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Published: 2021-09-03

Freshwater Sordariomycetes: new species and new records in Pleurotheciaceae, Pleurotheciales

Faculty of Environmental Science and Engineering, Kunming University of Science & Technology, Kunming 650500, P.R. China
Faculty of Agriculture and Food, Kunming University of Science & Technology, Kunming 650500, P.R. China
Center of Excellence in Fungal Research, Mae Fah Luang University, Chiang Rai 57100, Thailand; Institute of Plant Health, Zhongkai University of Agriculture and Engineering, Haizhu District, Guangzhou 510225, P.R. China
Center for Yunnan Plateau Biological Resources Protection and Utilization, College of Biological Resource and Food Engineering, Qujing Normal University, Qujing 655011, P.R. China
Faculty of Environmental Science and Engineering, Kunming University of Science & Technology, Kunming 650500, P.R. China
Faculty of Agriculture and Food, Kunming University of Science & Technology, Kunming 650500, P.R. China; School of Life Science and Technology, University of Electronic Science and Technology of China, Chengdu 611731, P.R. China
Faculty of Agriculture and Food, Kunming University of Science & Technology, Kunming 650500, P.R. China; Department of Botany, University of British Columbia, Vancouver, Canada; Yunnan Key Lab of Soil Carbon Sequestration and Pollution Control, Kunming University of Science and Technology, Kunming 650500, P.R. China
2 new taxa asexual morphs denticulate morphology Phaeoisaria phylogeny Pleurotheciella Pleurothecium Fungi


Eight freshwater asexual strains collected from China and Thailand were subjected to morpho-molecular analyses. BLAST search results showed that the new strains belonged to Pleurotheciales, Sordariomycetes. Phylogenetic analyses based on a combined LSU, SSU, ITS and RPB2 sequence dataset are used to clarify the placement of the new strains within Pleurotheciales. Pleurothecium guttulatum and Pleurotheciella sympodia are introduced as two new species from freshwater habitats. New geographical record of Pleurotheciella nilotica from Asia (Thailand and China) is reported. New habitat and geographical records of Phaeoisaria annesophieae (from freshwater habitats in Asia) are reported. Descriptions and illustrations of the new species and records, as well as known species Rhexoacrodictys erecta and Phaeoisaria annesophieae, are provided. A key to Pleurotheciella species is provided.


  1. Abdel-Aziz, F.A., Bahkali, A.H., Elgorban, A.M. & Abdel-Wahab, M.A. (2020) Pleurotheciella nilotica sp. nov. (Pleurotheciales, Ascomycota) from freshwater habitats in Egypt. Nova Hedwigia 110: 91–98.

  2. Baker, W.A., Partridge, E.C. & Morgan-Jones, G. (2002) Notes on hyphomycetes LXXXVII. Rhexoacrodictys, a new segregate genus to accommodate four species previously classified in Acrodictys. Mycotaxon 82: 95–113. [Accession Number: WOS:000177167900003]

  3. Bao, D.F., McKenzie, E.H.C., Bhat, D.J., Hyde, K.D., Luo, Z.L., Shen, H.W. & Su, H.Y. (2020) Acrogenospora (Acrogenosporaceae, Minutisphaerales) appears to be a very diverse genus. Frontiers in Microbiology 11: 1606–1629.

  4. Cheng, X.L., Li, W. & Zhang, T.Y. (2014) A new species of Phaeoisaria from intertidal marine sediment collected in Weihai, China. Mycotaxon 127: 17–24.

  5. Chomnunti, P., Hongsanan, S., Aguirre-Hudson, B., Tian, Q., Persoh, D., Dhami, M., Alias, S.A., Xu, J.C., Liu, X.Z., Stadler, M. & Hyde, K.D. (2014) The sooty moulds. Fungal Diversity 66: 1–36.

