Abstract
Helminthosporium taiwanense sp. nov., isolated from a submerged decaying woody twig in a freshwater stream in Taiwan, is described and illustrated. While morphologically similar to Ellismarsporium due to its production of cylindrical conidia in short chains, multi-locus phylogenetic analyses based on a combined dataset of ITS, LSU, SSU, rpb2, and tef1-α sequences revealed it to be a distinct and undescribed species of Helminthosporium. The morphological boundaries and phylogenetic placement of the genus Ellismarsporium are discussed in light of this discovery.
References
- Alcorn, J.L. (1983) Generic concepts in Drechslera, Bipolaris and Exserohilum. Mycotaxon 17: 1–86. https://doi.org/10.5962/p.417485
- Alcorn, J.L. (1988) The taxonomy of “Helminthosporium” species. Annual Review of Phytopathology 26: 37–56. https://doi.org/10.1146/annurev.py.26.090188.000345
- Alves-Barbosa, M., Costa, P.M.O., Malosso, E. & Castaneda-Ruiz, R.F. (2017) Two new species of Dictyosporium and Helminthosporium from the Brazilian Atlantic Forest. Nova Hedwigia 105: 65–73. https://doi.org/10.1127/nova_hedwigia/2017/0401
- Bao, D.F., Luo, Z.L., Liu, J.K., Bhat, D.J., Sarunya, N., Li, W.L., Su, H.Y. & Hyde, K.D. (2018) Lignicolous freshwater fungi in China III: New species and record of Kirschsteiniothelia from northwestern Yunnan Province. Mycosphere 9 (4): 755–768. https://doi.org/10.5943/mycosphere/9/4/4
- Boonmee, S., Ko Ko, T.W., Chukeatirote, E., Hyde, K.D., Chen, H., Cai, L., McKenzie, E.H.C., Jones, E.B.G., Kodsueb, R. & Hassan, B.A. (2012) Two new Kirschsteiniothelia species with Dendryphiopsis anamorphs cluster in Kirschsteiniotheliaceae fam. nov. Mycologia 104: 698–714. https://doi.org/10.3852/11-089
- Boonmee, S., Wanasinghe, D.N., Calabon, M.S., Huanraluek, N., Chandrasiri, S.K.U., Jones, G.E.B., Rossi, W., Leonardi, M., Singh, S.K., Rana, S., Singh, P.N., Maurya, D.K., Lagashetti, A.C., Choudhary, D., Dai, Y.-C., Zhao, C.-L., Mu, Y.-H., Yuan, H.-S., He, S.-H., Phookamsak, R., Jiang, H.-B., Martín, M.P., Dueñas, M., Telleria, M.T., Kałucka, I.L., Jagodziński, A.M., Liimatainen, K., Pereira, D.S., Phillips, A.J.L., Suwannarach, N., Kumla, J., Khuna, S., Lumyong, S., Potter, T.B., Shivas, R.G., Sparks, A.H., Vaghefi, N., Abdel-Wahab, M.A., Abdel-Aziz, F.A., Li, G.-J., Lin, W.-F., Singh, U., Bhatt, R.P., Lee, H.B., Nguyen, T.T.T., Kirk, P.M., Dutta, A.K., Acharya, K., Sarma, V.V., Niranjan, M., Rajeshkumar, K.C., Ashtekar, N., Lad, S., Wijayawardene, N.N., Bhat, D.J., Xu, R.-J., Wijesinghe, S.N., Shen, H.-W., Luo, Z.-L., Zhang, J.-Y., Sysouphanthong, P., Thongklang, N., Bao, D.-F., Aluthmuhandiram, J.V.S., Abdollahzadeh, J., Javadi, A., Dovana, F., Usman, M., Khalid, A.N., Dissanayake, A.J., Telagathoti, A., Probst, M., Peintner, U., Garrido-Benavent, I., Bóna, L., Merényi, Z., Boros, L., Zoltán, B., Stielow, J.B., Jiang, N., Tian, C.-M., Shams, E., Dehghanizadeh, F., Pordel, A., Javan-Nikkhah, M., Denchev, T.T., Denchev, C.M., Kemler, M., Begerow, D., Deng, C.-Y., Harrower, E., Bozorov, T., Kholmuradova, T., Gafforov, Y., Abdurazakov, A., Xu, J.-C., Mortimer, P.E., Ren, G.-C., Jeewon, R., Maharachchikumbura, S.S.N., Phukhamsakda, C., Mapook, A.& Hyde, K.D. (2021) Fungal diversity notes 1387–1511: taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 111 (1): 1–335. https://doi.org/10.1007/s13225-021-00489-3
- Castañeda-Ruiz, R.F., Li, D.W., Zhang, X.G., Kendrick, B., Ramos-García, B., Pérez-Martínez, S. & Sosa, D. (2017) Ellismarsporium gen. nov. and Stanhughesiella gen. nov. to accommodate atypical Helminthosporium and Corynesporella species. Mycotaxon 132 (4): 759–766. https://doi.org/10.5248/132.759
- Chen, Y.P., Tian, W.H., Guo, Y.B., Madrid, H. & Maharachchikumbura, S.S.N. (2022) Synhelminthosporium gen. et sp. nov. and two new species of Helminthosporium (Massarinaceae, Pleosporales) from Sichuan Province, China. Journal of Fungi 8 (7): 712. https://doi.org/10.3390/jof8070712
- Crous, P.W., Wingfield, M.J., Schumacher, R.K., Summerell, B.A., Giraldo, A., Gené, J., Guarro, J., Wanasinghe, D.N., Hyde, K.D., Camporesi, E., Garethjones, E.B., Thambugala, K.M., Malysheva, E.F., Malysheva, V.F., Acharya, K., Álvarez, J., Alvarado, P., Assefa, A., Barnes, C.W., Bartlett, J.S., Blanchette, R.A., Burgess, T.I., Carlavilla, J.R., Coetzee, M.P.A., Damm, U., Decock, C.A., Denbreeÿen, A., Devries, B., Dutta, A.K., Holdom, D.G., Rooney-Latham, S., Manjón, J.L., Marincowitz, S., Mirabolfathy, M., Moreno, G., Nakashima, C., Papizadeh, M., Shahzadehfazeli, S.A., Amoozegar, M.A., Romberg, M.K., Shivas, R.G., Stalpers, J.A., Stielow, B., Stukely, M.J.C., Swart, W.J., Tan, Y.P., Vanderbank, M., Wood, A.R., Zhang, Y. & Groenewald, J.Z. (2014) Fungal Planet description sheets: 281–319. Persoonia 33: 212–319. https://doi.org/10.3767/003158514x685680
- Crous, P.W., Wingfield, M.J., Richardson, D.M., Leroux, J.J., Strasberg, D., Edwards, J., Roets, F., Hubka, V., Taylor, P.W.J., Heykoop, M., Martín, M.P., Moreno, G., Sutton, D.A., Wiederhold, N.P., Barnes, C.W., Carlavilla, J.R., Gené, J., Giraldo, A., Guarnaccia, V., Guarro, J., Hernández-Restrepo, M., Kolařík, M., Manjón, J.L., Pascoe, I.G., Popov, E.S., Sandoval-Denis, M., Woudenberg, J.H.C., Acharya, K., Alexandrova, A.V., Alvarado, P., Barbosa, R.N., Baseia, I.G., Blanchette, R.A., Boekhout, T., Burgess, T.I., Cano-Lira, J.F., Čmoková, A., Dimitrov, R.A., Dyakov, M.Yu., Dueñas, M., Dutta, A.K., Esteve-Raventós, F., Fedosova, A.G., Fournier, J., Gamboa, P., Gouliamova, D.E., Grebenc, T., Groenewald, M., Hanse, B., Hardy, G.E.ST.J., Held, B.W., Jurjević, Ž, Kaewgrajang, T., Latha, K.P.D., Lombard, L., Luangsa-ard, J.J., Lysková, P., Mallátová, N., Manimohan, P., Miller, A.N., Mirabolfathy, M., Morozova, O.V., Obodai, M., Oliveira, N.T., Ordóñez, M.E., Otto, E.C., Paloi, S., Peterson, S.W., Phosri, C., Roux, J., Salazar, W.A., Sánchez, A., Sarria, G.A., Shin, H.-D., Silva, B.D.B., Silva, G.A., Smith, M.TH., Souza-Motta, C.M., Stchigel, A.M., Stoilova-Disheva, M.M., Sulzbacher, M.A., Telleria, M.T., Toapanta, C., Traba, J.M., Valenzuela-Lopez, N., Watling, R. & Groenewald, J.Z. (2016) Fungal Planet description sheets: 400–468. Persoonia 36 (1): 316–458. https://doi.org/10.3767/003158516x692185
- Crous, P., Wingfield, M., Burgess, T., Hardy, G., Gene, J., Guarro, J., Baseia, I., Garcia, D., Gusmao, L., Souza-Motta, C., Thangavel, R., Adamcik, S., Barili, A., Barnes, C., Bezerra, J., Bordallo, J., Cano-Lira, J., de Oliveira, R., Ercole, E., Hubka, V., Iturrieta-Gonzalez, I., Kubatova, A., Martin, M., Moreau, P.A., Morte, A., Ordonez, M., Rodriguez, A., Stchigel, A., Vizzini, A., Abdollahzadeh, J., Abreu, V., Adamcikova, K., Albuquerque, G., Alexandrova, A., Alvarez Duarte, E., Armstrong-Cho, C., Banniza, S., Barbosa, R., Bellanger, J.M., Bezerra, J., Cabral, T., Cabon, M., Caicedo, E., Cantillo, T., Carnegie, A., Carmo, L., Castaneda-Ruiz, R., Clement, C., Cmokova, A., Conceicao, L., Cruz, R., Damm, U., da Silva, B., da Silva, G., da Silva, R., Santiago, Ad.A., de Oliveira, L., de Souza, C., Deniel, F., Dima, B., Dong, G., Edwards, J., Felix, C., Fournier, J., Gibertoni, T., Hosaka, K., Iturriaga, T., Jadan, M., Jany, J.-L., Jurjevic, Z., Kolarik, M., Kusan, I., Landell, M., Leite Cordeiro, T., Lima, D., Loizides, M., Luo, S., Machado, A., Madrid, H., Magalhaes, O., Marinho, P., Matocec, N., Mesic, A., Miller, A., Morozova, O., Neves, R., Nonaka, K., Novakova, A., Oberlies, N., Oliveira-Filho, J., Oliveira, T., Papp, V., Pereira, O., Perrone, G., Peterson, S., Pham, T. & Raja, H. (2018) Fungal Planet description sheets: 716–784. Persoonia 40 (1): 239–392. https://doi.org/10.3767/persoonia.2018.40.10
- Crous, P.W., Carnegie, A.J., Wingfield, M.J., Sharma, R., Mughini, G., Noordeloos, M.E., Santini, A., Shouche, Y.S., Bezerra, J.D.P., Dima, B., Guarnaccia, V., Imrefi, I., Jurjević, Ž., Knapp, D.G., Kovács, G.M., Magistà, D., Perrone, G., Rämä, T., Rebriev, Y.A., Shivas, R.G., Singh, S.M., Souza-Motta, C.M., Thangavel, R., Adhapure, N.N., Alexandrova, A.V., Alfenas, A.C., Alfenas, R.F., Alvarado, P., Alves, A.L., Andrade, D.A., Andrade, J.P., Barbosa, R.N., Barili, A., Barnes, C.W., Baseia, I.G., Bellanger, J.-M., Berlanas, C., Bessette, A.E., Bessette, A.R., Biketova, A.Yu., Bomfim, F.S., Brandrud, T.E., Bransgrove, K., Brito, A.C.Q., Cano-Lira, J.F., Cantillo, T., Cavalcanti, A.D., Cheewangkoon, R., Chikowski, R.S., Conforto, C., Cordeiro, T.R.L., Craine, J.D., Cruz, R., Damm, U., de Oliveira, R.J.V., de Souza, J.T., de Souza, H.G., Dearnaley, J.D.W., Dimitrov, R.A., Dovana, F., Erhard, A., Esteve-Raventós, F., Félix, C.R., Ferisin, G., Fernandes, R.A., Ferreira, R.J., Ferro, L.O., Figueiredo, C.N., Frank, J.L., Freire, K.T.L.S., García, D., Gené, J., Gęsiorska, A., Gibertoni, T.B., Gondra, R.A.G., Gouliamova, D.E., Gramaje, D., Guard, F., Gusmão, L.F.P., Haitook, S., Hirooka, Y., Houbraken, J., Hubka, V., Inamdar, A., Iturriaga, T., Iturrieta-González, I., Jadan, M., Jiang, N., Justo, A., Kachalkin, A.V., Kapitonov, V.I., Karadelev, M., Karakehian, J., Kasuya, T., Kautmanová, I., Kruse, J., Kušan, I., Kuznetsova, T.A., Landell, M.F., Larsson, K.-H., Lee, H.B., Lima, D.X., Lira, C.R.S., Machado, A.R., Madrid, H., Magalhães, O.M.C., Majerova, H., Malysheva, E.F., Mapperson, R.R., Marbach, P.A.S., Martín, M.P., Martín-Sanz, A., Matočec, N., McTaggart, A.R., Mello, J.F., Melo, R.F.R., Mešič, A., Michereff, S.J., Miller, A.N., Minoshima, A., Molinero-Ruiz, L., Morozova, O.V., Mosoh, D., Nabe, M., Naik, R., Nara, K., Nascimento, S.S., Neves, R.P., Olariaga, I., Oliveira, R.L., Oliveira, T.G.L., Ono, T., Ordoñez, M.E., de M. Ottoni, A., Paiva, L.M., Pancorbo, F., Pant, B., Pawłowska, J., Peterson, S.W., Raudabaugh, D.B., Rodríguez-Andrade, E., Rubio, E., Rusevska, K., Santiago, A.L.C.M.A., Santos, A.C.S., Santos, C., Sazanova, N.A., Shah, S., Sharma, J., Silva, B.D.B., Siquier, J.L., Sonawane, M.S., Stchigel, A.M., Svetasheva, T., Tamakeaw, N., Telleria, M.T., Tiago, P.V., Tian, C.M., Tkalčec, Z., Tomashevskaya, M.A., Truong, H.H., Vecherskii, M.V., Visagie, C.M., Vizzini, A., Yilmaz, N., Zmitrovich, I.V., Zvyagina, E.A., Boekhout, T., Kehlet, T., Læssøe, T. & Groenewald, J.Z. (2019) Fungal Planet description sheets: 868–950. Persoonia 42: 291–473. https://doi.org/10.3767/persoonia.2019.42.11
- Dokhanchi, H., Babai-Ahari, A., Arzanlou, M. & Abed-Ashtiani, F. (2022) Characterization of Pyrenophora, Bipolaris, Curvularia and Exserohilum species associated with leaf spot diseases on barley in East Azerbaijan province, using morphological and molecular data. Journal of Applied Research in Plant Protection 11 (1): 1–18.
- Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32: 1792–1797. https://doi.org/10.1093/nar/gkh340
- Ellis, M.B. (1971) Dematiaceous Hyphomycetes. Commonwealth Mycological Institute, Kew, Surrey, England, 608 pp. https://doi.org/10.1079/9780851986180.0000
- Goh, T.K. (1998) Generic distinction in the Helminthosporium-complex based on restriction analysis of the nuclear ribosomal RNA gene. Fungal Diversity 1: 109–113.
- Goh, T.K. (1999) Single-spore isolation using a hand-made glass needle. Fungal Diversity 2: 47–63.
- Gomes de Farias, A.R., Afshari, N., Silva, V., Louangphan, J., Karimi, O. & Boonmee, S. (2024) Three novel species and new records of Kirschsteiniothelia (Kirschsteiniotheliales) from northern Thailand. MycoKeys 101: 347–370. https://doi.org/10.3897/mycokeys.101.115286
- Gornostaĭ, V.I. (1972) New and uncommon fungal species in the maritime area. Mikologiya i Fitopatologiya 6: 154–156.
- Hu, Y.-F., Liu, J.-W., Xu, Z.-H., Castaneda-Ruiz, R.F., Zhang, K. & Ma, J. (2023) Morphology and multigene phylogeny revealed three new species of Helminthosporium (Massarinaceae, Pleosporales) from China. Journal of Fungi 9 (2): 280. https://doi.org/10.3390/jof9020280
- Hyde, K.D., Jones, E.B.G., Liu, J.-K., Ariyawansa, H., Boehm, E., Boonmee, S., Braun, U., Chomnunti, P., Crous, P.W., Dai, D.-Q., Diederich, P., Dissanayake, A., Doilom, M., Doveri, F., Hongsanan, S., Jayawardena, R., Lawrey, J.D., Li, Y.-M., Liu, Y.-X., Lücking, R., Monkai, J., Muggia, L., Nelsen, M.P., Pang, K.-L., Phookamsak, R., Senanayake, I.C., Shearer, C.A., Suetrong, S., Tanaka, K., Thambugala, K.M., Wijayawardene, N.N., Wikee, S., Wu, H.-X., Zhang, Y., Aguirre-Hudson, B., Alias, S.A., Aptroot, A., Bahkali, A.H., Bezerra, J.L., Bhat, D.J., Camporesi, E., Chukeatirote, E., Gueidan, C., Hawksworth, D.L., Hirayama, K., De Hoog, S., Kang, J.-C., Knudsen, K., Li, W.-J., Li, X.-H., Liu, Z.-Y., Mapook, A., McKenzie, E.H.C., Miller, A.N., Mortimer, P.E., Phillips, A.J.L., Raja, H.A., Scheuer, C., Schumm, F., Taylor, J.E., Tian, Q., Tibpromma, S., Wanasinghe, D.N., Wang, Y., Xu, J.-C., Yacharoen, S., Yan, J.-Y. & Zhang, M. (2013) Families of Dothideomycetes. Fungal Diversity 63: 1–313. https://doi.org/10.1007/s13225-013-0263-4
- Hyde, K.D., Norphanphoun, C., Abreu, V.P., Bazzicalupo, A., Chethana, K.W.T., Clericuzio, M., Dayarathne, M.C., Dissanayake, A.J., Ekanayaka, A.H., He, M.-Q., Hongsanan, S., Huang, S.-K., Jayasiri, S.C., Jayawardena, R.S., Karunarathna, A., Konta, S., Kušan, I., Lee, H., Li, J., Lin, C.-G., Liu, N.-G., Lu, Y.-Z., Luo, Z.-L., Manawasinghe, I.S., Mapook, A., Perera, R.H., Phookamsak, R., Phukhamsakda, C., Siedlecki, I., Soares, A.M., Tennakoon, D.S., Tian, Q., Tibpromma, S., Wanasinghe, D.N., Xiao, Y.-P., Yang, J., Zeng, X.-Y., Abdel-Aziz, F.A., Li, W.-J., Senanayake, I.C., Shang, Q.-J., Daranagama, D.A., de Silva, N.I., Thambugala, K.M., Abdel-Wahab, M.A., Bahkali, A.H., Berbee, M.L., Boonmee, S., Bhat, D.J., Bulgakov, T.S., Buyck, B., Camporesi, E., Castañeda-Ruiz, R.F., Chomnunti, P., Doilom, M., Dovana, F., Gibertoni, T.B., Jadan, M., Jeewon, R., Jones, E.B.G., Kang, J.-C., Karunarathna, S.C., Lim, Y.W., Liu, J.-K., Liu, Z.-Y., Plautz, H.L.Jr., Lumyong, S., Maharachchikumbura, S.S.N., Matočec, N., McKenzie, E.H.C., Mešić, A., Miller, D., Pawłowska, J., Pereira, O.L., Promputtha, I., Romero, A.I., Ryvarden, L., Su, H.-Y., Suetrong, S., Tkalčec, Z., Vizzini, A., Wen, T.-C., Wisitrassameewong, K., Wrzosek, M., Xu, J.-C., Zhao, Q., Zhao, R.-L. & Mortimer, P.E. (2017) Fungal diversity notes 603–708: taxonomic and phylogenetic notes on genera and species. Fungal Diversity 87: 1–235. https://doi.org/10.1007/s13225-017-0391-3
- Hyde, K.D., Norphanphoun, C., Ma, J., Yang, H.D., Zhang, J.Y., Du, T.Y., Gao, Y., Gomes de Farias, A.R., He, S.C., He, Y.K., Li, C.J.Y., Liu, X.F., Lu, L., Su, H.L., Tang, X., Tian, X.G., Wang, S.Y., Wei, D.P., Xu, R.F., Xu, R.J., Yang, Q., Yang, Y.Y., Zhang, F., Zhang, Q., Bahkali, A.H., Boonmee, S., Chethana, K.W.T., Jayawardena, R.S., Lu, Y.Z., Karunarathna, S.C., Tibpromma, S., Wang, Y. & Zhao, Q. (2023) Mycosphere notes 387–412: novel species of fungal taxa from around the world. Mycosphere 14 (1): 663–744. https://doi.org/10.5943/mycosphere/14/1/8
- Hyde, K.D., Wijesinghe, S.N., Afshari, N., Aumentado, H.D., Bhunjun, C.S., Boonmee, S., Camporesi, E., Chethana, K.W.T., Doilom, M., Dong, W., Du, T.Y., Farias, A.R.G., Gao, Y., Jayawardena, R.S., Karimi, O., Karunarathna, S.C., Kularathnage, N.D., Lestari, A.S., Li, C.J.Y., Li, Y.X., Liao, C.F., Liu, X.F., Lu, L., Lu, Y.Z., Luo, Z.L., Ma, J., Mamarabadi, M., Manawasinghe, I.S., Mapook, A., Mi, L.X., Niranjan, M., Senanayake, I.C., Shen, H.W., Su, H.L., Tibpromma, S., Xu, R.J., Yan, J.Y., Yang, Y.H., Yang, Y.Y., Yu, F.Q., Kang, J.C. & Zhang, J.Y. (2024) Mycosphere Notes 469–520. Mycosphere 15 (1): 1294–1454. https://doi.org/10.5943/mycosphere/15/1/11
- Index Fungorum (2025) Available from: http://www.indexfungorum.org/names/Names.asp (accessed 16 September 2025)
- Ilyukhin, E. & Markovskaja, S. (2023) Helminthosporium paraoligosporum sp. nov., a new ascomycetous fungus from Ontario, Canada. Phytotaxa 607 (2): 151–159. https://doi.org/10.11646/phytotaxa.607.2.4
- Jayawardena, R.S., Hyde, K.D., Wang, S., Sun, Y.-R., Suwannarach, N., Sysouphanthong, P., Abdel-Wahab, M.A., Abdel-Aziz, F.A., Abeywickrama, P.D., Abreu, V.P., Armand, A., Aptroot, A., Bao, D.-F., Begerow, D., Bellanger, J.-M., Bezerra, J.D.P., Bundhun, D., Calabon, M.S., Cao, T., Cantillo, T., Carvalho, J.L.V.R., Chaiwan, N., Chen, C.-C., Courtecuisse, R., Cui, B.-K., Damm, U., Denchev, C.M., Denchev, T.T., Deng, C.Y., Devadatha, B., de Silva, N.I., dos Santos, L.A., Dubey, N.K., Dumez, S., Ferdinandez, H.S., Firmino, A.L., Gafforov, Y., Gajanayake, A.J., Gomdola, D., Gunaseelan, S., He, S., Htet, Z.H., Kaliyaperumal, M., Kemler, M., Kezo, K., Kularathnage, N.D., Leonardi, M., Li, J.-P., Liao, C., Liu, S., Loizides, M., Luangharn, T., Ma, J., Madrid, H., Mahadevakumar, S., Maharachchikumbura, S.S.N., Manamgoda, D.S., Martín, M.P., Mekala, N., Moreau, P.-A., Mu, Y.-H., Pahoua, P., Pem, D., Pereira, O.L., Phonrob, W., Phukhamsakda, C., Raza, M., Ren, G.-C., Rinaldi, A.C., Rossi, W., Samarakoon, B.C., Samarakoon, M.C., Sarma, V.V., Senanayake, I.C., Singh, A., Souza, M.F., Souza-Motta, C.M., Spielmann, A.A., Su, W., Tang, X., Tian, X., Thambugala, K.M., Thongklang, N., Tennakoon, D.S., Wannathes, N., Wei, D., Welti, S., Wijesinghe, S.N., Yang, H., Yang, Y., Yuan, H.-S., Zhang, H., Zhang, J., Balasuriya, A., Bhunjun, C.S., Bulgakov, T.S., Cai, L., Camporesi, E., Chomnunti, P., Deepika, Y.S., Doilom, M., Duan, W.-J., Han, S.-L., Huanraluek, N., Jones, E.B.G., Lakshmidevi, N., Li, Y., Lumyong, S., Luo, Z.-L., Khuna, S., Kumla, J., Manawasinghe, I.S., Mapook, A., Punyaboon, W., Tibpromma, S., Lu, Y.-Z., Yan, J.-Y. & Wang, Y. (2022) Fungal diversity notes 1512–1610: taxonomic and phylogenetic contributions on genera and species of fungal taxa. Fungal Diversity 117 (1): 1–272. https://doi.org/10.1007/s13225-022-00513-0
- Jiang, S.H., Hawksworth, D.L., Lücking, R. & Wei, J.C. (2020) A new genus and species of foliicolous lichen in a new family of Strigulales (Ascomycota: Dothideomycetes) reveals remarkable class-level homoplasy. IMA Fungus 11 (1): 1. https://doi.org/10.1186/s43008-019-0026-2
- Konta, S., Hyde, K.D., Karunarathna, S.C., Mapook, A., Senwanna, C., Dauner, L.A., Nanayakkara, C.M., Xu, J., Tibpromma, S. & Lumyong, S. (2021) Multi-gene phylogeny and morphology reveal Haplohelminthosporium gen. nov. and Helminthosporiella gen. nov. associated with palms in Thailand and a checklist for Helminthosporium reported worldwide. Life 11 (5): 454. https://doi.org/10.3390/life11050454
- Konta, S., Tibpromma, S., Karunarathna, S.C., Samarakoon, M.C., Steven, L.S., Mapook, A., Boonmee, S., Senwanna, C., Balasuriya, A., Eungwanichayapant, P.D. & Hyde, K.D. (2023) Morphology and multigene phylogeny reveal ten novel taxa in Ascomycota from terrestrial palm substrates (Arecaceae) in Thailand. Mycosphere 14 (1): 107–152. https://doi.org/10.5943/mycosphere/14/1/2
- Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution 33: 1870–1874. https://doi.org/10.1093/molbev/msw054
- Kumar, P., Akhtar, J., Kiran, R., Meena, B.R., Sadhana & Chalam, V.C. (2023) Phylogenetic analysis and species level identification of seed-borne Bipolaris, Curvularia and Exserohilum fungal pathogens. Indian Phytopathology 76 (1): 115–121. https://doi.org/10.1007/s42360-022-00585-7
- Liao, M.G., Luo, X.X., Xia, J.W., Hu, Y.F., