang'® 1 Engineering and Research Center for Southwest Biopharmaceutical Resource of National Education Ministry of China, Guizhou University, Guiyang, 550025, Guizhou Province, China ND oo Fe W PD Qujing 6550171, China Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai, 57100, Thailand School of Science, Mae Fah Luang University, Chiang Rai, 57100, Thailand Department of Health Sciences, Faculty of Medicine and Health Sciences, University of Mauritius, Reduit, Mauritius Department of Zoology, College of Science, King Saud University, PO. Box 2455, Riyadh 11451, Saudi Arabia School of Food and Pharmaceutical Engineering, Guizhou Institute of Technology, Guiyang, Guizhou Province550003, China Center for Yunnan Plateau Biological Resources Protection and Utilization, College of Biological Resource and Food Engineering, Qujing Normal University, Corresponding authors: Rajesh Jeewon (r.jeewon@uom.ac.mu); Ji-Chuan Kang (jckang@gzu.edu.cn) OPEN Qrceess Academic editor: C. S. Bhunjun Received: 22 September 2023 Accepted: 9 November 2023 Published: 6 December 2023 Citation: Tang X, Jeewon R, Lu Y-Z, Alrefaei AF, Jayawardena RS, Xu R-J, Ma J, Chen X-M, Kang J-C (2023) Morphophylogenetic evidence reveals four new fungal species within Tetraplosphaeriaceae (Pleosporales, Ascomycota) from tropical and subtropical forest in China. Mycokeys 100: 171-204. https://doi. org/10.3897/mycokeys.100.113141 Copyright: © Xia Tang et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract Tetraplosphaeriaceae (Pleosporales, Ascomycota) is a family with many saprobes re- corded from various hosts, especially bamboo and grasses. During a taxonomic inves- tigation 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 com- bined dataset of LSU, ITS, SSU, and tub2 DNA sequence data. Detailed illustrations, de- scriptions 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 cli- mate 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 di- versity. 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; 171 Xia Tang et al.: Novel fungi from China Bao et al. 2021; Mortimer et al. 2021). So far more than 6000 fungal species de- scribed alone from Yunnan province (Feng and Yang 2018). Guizhou, as a prom- inent 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 spe- cial climate that fosters the growth of numerous rare, endangered, and indige- nous plant, animal, and fungal species. Over the past few decades, extensive research has focused on fungi, encompassing both macro and microfungi, lead- ing 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 in- vestigate microfungi in this region, and potentially discover new fungal species. Tetraplosphaeriaceae was introduced by Tanaka et al. (2009) to accommo- date the massarina-like species that produced tetraploa-like anamorphs in cul- ture 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). Tetraplos- phaeriaceae 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 Tetraplosphaeriace- ae based on morphological and molecular evidence from S. longiappendiculata. Later, Ernakulamia was accommodated in Tetraplosphaeriaceae based on mor- phology and phylogenetic analyses of combined ITS, LSU and tub2 sequence data (Delgado et al. 2017; Hyde et al. 2020a). Pem et al. (2019) transferred Bys- solophis from the genera incertae sedis to Tetraplosphaeriaceae based on its massarina-like morphology and phylogenetic analyses based on combined LSU, SSU, tef7-a, and rpbb2 sequence data. Hongsanan et al. (2020) provided a taxo- nomic update on families of Dothideomycetes and eight genera were accepted in Tetraplosphaeriaceae. Recently, Li et al. (2021) discovered a freshwater fun- gus 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 (Wi- jayawardene et al. 2022). However, Pseudotetraploa, Tetraploa, and Triplos- phaeria 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; Mycokeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 172 Xia Tang et al.: Novel fungi from China Jayawardena et al. 2023). Tetraplosphaeriaceae is characterized by massari- na-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 as- cospores, 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 (Tana- ka et al. 2009; Hyde et al. 2013; Hongsanan et al. 2020). In this study, the aim is to characterize anamorphic fungal species collect- ed 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 morpholog- ical differences and similarities with extant species; 3) to establish four new species within the family Tetraplosphaeriaceae with support from results gen- erated 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 col- lected in Guizhou (Xingyi city, Xianheping National Forest Park), Hainan (Wuzhis- han city, Wuzhishan National Nature Reserve), and Yunnan (Puer city, Ailao moun- tains) 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; Yun- nan-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 her- baria of Kunming Institute of Botany, Chinese Academy of Sciences (HKAS), Kun- ming, 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 MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 173 Xia Tang et al.: Novel fungi from China 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), LROR/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, sequenc- ing, and phylogenetic analyses, see Tang et al. (2022). The polymerase chain reaction was carried out in a volume of 50 uL, and the reagents that were used were as follows: DNA template (2 uL), forward primers (2 uL), reverse primers (2 pL), 2 xTaq PCR Master Mix (25 uL) and 19 uL of ddH,0 (double-distilled wa- ter). 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 as- sembled 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. 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; Sta- matakis 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 1000" generation. The Posterior Probabilities (PP) for the remaining MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 174 Xia Tang et al.: Novel fungi from China Table 1. Taxa used in this study and their GenBank accession numbers for LSU, ITS, SSU and tub2 sequence data. GenBank Accession Numbers a eas 08st —Ne.ose868 | NGosaree Aq. Thailandica DLUCC B151 | Mws90764 «=| «=Mwe90968 Taxa name Strain Numbers Amniculicola immersa E. krabiensis MELUCC 18-0237T MK347990 MK347880 MK347773 E. tanakae NFCCI 4615T MN937211 Po MN937229 ezec 2a-059eT oRaseee on a2T27 ~ezcc.23-0s00 —oRsa6890 |= =| =COR ATTN Twrwuce1s-oeeoT | Kuzee7e7 | —~«|~——KU2AB766 CGMCC 3.20939T ON332933 ON332923 ON332915 nrcerasier | noa7200 || N72 RuncetoaseT ——oRase891 OR MMBO79 E. xishuangbannaensis Polyplosphaeria fusca Po. Fusca Po. guizhouensis Po. Hainanensis Po. Hainanensis Po. Thailandica Po. nabanheensis Po. pandanicola Pseudotetraploa bambusicola Ps. bambusicola Ps. curviappendiculata Ps. Javanica Ps. Longissima Ps. rajmachiensis Ps. Yunnanensis | Quadricrura bicornis Q. meridionalis TwFLuce Te-os6er | wTez76sa | | MToarz20 pec 7etes | kraveera | | _xTaTGA7S Q. septentrionalis Shrungabeeja aqutica S. longiappendiculata S. longiappendiculata S. fluviatilis GZCC 19-0511 MW133853 MW134631 Lo = I MFLUCC 17-2362 MN913685 Pe MT627681 MELU 19-0996 MT530453 MT530454 MT530449 S. vadirajensis Tetraploa aquatica T. aquatica MFLUCC 19-0995T MT530452 | = =- | MT530448 T. aristata CBS 996.70 AB524627 AB524486 AB524805 T. bambusae KUMCC 21-0844T ONO77067 ON077073 ONO77078 ~nFociasarT | MNse7207 |) N25 ~nroorasca | noava0e |= =| ~CND7925 eestariiat | mwosoies || zT0079 ~ezcc2a-0so2 —orsae899 | =| _oaz7a26 cas aso4s | ONeose00 | ON G7E0 MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 T. dwibahubeeja T. dwibahubeeja T. endophytica T. hainanensis T. hainanensis T. juncicola T. nagasakiensis T. nagasakiensis CBS 125427T AB524613 AB524472 AB524797 tub2 MN938312 AB524853 AB524851 OR449118 OR449115 MH412745 AB524854 AB524857 AB524858 AB524859 AB524860 AB524861 AB524867 ONO75065 MN938308 MN938309 OR449116 OR449117 AB524868 175 Xia Tang et al.: Novel fungi from China Taxa name T. pseudoaristata T. pseudoaristata T. puzheheiensis T. sasicola T. thailandica T. thrayabahubeeja T. thrayabahubeeja T. yunnanensis T. yakushimensis T. cylindrica T. cylindrica T. dashaoensis T. obpyriformis Tetraploa sp.1 Tetraploa sp.2 T. tetraploa Triplosphaeria acuta Tr. Cylindrica Tr. cylindrica Tr. maxima Tr. yezoensis Tr. yezoensis Triplosphaeria sp. Triplosphaeria sp. Strain Numbers NFCCI 4624T NFCCI 4625 MFLUCC 20-0151T KT 563T MFLUCC 21-0030T NFCCI 4627T NFCCI 4628 MFLUCC 19-0319T KT 1906T KUMCC 20-0205T ZHKUCC 22-0087 KUMCC 21-0010T KUMCC 21-0011T KT 1684 KT 2578 CY112 KT 1170T NBRC 106247 KT 1800 KT 870T KT 1715T KT 1732 HHUF 27481 KT 2546 LSU MN937214 MN937212 MT627655 AB524631 MZ412530 MN937217 MN937215 MN913735 AB524632 MT893204 ON555688 OL473555 OL473554 AB524628 AB524629 AB524633 AB524636 AB524635 AB524637 AB524638 AB524639 AB524640 AB524641 GenBank Accession Numbers SSU AB524490 MZ413274 MT864341 AB524491 MT893203 ON555690 OL473556 OL473557 AB524487 AB524488 AB524492 AB524495 AB524494 AB524496 AB524497 AB524498 AB524499 AB524500 ITS MN937232 MN937230 MT627744 AB524807 MZ412518 MN937235 MN937233 MT627743 AB524808 MT893205 ON555689 OL473549 0L473558 HQ607964 AB524809 AB524811 AB524810 AB524812 AB524813 AB524814 AB524815 AB524816 tub2 MN938315 MN938313 AB524869 MN938318 MN938316 AB524870 MT899417 ON564477 OL505601 OL505600 AB524871 AB524873 AB524872 AB524874 AB524875 AB524876 AB524877 AB524878 Notes: Ex-type strains are indicated by “T” at the end of the strain number, and newly generated sequences are in bold. Abbreviations: BCC: Biotec Culture Collection, Bangkok, Thailand; CBS: Westerdijk Fungal Biodiversity Institute, Utrecht, the Netherlands; CGMCC: China General Microbiological Culture Collection Centre, Beijing, China; DLUCC: Dali University Culture Collection, Yunnan, China; GZCC: Guizhou Culture Collection, Guizhou, China; HHUF: Herbaria of Hirosaki University; IFRDCC: Culture Collection, International Fungal Re- search and Development Centre, Chinese Academy of Forestry, Kunming, China; JCM: the Japan Collection of Microorganisms, Japan; KUNCC: Kunming Institute of Botany Culture Collection; KT: Kazuaki Tanaka; MFLUCC: Mae Fah Luang University Culture Collection, Chiang Rai, Thailand; MFLU: Mae Fah Luang University Herbarium Collection; NBRC: Nite Biological Resource Center, Department of Bio- technology, National Institute of Technology and Evaluation, Japan; NFCCI: National Fungal Culture Collection of India NFCCI-A National Facility; UESTCC: University of Electronic Science and Technology Culture Collection, Chengdu, China; ZHKUCC: Zhongkai University of Agriculture and Engineering Culture Collection, Guangzhou, China. A. = Amniculicola. Aq. = Aquatisphaeria. E. = Ernakulamia. Po. = Polyplosphaeria. Ps. = Pseudotetraploa. Q. = Quadricrura. S. = Shrungabeeja. T. = Tetraploa. Tr. = Triplosphaeria. 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 cri- teria. 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 re- covered from the combined dataset of ML and PP analyses. The top-scoring MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 176 Xia Tang et al.: Novel fungi from China 7510.99 73/0.99 86/0.99 a Pseudotetraploa javanica JCM 12854 Pseudotetraploa curviappendiculata JCM 12852" Pseudotetraploa yunnanensis KUNCC 10464! Pseudotetraploa rajmachiensis NFCCI 46187 Pseudotetraploa bambusicola UESTCC 22-0005 Pseudotetraploa bambusicola CGMCC 3.20939! eudotetraploa longissima JCM 12853" Pseudotetrapl 77/0.99 81/0.99 100/1 Triplosphae Triplosphaeria | 100/1p 2riplosphaeria Triplosphaeria sp. ae 92/1 Shrungabeeja fluviatilis GZCC19-0511 97/1 78/9.99 Shrungabeeja fluviatilis GZCC20—-0505" bh} Shrungabeeja longiappendiculata BCC 76464 "7 Shrungabeeja longiappendiculata BCC 76463" Shrung abeej 3 Shrungabeeja aqutica MFLUCC 18-0664" Shrungabeeja vadirajensis MFLUCC 17-2362 100/1_pAquatisphe A quatisphae ailc Polyplosphaeria thailandica MFLUCC 15-0840! -/0.95,--Ernakulamia cochinensis MFLUCC 18-1237 Jeu 0.99 Ernakulamia xishuangbannaensis KUMCC 17-0187" Ernakulamia tanakae NFCCI 4615" , Ernakulamia krabiensis MFLUCC 18-0237" Ernakulamia 100/1 1aeria guizhouensis G : | 96/1 75/909 100/1_,Pol Poly DI Lod/1p Po! Polyp 82/1 90/1 00/1 99/] Quadricrura meridionalis CBS 125684" ; Quadricrura bicornis CBS 125427" Quadricrura Quadricrura septentrionalis CBS 125429 Byssolophis sphaerioides IFRDCC2053 100/1 Amniculicola parva CBS 123092? Amniculicola immersa CBS 123083" Outgroups 100/1 0.03 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. MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 177 Xia Tang et al.: Novel fungi from China RAXxML tree is shown in Fig. 1, with a final ML optimization likelihood value of -17569.286960 (In). 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 mod- els 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 Tetraplosphae- riaceae clade based on the results of the combined LSU, ITS, SSU, and tub2 sequence data analysis. Taxonomy Tetraplosphaeriaceae Kaz. Tanaka & K. Hiray, Studies in Mycology 64: 177 (2009) MycoBank No: 515253 Facesoffungi Number: FoF06665 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 ac- commodate the species which has massarina-like teleomorphic morph and tet- raploa-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 Facesoffungi Number: FoF06668 Type species. Polyplosphaeria fusca Kaz. Tanaka & K. Hirayama, Studies in My- cology 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). MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 178 Xia Tang et al.: Novel fungi from China 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 Pleio- blastus 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 Ernakula- mia 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). Polyplosphaeria guizhouensis X. Tang, Jayaward., R. Jeewon & J.C. Kang, sp. nov. MycoBank No: 900950 Facesoffungi Number: FoF 14571 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 di- rectly on creeping hyphae, integrated, monoblastic,determinate. Conidia 34-61 x 41-63 um (x= 51 x 51 um, n = 20), globose to subglobose to turbinate, solitary, ol- ivaceous-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 x 3-5 um (x = 89 x 4.0 um, n = 20), wide at the base, 2—6-septate, arising from api- cal part of conidia; short appendages 13-38 x 2.5-6 um (x = 25 x 4 um, n = 20), wide at the base, 0-3-septate, arising randomly from conidial apex. 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 re- verse side is greenish grey in the centre, with a dark brown margin and pigment. Material examined. CHINA, Guizhou Province, Xingyi City, Xianheping Nation- al 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 sup- port (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 recommen- dations proposed by Jeewon and Hyde (2016) and Chethana et al. (2021). MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 179 Xia Tang et al.: Novel fungi from China Figure 2. Polyplosphaeria guizhouensis (GZAAS 23-0600, holotype) a colonies on decaying wood b, c colonies on natu- ral substrates d-n conidia bearing appendages o germinating conidium p colony on PDA (front at right, reverse at left). Scale bars: 50 um (d-o). MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 180 Xia Tang et al.: Novel fungi from China Table 2. Tetraplosphaeriaceae species and their country, life cycle, habitat, host and reference. Species name Country Aquatisphaeria Thailand thailandica Byssolophis byssiseda France B. sphaerioides Finland, UK Ernakulamia Argentina, Cuba, cochinensis India, Japan, Malaysia, Mexico, Panama, Thailand E. krabiensis Thailand E. tanakae India E. xishuangbannaensis China Polyplosphaeria China guizhouensis Po. Hainanensis China Po. Fusca Japan Po. thailandica Thailand Po. nabanheensis China Po. pandanicola China Pseudotetraploa China bambusicola Ps. curviappendiculata Japan Ps. yunnanensis China Ps. Javanica Indonesia, Japan Ps. longissima Japan Ps. rajmachiensis India Quadricrura bicornis Japan Q. meridionalis Japan Q. septentrionalis Japan Shrungabeeja Thailand aquatica Life cycle Habitat saprobic | freshwater saprobic | terrestrial saprobic | terrestrial saprobic | freshwater, terrestrial saprobic | terrestrial saprobic | terrestrial saprobic | terrestrial saprobic | terrestrial saprobic _ terrestrial saprobic | terrestrial saprobic | terrestrial saprobic | terrestrial saprobic | terrestrial saprobic | terrestrial saprobic | terrestrial saprobic | freshwater saprobic | terrestrial saprobic | terrestrial saprobic | terrestrial saprobic | terrestrial saprobic | terrestrial saprobic | terrestrial saprobic | freshwater MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 Host decaying wood branch of Carpinus, decaying wood decaying stemp of Rubus, decaying wood birch Astrocaryum standleyanum, Benthamidia japonica, dead leaves, dead spathes of Cocos nucifera, decomposing leaves of Satakentia liukivensis, Freycinetia multi, palm tree, /lex sp., Ocotea leucoxylon, Pandanus tectorius, P monticola, submerged wood, Syagrus romanzoffiana, Stewartia monadelpha, Vitex sp. Acacia sp. dead spathes of Cocos nucifera dead leaves of Pandanus sp. unidentified decaying wood unidentified decaying wood culms of Chimonobambusa marmorea, culms of Phyllostachys bambusoides, culms of Pleioblastus chino, culms of Sasa kurilensis decaying bamboo decaying leaves of Pandanus sp. decaying leaves of Pandanus sp. dead branches of Bamboo culms of Sasa kurilensis bamboo 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 culms of Pleioblastus chino decaying bamboo culms, Dendrocalamus stocksii (Poaceae) culms of Sasa kurilensis, leaf litter of a conifer bamboo culms of Sasa kurilensis submerged wood Reference Li et al. (2021) Zhang et al. (2012) Berkeley and Broome (1854); Karsten (1870) Ellis (1976); Holubova- Jechova and Mercado (1986); Holubova-Jechova (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) Jayasiri et al. (2019) Hyde et al. (2020a) Tibpromma et al. (2018); Dong et al. (2020) This study This study Tanaka et al. (2009) Li et al. (2016) Tibpromma et al. (2018) Tibpromma et al. (2018) Yu et al. (2022) Tanaka et al. (2009) This study Hatakeyama et al. (2005); Tanaka et al. (2009); Rifai et al. (2014) Tanaka et al. (2009) Hyde et al. (2020a) Tanaka et al. (2009) Tanaka et al. (2009) Tanaka et al. (2009) Dong et al. (2020) 181 Xia Tang et al.: Novel fungi from China Species name S. longiappendiculata S. vadirajensis S. begoniae S. melicopes S. piepenbringiana S. fluviatilis Tetraploa abortiva T. aquatica T. aristata T. bambusae T. biformis T. circinata T. conata F T. cylindrica T. dashaoensis T. divergens T. dwibahubeeja T. ellisii T. endophytica T. hainanensis T. indica T. josettae™ T. juncicola Country Thailand Brazil, China, India China China Panama China Argentina China 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) China Japan India India China China USA (Mississippi) India Argentina, USA (New Jersey), Zimbabwe Germany China India France The Netherlands Life cycle saprobic saprobic saprobic saprobic saprobic saprobic saprobic saprobic pathogenic (human), saprobic saprobic saprobic saprobic N/A saprobic saprobic saprobic saprobic saprobic endophytic Saprobic N/A N/A saprobic Habitat terrestrial terrestrial terrestrial terrestrial terrestrial freshwater freshwater freshwater terrestrial terrestrial terrestrial terrestrial N/A terrestrial terrestrial terrestrial terrestrial terrestrial terrestrial terrestrial N/A N/A terrestrial MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 Host dead culm of Bambusa sp. (Poaceae) dead branches of unidentified plant dead branches of Begonia semperflorens dead branches of Melicope triphylla dead Poaceae submerged decaying twig N/A submerged decaying wood 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 dead twigs of bamboo dead bark of broad-leaved tree decaying bamboo twig N/A decaying stems of Saccharum arundinaceum (Poaceae) dead stem of Saccharum arundinaceum leaves of Panicum agrostidiforme decaying spathes of Cocos nucifera Chloris, Dactylis, Hevea brasiliensis, stalks of Zea mays roots of Microthlaspi perfoliatum unidentified decaying wood N/A N/A dead culm of Juncus inflexus (Juncaceae) Reference Ariyawansa et al. (2015) Rao and Reddy (1981); Zhang et al. (2009) Zhang et al. (2009) Zhang et al. (2009) Kirschner et al. (2017) Yang et al. (2023) Arambarri et al. (1987) Li et al. (2020) Ellis (1949, 1971); Markham et al. (1990); Tanaka et al. (2009); Senwanna et al. (2021) Phookamsak et al. (2022) Matsushima and Matsushima (1996) Pratibha and Bhat (2008) Saxena and Sarkar (1986); Gupta (2002) Liao et al. (2022) Jayawardena et al. (2023) Tracy and Earle (1895) Hyde et al. (2020a) Cooke and Ellis (1879); Ellis (1971); Senwanna et al. (2021) Crous et al. (2021) This study Saxena and Khare (1991) Nufiez Otafo et al. (2022) Crous et al. (2022) 182 Xia Tang et al.: Novel fungi from China Species name Country T. muscicola Spain T. nagasakiensis Japan, China T. obpyriformis China T. opaca China T. pseudoaristata India T. puzheheiensis China T. sasicola China, Japan T. scabra USA T. scheueri UK T. setifera Hungary T. siwalika N/A T. taugourdeaui* India T. thailandica Thailand T. thrayabahubeeja India T. yakushimensis Japan T. yunnanensis Tetraploa sp. 1 Tetraploa sp. 2 Triplosphaeria cylindrica Tr. maxima Tr. yezoensis Tr. acuta Triplosphaeria sp. China, Thailand Japan Japan Japan Japan Japan Japan Japan Life cycle N/A saprobic saprobic saprobic saprobic saprobic saprobic N/A saprobic saprobic N/A N/A saprobic saprobic saprobic saprobic saprobic saprobic saprobic saprobic saprobic saprobic saprobic Habitat Host Reference N/A fronds of Aneura multifida, Gonzalez Fragoso (1916) Lophozia quinquedentata terrestrial culms of bamboo Hyde et al. (2013, 2019) terrestrial dead grass under Saccharum Unpublished arundinaceum (Gramineae) terrestrial dead culms of bamboo, Zhao et al. (2009) decaying branches of unidentified tree terrestrial decaying spathes of Cocos Hyde et al. (2020a) nucifera (Arecacceae) freshwater submerged wood Dong et al. (2020) terrestrial | culms of Sasa senanensis, dead | Tanaka et al. (2009); Hyde leaves of Pennisetum purpureum et al. (2020a) (Poaceae) terrestrial Scirpus sp. Harkness (1885) freshwater, leaves of Carex acutiformis, Scheuer (1991); Hyde et terrestrial rotten leaves al. (2013) terrestrial rotten wood Révay (1993) N/A N/A Saxena et al. (1987) N/A N/A Saxena and Sarkar (1986) freshwater Submerged decaying wood Bao et al. (2021) terrestrial decaying spathes of Cocos Hyde et al. (2020a) nucifera (Arecacceae) terrestrial culms of Arundo donax Tanaka et al. (2009); Hyde et al. (2013) freshwater submerged wood Dong et al. (2020) terrestrial culms of bamboo Tanaka et al. (2009) terrestrial culms of Gramineae Tanaka et al. (2009) terrestrial culms of Sasa kurilensis Tanaka et al. (2009) terrestrial culms of Sasa kurilensis Tanaka et al. (2009) terrestrial culms of Sasa palmata Tanaka et al. (2009) freshwater | submerged culms of bamboo Tanaka et al. (2009) terrestrial culms of Sasa kurilensis Tanaka et al. (2009) Fossil fungi were indicated “F”; N/A: Not available or cannot find; Newly species indicate in bold. Polyplosphaeria hainanensis X. Tang, Jayaward., R. Jeewon & J.C. Kang, sp. nov. MycoBank No: 900951 Facesoffungi Number: FoF 14665 Figs 3,4 Etymology. The specific epithet ‘hainanensis’ refers to the place where the fun- gus was collected, Hainan Province, China. Holotype. GZAAS 23-0601 Description. Saprobic on unidentified decaying wood in the forest. Teleo- morph not observed. Anamorph Hyphomycetous. Colonies effuse, gregarious on host substrate, brown to blackish brown. Mycelium semi-immersed or im- mersed, dark brown, branched, septate. Conidiophores absent. Conidiogenous MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 183 Xia Tang et al.: Novel fungi from China Figure 3. Polyplosphaeria hainanensis (GZAAS 23-0601, holotype) a colonies on decay wood b colonies on natural sub- strates c—-m conidia bearing appendages n germinating conidium o colony on PDA (from front) p colony on PDA (from reverse). Scale bars: 50 um (c=n). MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 184 Xia Tang et al.: Novel fungi from China Figure 4. Polyplosphaeria hainanensis (GZAAS 23-0602, paratype) a colonies on decay wood b, ¢ colonies on natural substrates d-o conidia bearing appendages p colony on PDA (from front) q colony on PDA (from reverse). Scale bars: 100 um (d-i, k-m); 50 um (j, n, 0). MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 185 Xia Tang et al.: Novel fungi from China cells indistinguishable from creeping hyphae, integrated, monoblastic, determi- nate. Conidia 49-134.5 x 52—-90.5 um (x = 86 x 71 um, n = 20), globose, subglo- bose, obconical, broadly ellipsoidal to broadly pyriform, variable in shape, some- times 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 x 3-5.5 um (x = 44.5 x 4 um, n = 20), cylindrical, solitary, straight or flexuous, unbranched and almost hyaline at the apex, 0-4-septate, smooth, round at apex, pervasive. Culture characteristics. Conidia germinated from both ends on PDA and in- cubated at room temperature (25 °C). Colonies circular, cottony, flat, olivaceous with a slightly grey entire margin. The reverse side is an olive drab, which grad- ually extends outwards to form a deep colour ring in the centre with a pale grey margin and no pigment. Material examined. CHINA, Hainan Province, Wuzhishan City, Wuzhishan Na- tional 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), re- spectively, 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 mor- phological 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 Polyplosphae- ria 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 Facesoffungi Number: FoF06669 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 super- ficial. Conidiophores absent. Conidiogenous cells monoblastic, indistinguish- able 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 etal. (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 MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 186 Xia Tang et al.: Novel fungi from China are reported as saprobes on bamboo, dead bark of the broad-leaved tree, and un- identified 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 Facesoffungi Number: FoF 14666 FIg.<5 Etymology. The specific epithet ‘yunnanensis’ refers to the place where the fun- gus 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. Co- nidiogenous cells micronematous, mononematous, monoblastic, integrated, usually undistinguishable from superficial hyphae. Conidia 67-120 x 16.5- 35 um (x = 95 x 24 um, 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 um wide (x = 26 um, n = 20), slightly constricted at septa, rarely branched and make V-shaped conidia; setose appendages at the apical part 15-87 x 3.5-7 um (x = 37 x 5 um, n = 20), appendages 3-6 in number, 1-8-septate, brown at the base and almost hyaline at the apex, smooth, un- branched, shorter appendage is straight and longer appendage is curved. Culture characteristics. Conidia germinated from both ends on PDA and in- cubated 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. curviappen- diculata in having branched and V-shaped conidia, consisting of 3-6 columns of cells with 3-6 apical appendages, larger conidia [67-120 um vs. 52-67(-75) um] in length and [16-35 um vs. 15-22 um] in width, while Ps. curviappendiculata consists of 4-5 columns of cells with 4 apical appendages; Pseudotetraploa yun- nanensis differs from Ps. longissima in having smaller conidia [67-120 um vs. (98-)110-148(-155) um] in length and [16-35 um vs. 18-25 um] in width, with- out verrucose at the base. The phylogenetic analysis showed that Pseudotetrap- loa yunnanensis is sister to Ps. rajmachiensis and Ps. javanica. The comparison of MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 187 Xia Tang et al.: Novel fungi from China ? 7 ao ~ wr yas fed ‘c Sy, -) oe pone Tet. ~ Figure 5. Pseudotetraploa yunnanensis (HKAS 129442, holotype) a, b colonies on natural substrates c—n conidia. Scale bars: 20 um (c); 50 um (d=n). 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; Pseudotet- raploa 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). MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 188 Xia Tang et al.: Novel fungi from China Tetraploa Berk. & Broome, Ann. Mag. Nat. Hist. 5: 459, t. 11:6 (1850) MycoBank No: 10199 Facesoffungi Number: FoF06666 = 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 sen- su stricto Conidiophores absent. Conidiogenous cells monoblastic. Conidia composed of 4 columns, short-cylindrical, brown, verrucose at the base, eusep- tate, 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 Tetrap/oa sensu stricto based on a combined SSU and LSU DNA sequence data. Later, Hyde et al. (2013) treated Tetraploa as asynonym of Tetraplosphaeria, which has been applied previously to anamorphic species and used Tetraploa instead of Tetraplosphaeria. Species of Tetraploa are main- ly 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 Tet- raploa. Nufiez Otafo 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 Facesoffungi Number: FoF 14667 Figs 6, 7 Etymology. The specific epithet ‘hainanensis’ refers to the place where the fun- gus 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 inte- grated, monoblastic, determinate. Conidia 30-46 x 18-36 um (x = 38 x 27 um, 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 MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 189 Xia Tang et al.: Novel fungi from China Figure 6. Tetraploa hainanensis (GZAAS 23-0603, holotype) a, b colonies on natural substrates e-n conidia bearing 1-5 appendages 0 germinating conidium p colony on PDA (from reverse) q colony on PDA (from front). Scale bars 20 um (c); 50 pm (dg); 100 pm (h-o). MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 190 Xia Tang et al.: Novel fungi from China Figure 7. Tetraploa hainanensis (GZAAS 23-0604, paratype) a colonies on decay wood b, ¢ colonies on natural sub- strates 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 um (dg); 100 um (h-I, 0, q); 50 um (m, n, p). MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 191 Xia Tang et al.: Novel fungi from China column, smooth, mostly with four apical appendages, some with one or two or five appendages. Appendages 52-209 x 3-6 um (x = 140 x 4 um, 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. Culture characteristics. Conidia germinated from both ends on PDA and in- cubated 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. 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 x 18-36 um vs. 22-31 x 15-20 um) with four col- umns of cells, sometimes five columns of cells, and longer appendages (52-209 x 3-6 um vs. 23-107 x 2-5 um), 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. yakushi- mensis has only four columns and appendages; Tetraploa hainanensis differs from T. tetraploa in having four or five columns and shorter appendages (52-209 x 3-6 um vs. 263-350 x 2-3 um), while T. tetraploa has only four columns and slen- der 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 phy- logeny, 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 north- ern 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 es- timates. 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 colonis- ing them. Our study reveals four anamorphic fungal species that belong to the family Tetraplosphaeriaceae. In this study, we characterise two new anamor- phic species collected from unidentified decayed wood samples that belong to Polyplosphaeria. In Polyplosphaeria this study brings the number of species to MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 192 Xia Tang et al.: Novel fungi from China six. The first new species is described as Po. guizhouensis and our multigene phylogeny depict a close relationship to Po. pandanicola. The latter was collect- ed 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 gut- tules, 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 re- gard to DNA sequence data comparison, Po. guizhouensis differs from Po. pan- danicola (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% dif- ference) 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 Polyplos- phaeria 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 sig- nificant statistical support, a pattern observed in various previous studies as well (Li et al. 2021; Liao et al. 2022). Aquatisphaeria thailandica has been report- ed 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 phylogenet- ic analyses, it seems that Po. thailandica is a member of Aquatisphaeria. There is not much taxonomic data available for Aguatisphaeria, 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 MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 193 Xia Tang et al.: Novel fungi from China reported, and this study extends the known species count to six. The new spe- cies 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 spe- cies, the genus now comprises a total of 35 species. The new species is de- scribed as T. hainanensis, and the multigene phylogeny depicts a close relation- ship 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 most- ly 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 Tetrap- losphaeriaceae, 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 Lu- ang University for its support in the tuition fee scholarship. This work was sup- ported 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. MycoKeys 100: 171-204 (2023), DOI: 10.3897/mycokeys.100.113141 194 Xia Tang et al.: Novel fungi from China 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 Re- search 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. References Ando K (1992) A study of terrestrial aquatic hyphomycetes. Nippon Kingakkai Kaiho 33: 415-425. [In Japanese] Arambarri A, Cabello M, Mengascini A (1987) New hyphomycetes from Santiago River. Il. (Buenos Aires Province, Argentina). Mycotaxon 30: 263-267. 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