  6. Crous, P.W., Wingfield, M.J., Burgess, T.I., Carnegie, A.J., Hardy, G., Smith, D., Summerell, B.A., Cano-Lira, J.F., Guarro, J., Houbraken, J., Lombard, L.,. Martin, M.P., Sandoval-Denis, M., Alexandrova, A.V., Barnes, C.W., Baseia, I.G., Bezerra, J.D.P., Guarnaccia, V., May, T.W., Hernandez-Restrepo, M., Stchigel, A.M., Miller, A.N., Ordonez, M.E., Abreu, V.P., Accioly, T., Agnello, C., Agustin Colman, A., Albuquerque, C.C., Alfredo, D.S., Alvarado, P., Araujo-Magalhaes, G.R., Arauzo, S., Atkinson, T., Barili, A., Barreto, R.W., Bezerra, J.L., Cabral, T.S., Camello Rodriguez, F., Cruz, R., Daniels, P.P., da Silva, B.D.B., de Almeida, D.A.C., de Carvalho Junior, A.A., Decock, C.A., Delgat, L., Denman, S., Dimitrov, R.A., Edwards, J., Fedosova, A.G., Ferreira, R.J., Firmino, A.L., Flores, J.A., Garcia, D., Gene, J., Giraldo, A., Gois, J.S., Gomes, A.A.M., Goncalves, C.M., Gouliamova, D.E., Groenewald, M., Gueorguiev, B.V., Guevara-Suarez, M., Gusmao, L.F.P., Hosaka, K., Hubka, V., Huhndorf, S.M., Jadan, M., Jurjevic, Z., Kraak, B., Kucera, V., Kumar, T.K.A., Kusan, I., Lacerda, S.R., Lamlertthon, S., Lisboa, W.S., Loizides, M., Luangsa-Ard, J.J., Lyskova, P., Mac Cormack, W.P., Macedo, D.M., Machado, A.R., Malysheva, E.F., Marinho, P., Matocec, N., Meijer, M., Mesic, A., Mongkolsamrit, S., Moreira, K.A., Morozova, O.V., Nair, K.U., Nakamura, N., Noisripoom, W., Olariaga, I., Oliveira, R.J.V., Paiva, L.M., Pawar, P., Pereira, O.L., Peterson, S.W., Prieto, M., Rodriguez-Andrade, E., Rojo De Blas, C., Roy, M., Santos, E.S., Sharma, R., Silva, G.A., Souza-Motta, C.M., Takeuchi-Kaneko, Y., Tanaka, C., Thakur, A., Smith, M.T., Tkalcec, Z., Valenzuela-Lopez, N., van der Kleij, P., Verbeken, A., Viana, M.G., Wang, X.W. & Groenewald, J.Z. (2017) Fungal Planet description sheets: 625–715. Persoonia 39: 270–467.

  7. Dayarathne, M.C., Maharachchikumbura, S.S.N., Jones, E.B.G., Dong, W., Devadatha, B., Yang, J., Ekanayaka, A.H., De Silva, W., Sarma, V.V., Al-Sadi, A.M., Khongphinitbunjong, K., Hyde, K.D. & Zhao, R.L. (2019) Phylogenetic revision of Savoryellaceae and evidence for its ranking as a subclass. Frontiers in Microbiology 10: 840–867.

  8. Dong, W., Hyde, K.D., Bhat, D.J. & Zhang, H. (2018) Introducing Aculeata aquatica gen. et sp. nov., Minimelanolocus thailandensis sp. nov. and Thysanorea aquatica sp. nov. (Herpotrichiellaceae, Chaetothyriales) from freshwater in northern Thailand. Mycological Progress 17: 617–629.

  9. Dong, W., Hyde, K.D., Doilom, M., Yu, X.D., Bhat, D.J., Jeewon, R., Boonmee, S., Wang, G.N., Nalumpang, S. & Zhang, H. (2020) Pseudobactrodesmium (Dactylosporaceae, Eurotiomycetes, Fungi) a novel lignicolous genus. Frontiers in Microbiology 11: 456–469.

  10. Dong, W., Wang, B., Hyde, K.D., McKenzie, E.H.C., Raja, H.A., Tanaka, K., Abdel-Wahab, M.A., Abdel-Aziz, F.A., Doilom, M., Phookamsak, R., Hongsanan, S., Wanasinghe, D.N., Yu, X.D., Wang, G.N., Yang, H., Yang, J., Thambugala, K.M., Tian, Q., Luo, Z.L., Yang, J.B., Miller, A.N., Fournier, J., Boonmee, S., Hu, D.M., Nalumpang, S. & Zhang, H. (2021) Freshwater Dothideomycetes. Fungal Diversity 105: 319–575.

  11. Fernández, F.A., Lutzoni, F.M. & Huhndorf, S.M. (1999) Teleomorph-anamorph connections: the new pyrenomycetous genus Carpoligna and its Pleurothecium anamorph. Mycologia 91: 251–262.

  12. Guo, J.S., Zhang, Z., Qiao, M. & Yu, Z.F. (2019) Phalangispora sinensis sp. nov. from Yunnan, China and two new members of Wiesneriomycetaceae. International Journal of Systematic and Evolutionary Microbiology 69: 3217–3223.

  13. Hall, T. (1999) BioEdit: A user-friendly biological sequence alignment program for Windows 95/98/NT. Nucleic Acids Symposium Series 41: 95–98.

  14. Hernández-Restrepo, M., Gene, J., Castaneda-Ruiz, R.F., Mena-Portales, J., Crous, P.W. & Guarro, J. (2017) Phylogeny of saprobic microfungi from Southern Europe. Studies in Mycology 86: 53–97.

  15. Höhnel, F. (1919) Fünfte vorlaüfige Mitteilungen mykologischer Ergebnisse (Nr. 399–500). Berichte der Deutschen Botanischen Gesellschaft 37: 153–161.

  16. Höhnel, F. (1923) Studien über Hyphomyceten. Zentralbl Bakteriol Parasitenk Infektionskrankh 60: 1–26.

  17. Hongsanan, S., Maharachchikumbura, S.S.N., Hyde, K.D., Samarakoon, M.C., Jeewon, R., Zhao, Q., Al-Sadi, AM. & Bahkali, A.H. (2017) An updated phylogeny of Sordariomycetes based on phylogenetic and molecular clock evidence. Fungal Diversity 84: 25–41.

  18. Ho, W.H., Hyde, K.D. & Hodgkiss, J.I. (2001) Fungal communities on submerged wood from streams in Brunei, Hong Kong, and Malaysia. Mycological Research 105: 1492–1501.

  19. Ho, W.H., Hyde, K.D. & Hodgkiss, I.J. (2002) Seasonality and sequential occurrence of fungi on wood submerged in Tai Po Kau Forest Stream, Hong Kong. Fungal Diversity 10: 21–43. [Accession Number: WOS:000176727700004]

  20. Hyde, K.D. & Goh, T.K. (1998a) Fungi on submerged wood in Lake Barrine, north Queensland, Australia. Mycological Research 102: 739–749.

  21. Hyde, K.D. & Goh, T.K. (1998b) Fungi on submerged wood in the Riviere St Marie-Louis, The Seychelles. South African Journal of Botany 64: 330–336.

  22. Hyde, K.D., Fryar, S., Tian, Q., Bahkali, A.H. & Xu, J. (2016a) Lignicolous freshwater fungi along a north–south latitudinal gradient in the Asian/Australian region; can we predict the impact of global warming on biodiversity and function? Fungal Ecology 19: 190–200.

  23. Hyde, K.D., Hongsanan, S., Jeewon, R., Bhat, D.J., McKenzie, E.H.C., Jones, E.B.G., Phookamsak, R., Ariyawansa, H.A., Boonmee, S., Zhao, Q., Abdel-Aziz, F.A., Abdel-Wahab, M.A., Banmai, S., Chomnunti, P., Cui, B.K., Daranagama, D.A., Das, K., Dayarathne, M.C., de Silva, N.I., Dissanayake, A.J., Doilom, M., Ekanayaka, A.H., Gibertoni, T.B., Goes-Neto, A., Huang, S.K., Jayasiri, S.C., Jayawardena, R.S., Konta, S., Lee, H.B., Li, W.J., Lin, C.G., Liu, J.K., Lu, Y.Z., Luo, Z.L., Manawasinghe, I.S., Manimohan, P., Mapook, A., Niskanen, T., Norphanphoun, C., Papizadeh, M., Perera, R.H., Phukhamsakda, C., Richter, C., Santiago, A., Drechsler-Santos, E.R., Senanayake, I.C., Tanaka, K., Tennakoon, T., Thambugala, K.M., Tian, Q., Tibpromma, S., Thongbai, B., Vizzini, A., Wanasinghe, D.N., Wijayawardene, N.N., Wu, H.X., Yang, J., Zeng, X.Y., Zhang, H., Zhang, J.F., Bulgakov, T.S., Camporesi, E., Bahkali, A.H., Amoozegar, M.A., Araujo-Neta, L.S., Ammirati, J.F., Baghela, A., Bhatt, R.P., Bojantchev, D., Buyck, B., da Silva, G.A., de Lima, C.L.F., de Oliveira, R.J.V., de Souza, C.A.F., Dai, Y.C., Dima, B., Duong, T.T., Ercole, E., Mafalda-Freire, F., Ghosh, A., Hashimoto, A., Kamolhan, S., Kang, J.C., Karunarathna, S.C., Kirk, P.M., Kytovuori, I., Lantieri, A., Liimatainen, K., Liu, Z.Y., Liu, X.Z., Lucking, R., Medardi, G., Mortimer, P.E., Nguyen, T.T.T., Promputtha, I., Raj, K.N.A., Reck, M.A., Lumyong, S., Shahzadeh-Fazeli, S.A., Stadler, M., Soudi, M.R., Su, H.Y., Takahashi, T., Tangthirasunun, N., Uniyal, P., Wang, Y., Wen, T.C., Xu, J.C., Zhang, Z.K., Zhao, Y.C., Zhou, J.L. & Zhu, L. (2016b) Fungal diversity notes 367–490: taxonomic and phylogenetic contributions to fungal taxa. Fungal Diversity 80: 1–270.

  24. Hyde, K.D., Chaiwan, N., Norphanphoun, C., Boonmee, S., Camporesi, E., Chethana, K.W.T., Dayarathne, M.C., de Silva, N.I., Dissanayake, A.J., Ekanayaka, A.H., Hongsanan, S., Huang, S.K., Jayasiri, S.C., Jayawardena, R.S., Jiang, H.B., Karunarathna, A., Lin, C.G., Liu, J.K., Liu, N.G., Lu, Y.Z., Luo, Z.L., Maharachchimbura, S.S.N., Manawasinghe, I.S., Pem, D., Perera, R.H., Phukhamsakda, C., Samarakoon, M.C., Senwanna, C., Shang, Q.J., Tennakoon, D.S., Thambugala, K.M., Tibpromma, S., Wanasinghe, D.N., Xiao, Y.P., Yang, J., Zeng, X.Y., Zhang, J.F., Zhang, S.N., Bulgakov, T.S., Bhat, D.J., Cheewangkoon, R., Goh, T.K., Jones, E.B.G., Kang, J.C., Jeewon, R., Liu, Z.Y., Lumyong, S., Kuo, C.H., McKenzie, E.H.C., Wen, T.C., Yan, J.Y. & Zhao, Q. (2018) Mycosphere notes 169–224. Mycosphere 9: 271–430.

  25. Hyde, K.D., Tennakoon, D.S., Jeewon, R., Bhat, D.J., Maharachchikumbura, S.S.N., Rossi, W., Leonardi, M., Lee, H.B., Mun, H.Y., Houbraken, J., Nguyen, T.T.T., Jeon, S.J., Frisvad, J.C., Wanasinghe, D.N., Luecking, R., Aptroot, A., Caceres, M.E.S., Karunarathna, S.C., Hongsanan, S., Phookamsak, R., de Silva, N.I., Thambugala, K.M., Jayawardena, R.S., Senanayake, I.C., Boonmee, S., Chen, J., Luo, Z.L., Phukhamsakda, C., Pereira, O.L., Abreu, V.P., Campos Rosado, A.W., Bart, B., Randrianjohany, E., Hofstetter, V., Gibertoni, T.B., da Silva Soares, A.M., Plautz, H.L., Jr., Pontes Sotao, H.M., Silva Xavier, W.K., Pereira Bezerra, J.D., Lins de Oliveira, T.G., de Souza-Motta, C.M., Correia Magalhaes, O.M., Bundhun, D., Harishchandra, D., Manawasinghe, I.S., Dong, W., Zhang, S.N., Bao, D.F., Samarakoon, M.C., Pem, D., Karunarathna, A., Lin, C.G., Yang, J., Perera, R.H., Kumar, V., Huang, S.K., Dayarathne, M.C., Ekanayaka, A.H., Jayasiri, S.C., Xiao, Y., Konta, S., Niskanen, T., Liimatainen, K., Dai, Y.C., Ji, X.H., Tian, X.M., Mesic, A., Singh, S.K., Phutthacharoen, K., Cai, L., Sorvongxay, T., Thiyagaraja, V., Norphanphoun, C., Chaiwan, N., Lu, Y.Z., Jiang, H.B., Zhang, J.F., Abeywickrama, P.D., Aluthmuhandiram, J.V.S., Brahmanage, R.S., Zeng, M., Chethana, T., Wei, D.P., Reblova, M., Fournier, J., Nekvindova, J., Barbosa, R.d.N., Felinto dos Santos, J.E., de Oliveira, N.T., Li, G.J., Ertz, D., Shang, Q.J., Phillips, A.J.L., Kuo, C.-H., Camporesi, E., Bulgakov, T.S., Lumyong, S., Jones, E.B.G., Chomnunti, P., Gentekaki, E., Bungartz, F., Zeng, X.Y., Fryar, S., Tkalcec, Z., Liang, J.M., Li, G., Wen, T.C., Singh, P.N., Gafforov, Y., Promputtha, I., Yasanthika, E., Goonasekara, I.D., Zhao, R.L., Zhao, Q., Kirk, P.M., Liu, J.K., Yan, J.Y.., Mortimer, P.E., Xu, J.C. & Doilom, M. (2019) Fungal diversity notes 1036–1150: taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 96: 1–242.

  26. Hyde, K.D., Norphanphoun, C., Maharachchikumbura, S.S.N., Bhat, D.J., Jones, E.B.G., Bundhun, D., Chen, Y.J., Bao, D.F., Boonmee, S., Calabon, M.S., Chaiwan, N., Chethana, K.W.T., Dai, D.Q., Dayarathne, M.C., Devadatha, B., Dissanayake, A.J., Dissanayake, L.S., Doilom, M., Dong, W., Fan, X.L., Goonasekara, I.D., Hongsanan, S., Huang, S.K., Jayawardena, R.S., Jeewon, R., Karunarathna, A., Konta, S., Kumar, V., Lin, C.G., Liu, J.K., Liu, N.G., Luangsaard, J., Lumyong, S., Luo, Z.L., Marasinghe, D.S., McKenzie, E.H.C., Niego, A.G.T., Niranjan, M., Perera, R.H., Phukhamsakda, C., Rathnayaka, A.R., Samarakoon, M.C., Samarakoon, S.M.B.C., Sarma, V.V., Senanayake, I.C., Shang, Q.J., Stadler, M., Tibpromma, S., Wanasinghe, D.N., Wei, D.P., Wijayawardene, N.N., Xiao, Y.P., Yang, J., Zeng, X.Y., Zhang, S.N. & Xiang, M.M. (2020a) Refined families of Sordariomycetes. Mycosphere 11: 305–1059.

  27. Hyde, K.D., Dong, Y., Phookamsak, R., Jeewon, R., Bhat, D.J., Jones, E.B.G., Liu, N.G., Abeywickrama, P.D., Mapook, A., Wei, D.P., Perera, R.H., Manawasinghe, I.S., Pem, D., Bundhun, D., Karunarathna, A., Ekanayaka, A.H., Bao, D.F., Li, J., Samarakoon, M.C., Chaiwan, N., Lin, C.G., Phutthacharoen, K., Zhang, S.N., Senanayake, I.C., Goonasekara, I.D., Thambugala, K.M., Phukhamsakda, C., Tennakoon, D.S., Jiang, H.B., Yang, J., Zeng, M., Huanraluek, N., Liu, J.K., Wijesinghe, S.N., Tian, Q., Tibpromma, S., Brahmanage, R.S., Boonmee, S., Huang, S.K., Thiyagaraja, V., Lu, Y.Z., Jayawardena, R.S., Dong, W., Yang, E.F., Singh, S.K., Singh, S.M., Rana, S., Lad, S.S., Anand, G., Devadatha, B., Niranjan, M., Sarma, V.V., Liimatainen, K., Aguirre-Hudson, B., Niskanen, T., Overall, A., Alvarenga, R.L.M., Gibertoni, T.B., Pfliegler, W.P., Horvath, E., Imre, A., Alves, A.L., Santos, A.C.d.S., Tiago, P.V., Bulgakov, T.S., Wanasinghe, D.N., Bahkali, A.H., Doilom, M., Elgorban, A.M., Maharachchikumbura, S.S.N., Rajeshkumar, K.C., Haelewaters, D., Mortimer, P.E., Zhao, Q., Lumyong, S., Xu, J.C. & Sheng, J. (2020b) Fungal diversity notes 1151–1276: taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 100: 5–277.

  28. Hyde, K.D., Jeewon, R., Chen, Y.J., Bhunjun, C.S., Calabon, M.S., Jiang, H.-B., Lin, C.G., Norphanphoun, C., Sysouphanthong, P., Pem, D., Tibpromma, S., Zhang, Q., Doilom, M., Jayawardena, R.S., Liu, J.-K., Maharachchikumbura, S.S.N., Phukhamsakda, C., Phookamsak, R., Al-Sadi, A.M., Thongklang, N., Wang, Y., Gafforov, Y., Jones, E.B.G. & Lumyong, S. (2020c) The numbers of fungi: is the descriptive curve flattening? Fungal Diversity 103: 219–271.

  29. Index Fungorum (2021) Available from: (accessed 3 September 2021)

  30. Jeewon, R. & Hyde, K.D. (2016) Establishing species boundaries and new taxa among fungi: Recommendations to resolve taxonomic ambiguities. Mycosphere 7: 1669–1677.

  31. Kurniawati, E., Zhang, H., Chukeatirote, E., Sulistyowati, L., Moslem, M.A. & Hyde, K.D. (2010) Diversity of freshwater ascomycetes in freshwater bodies at Amphoe Mae Chan, Chiang Rai. Cryptogamie Mycologie 3: 323–331. [Accession Number: WOS:000283517200006]

  32. Larget, B. & Simon, D.L. (1999) Markov chain Monte Carlo algorithms for the Bayesian analysis of phylogenetic trees. Molecular Biology and Evolution 16: 750–759.

  33. Liu, Y.J., Whelen, S. & Hall, B.D. (1999) Phylogenetic relationships among ascomycetes: Evidence from an RNA polymerse II subunit. Molecular Biology and Evolution 16: 1799–1808.

  34. Lu, Y.Z., Boonmee, S., Liu, J.K., Hyde, K.D., McKenzie, E.H., Eungwanichayapant, P.D. & Kang, J.C. (2018a) Multi-gene phylogenetic analyses reveals Neohelicosporium gen. nov. and five new species of helicosporous hyphomycetes from aquatic habitats. Mycological Progress 17: 631–646.

  35. Lu, Y.Z., Liu, J.K., Hyde, K.D., Jeewon, R., Kang, J.C., Fan, C., Boonmee, S., Bhat, D.J., Luo, Z.L., Lin, C.G. & Eungwanichayapant, P.D. (2018b) A taxonomic reassessment of Tubeufiales based on multi-locus phylogeny and morphology. Fungal Diversity 92: 131–344.

  36. Luo, Z.L., Maharachchikumbura, S.S.N., Liu, X.Y., Chen, L.J., Zhou, D.Q., Su, H.Y. & Hyde, K.D. (2015) Annulatascus saprophyticus sp. nov. and Pseudoannulatascus gen. nov. to accommodate Annulatascus biatriisporus (Annulatascales, Sordariomycetes) from Thailand. Phytotaxa 239: 174–182.

  37. Luo, Z.L., Bhat, D.J., Jeewon, R., Boonmee, S., Bao, D.F., Zhao, Y.C., Chai, H.M., Su, H.Y., Su, X.J. & Hyde, K.D. (2017) Molecular phylogeny and morphological characterization of asexual fungi (Tubeufiaceae) from Freshwater Habitats in Yunnan, China. Cryptogamie Mycologie 38: 27–53.

  38. Luo, Z.L., Hyde, K.D., Bhat, D.J., Jeewon, R., Maharachchikumbura, S.S.N., Bao, D.F., Li, W.L., Su, X.J., Yang, X.Y. & Su, H.Y. (2018) Morphological and molecular taxonomy of novel species Pleurotheciaceae from freshwater habitats in Yunnan, China. Mycological Progress: 1–20.

  39. Luo, Z.L., Hyde, K.D., Liu, J.K., Maharachchikumbura, S.S.N., Jeewon, R., Bao, D.F., Bhat, D.J., Lin, C.G., Li, W.L., Yang, J., Liu, N.G., Lu, Y.Z., Jayawardena, R.S., Li, J.F. & Su, H.Y. (2019) Freshwater Sordariomycetes. Fungal Diversity 99: 451–660.

  40. Maharachchikumbura, S.S.N., Hyde, K.D., Jones, E.B.G., McKenzie, E.H.C., Bhat, D.J., Dayarathne, M.C., Huang, S.K., Norphanphoun, C., Senanayake, I.C. & Perera, R.H. (2016) Families of Sordariomycetes. Fungal Diversity 79: 1–317.

  41. Matsushima, T. (1975) Icones Microfungorum a Matsushima lectorum. Matsushima lectorum. Nippon Printing Co, Osaka, 209 pp.

  42. Matsushima K, Matsushima T (1996) Fragmenta mycologica – II. In: Matsushima, T. (Ed.) Matsushima mycological memoirs, no. 9. Published by the author, Kobe, pp. 3140.

  43. Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Gateway Computing Environments Workshop 1: 1–8.

  44. Miller, M.A., Schwartz, T., Pickett, B.E., He, S., Klem, E.B., Scheuermann, R.H., Passarotti, M., Kaufman, S. & O’Leary, M.A. (2015) A RESTful API for access to phylogenetic tools via the CIPRES science gateway. Evolutionary Bioinformatics 11: 43–48.

  45. Monteiro, J.S., Gusmão, L.F.P. & Castañeda-Ruiz, R.F. (2016) Pleurothecium bicoloratum & Sporidesmiopsis pluriseptata spp. nov. from Brazil. Mycotaxon 131: 145–152.

  46. Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Tracer v1.7. Available from: (Accessed 8 May 2021)

  47. Page, R.D. (1996) TreeView: an application to display phylogenetic trees on personal computers. Computer Applications in the Biosciences 12: 357–358.

  48. Rannala, B. & Yang, Z. (1996) Probability distribution of molecular evolutionary trees: A new method of phylogenetic inference. Journal of Molecular Evolution 43: 304–311.

  49. Réblová, M. & Seifert, K.A. (2011) Discovery of the teleomorph of the hyphomycete, Sterigmatobotrys macrocarpa, and epitypification of the genus to holomorphic status. Studies in mycology 68: 193–202.

  50. Réblová, M., Seifert, K.A., Fournier, J. & Stepanek, V. (2012) Phylogenetic classification of Pleurothecium and Pleurotheciella gen. nov. and its dactylaria-like anamorph (Sordariomycetes) based on nuclear ribosomal and protein-coding genes. Mycologia 104: 1299–1314.

  51. Réblová, M., Seifert, K.A., Fournier, J. & Stepanek, V. (2016a) Newly recognised lineages of perithecial ascomycetes: the new orders Conioscyphales and Pleurotheciales. Persoonia 37: 57–81.

  52. Réblová, M., Miller, A.N., Rossman, A.Y., Seifert, K.A., Crous, P.W., Hawksworth, D.L., Abdel-Wahab, M.A., Cannon, P.F., Daranagama, D.A., De Beer, Z.W., Huang, S.K., Hyde, K.D., Jayawardena, R., Jaklitsch, W., Jones, E.B.G., Ju, Y.M., Judith, C., Maharachchikumbura, S.S.N., Pang, K.L., Petrini, L.E., Raja, H.A., Romero, A.I., Shearer, C., Senanayake, I.C., Voglmayr, H., Weir, B.S. & Wijayawarden, N.N. (2016b) Recommendations for competing sexual-asexually typified generic names in Sordariomycetes (except Diaporthales, Hypocreales, and Magnaporthales). IMA Fungus 7: 131–153.

  53. Réblová, M., Hernández-Restrepo, M., Fournier, J. & Nekvindová, J. (2020) New insights into the systematics of Bactrodesmium and its allies and introducing new genera, species and morphological patterns in the Pleurotheciales and Savoryellales (Sordariomycetes). Studies in Mycology, 95: 415–466.

  54. Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572–1574.

  55. Su, H.Y., Udayanga, D., Luo, Z.L., Manamgoda, D.S., Zhao, Y.C., Yang, J., Liu, X.Y., Mckenzie, E.H.C., Zhou, D.Q. & Hyde, K.D. (2015) Hyphomycetes from aquatic habitats in Southern China: Species of Curvularia (Pleosporaceae) and Phragmocephala (Melannomataceae). Phytotaxa 226: 201–216.

  56. Subramanian, C.V. & Bhat, D.J. (1989) Hyphomycetes from South India I. Some new taxa. Kavaka 15: 41–74.

  57. Tsui, C.K.M., Hyde, K.D. & Hodgkiss, I.J. (2001) Longitudinal and temporal distribution of freshwater ascomycetes and dematiaceous hyphomycetes on submerged wood in the Lam Tsuen River, Hong Kong. Journal of the North American Benthological Society 20: 533–549.

  58. Vilgalys, R. & Hester, M. (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. Journal of Bacteriology 172: 4238–4246.

  59. Wang, G.N., Yu, X.D., Dong, W., Jayarama Bhat, D.J., Boonmee, S., Zhang, D. & Zhang, H. (2019) Freshwater hyphomycetes in Eurotiomycetes: a new species of Minimelanolocus and a new collection of Thysanorea papuana (Herpotrichiellaceae). Mycological Progress 18: 511–522.

  60. Wei, M.J., Zhang, H., Dong, W., Boonmee, S. & Zhang, D. (2018) Introducing Dictyochaeta aquatica sp. nov. and two new species of Chloridium (Chaetosphaeriaceae , Sordariomycetes) from aquatic habitats. Phytotaxa 362: 187–199.

  61. White, T.J., Bruns, T., Lee, S. & Taylor, J. (1990) 38-Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis, M.A., Gelfand, D.H., Sninsky, J.J. & White, T.J. (Eds.) PCR protocols: a guide to methods and applications. Academic Press, San Diego, pp. 315–322

  62. Wijayawardene, N.N., Hyde, K.D., Rajeshkumar, K.C., Hawksworth, D.L., Madrid, H., Kirk, P.M., Braun, U., Singh, R.V., Crous, P.W., Kukwa, M., Lucking, R., Kurtzman, C.P., Yurkov, A., Haelewaters, D., Aptroot, A., Lumbsch, H.T., Timdal, E., Ertz, D., Etayo, J., Phillips, A.J.L., Groenewald, J.Z., Papizadeh, M., Selbmann, L., Dayarathne, M.C., Weerakoon, G., Jones, E.B.G., Suetrong, S., Tian, Q., Castaneda-Ruiz, R.F., Bahkali, A.H., Pang, K.L., Tanaka, K., Dai, D.Q., Sakayaroj, J., Hujslova, M., Lombard, L., Shenoy, B.D., Suija, A., Maharachchikumbura, S.S.N., Thambugala, K.M., Wanasinghe, D.N., Sharma, B.O., Gaikwad, S., Pandit, G., Zucconi, L., Onofri, S., Egidi, E., Raja, H.A., Kodsueb, R., Caceres, M.E.S., Perez-Ortega, S., Fiuza, P.O., Monteiro, J.S., Vasilyeva, L.N., Shivas, R.G., Prieto, M., Wedin, M., Olariaga, I., Lateef, A.A., Agrawal, Y., Fazeli, S.A.S., Amoozegar, M.A., Zhao, G.Z., Pfliegler, W.P., Sharma, G., Oset, M., Abdel-Wahab, M.A., Takamatsu, S., Bensch, K., de Silva, N.I., De Kesel, A., Karunarathna, A., Boonmee, S., Pfister, D.H., Lu, Y.Z., Luo, Z.L., Boonyuen, N., Daranagama, D.A., Senanayake, I.C., Jayasiri, S.C., Samarakoon, M.C., Zeng, X.Y., Doilom, M., Quijada, L., Rampadarath, S., Heredia, G., Dissanayake, A.J., Jayawardana, R.S., Perera, R.H., Tang, L.Z., Phukhamsakda, C., Hernandez-Restrepo, M., Ma, X.Y., Tibpromma, S., Gusmao, L.F.P., Weerahewa, D. & Karunarathna, S.C. (2017) Notes for genera: Ascomycota. Fungal Diversity 86: 1–594.

  63. Wijayawardene, N.N., Hyde, K.D., Al-Ani, L.K.T., Tedersoo, L., Haelewaters, D., Rajeshkumar, K.C., Zhao, R.L., Aptroot, A., Leontyev, D.V., Saxena, R.K., Tokarev, Y.S., Dai, D.Q., Letcher, P.M., Stephenson, S.L., Ertz, D., Lumbsch, H.T., Kukwa, M., Issi, I.V., Madrid, H., Phillips, A.J.L., Selbmann, L., Pfliegler, W.P., Horvath, E., Bensch, K., Kirk, P.M., Kolarikova, K., Raja, H.A., Radek, R., Papp, V., Dima, B., Ma, J., Malosso, E., Takamatsu, S., Rambold, G., Gannibal, P.B., Triebel, D., Gautam, A.K., Avasthi, S., Suetrong, S., Timdal, E., Fryar, S.C., Delgado, G., Reblova, M., Doilom, M., Dolatabadi, S., Pawlowska, J.Z., Humber, R.A., Kodsueb, R., Sanchez-Castro, I., Goto, B.T., Silva, D.K.A., de Souza, F.A., Oehl, F.R., da Silva, G.A., Silva, I.R., Blaszkowski, J., Jobim, K., Maia, L.C., Barbosa, F.R., Fiuza, P.O., Divakar, P.K., Shenoy, B.D., Castaneda-Ruiz, R.F., Somrithipol, S., Lateef, A.A., Karunarathna, S.C., Tibpromma, S., Mortimer, P.E., Wanasinghe, D.N., Phookamsak, R., Xu, J., Wang, Y., Tian, F., Alvarado, P., Li, D.W., Kusan, I., Matocec, N., Mesic, A., Tkalcec, Z., Maharachchikumbura, S.S.N., Papizadeh, M., Heredia, G., Wartchow, F., Bakhshi, M., Boehm, E., Youssef, N., Hustad, V.P., Lawrey, J.D., Santiago, A., Bezerra, J.D.P., Souza-Motta, C.M., Firmino, A.L., Tian, Q., Houbraken, J., Hongsanan, S., Tanaka, K., Dissanayake, A.J., Monteiro, J.S., Grossart, H.P., Suija, A., Weerakoon, G., Etayo, J., Tsurykau, A., Vazquez, V., Mungai, P., Damm, U., Li, Q.R., Zhang, H., Boonmee, S., Lu, Y.Z., Becerra, A.G., Kendrick, B., Brearley, F.Q., Motiejunaite, J., Sharma, B., Khare, R., Gaikwad, S., Wijesundara, D.S.A., Tang, L.Z., He, M.Q., Flakus, A., Rodriguez-Flakus, P., Zhurbenko, M.P., McKenzie, E.H.C., Stadler, M., Bhat, D.J., Liu, J.K., Raza, M., Jeewon, R., Nassonova, E.S., Prieto, M., Jayalal, R.G.U., Erdogdu, M., Yurkov, A., Schnittler, M., Shchepin, O.N., Novozhilov, Y.K., Silva, A.G.S., Gentekaki, E., Liu, P., Cavender, J.C., Kang, Y., Mohammad, S., Zhang, L.F., Xu, R.F., Li, Y.M., Dayarathne, M.C., Ekanayaka, A.H., Wen, T.C., Deng, C.Y., Pereira, O.L., Navathe, S., Hawksworth, D.L., Fan, X.L., Dissanayake, L.S., Kuhnert, E. & Thines, M. (2020) Outline of Fungi and fungi-like taxa. Mycosphere 11: 1060–1456.

  64. Wu, Y.M. & Zhang, T.Y. (2009) New species of Phialosporostilbe and Pleurothecium from soil. Mycotaxon 110: 1–4.

  65. Xia, J.W., Ma, Y.R., Li, Z. & Zhang, X.G. (2017) Acrodictys-like wood decay fungi from southern China, with two new families Acrodictyaceae and Junewangiaceae. Scientific Reports 7: 7888–7909.

  66. Xiao, Z.J., Li, X.X., Wang, H.D., Song, P.Y. & Tang, L. (2018) Rhexoacrodictys broussonetiae sp. nov. from Guizhou, China. Mycotaxon 133: 149–152.

  67. Yang, H,, Dong, W., Yu, X.D., Bhat, D.J., Boonmee, S. & Zhang, H. (2020) Four freshwater dematiaceous hyphomycetes in Sordariomycetes with two new species of Parafuscosporella. Phytotaxa 441: 19–34.

  68. Yu, X.D., Dong, W., Bhat, D.J., Boonmee, S., Zhang, D. & Zhang, H. (2018) Cylindroconidiis aquaticus gen. et sp nov., a new, lineage of aquatic hyphomycetes in Sclerococcaceae (Eurotiornycetes). Phytotaxa 372: 79–87.

  69. Zhang, H., Hyde, K.D., Mckenzie, E.H.C., Bahkali, A.H. & Zhou, D.Q. (2012) Sequence data reveals phylogenetic affinities of Acrocalymma aquatica sp. nov., Aquasubmersa mircensis gen. et sp. nov. and Clohesyomyces aquaticus (freshwater coelomycetes). Cryptogamie Mycologie 33: 333–346.

  70. Zhang, H., Hyde, K.D., Abdel-Wahab, M.A., Abdel-Aziz, F.A., Ariyawansa, H.A., Ko, T.W.K., Zhao, R.L., Alias, S.A., Bahkali, A.H. & Zhou, D.Q. (2013) A modern concept for Helicascus with a Pleurophomopsis-like asexual state. Sydowia 65: 147–166. [Accession Number: WOS:000323139100011]

  71. Zhang, H., Hyde, K.D., Zhao, Y.C., McKenzie, E.H.C. & Zhou, D.Q. (2014) Freshwater ascomycetes: Lophiostoma vaginatispora comb. nov. (Dothideomycetes, Pleosporales, Lophiostomaceae) based on morphological and molecular data. Phytotaxa 176: 184–191.

  72. Zhang, H., Dong, W., Hyde, K.D., Bahkali, A.H., Liu, J.K., Zhou, D.Q. & Zhang, D. (2016) Molecular data shows Didymella aptrootii is a new genus in Bambusicolaceae. Phytotaxa 247: 099–108.

  73. Zhang, H., Dong, W., Hyde, K.D., Maharachchikumbura, S.S.N., Hongsanan, S., Bhat, D.J., Al-Sadi, A.M. & Zhang, D. (2017) Towards a natural classification of Annulatascaceae-like taxa: introducing Atractosporales ord. nov. and six new families. Fungal Diversity 85: 75–110.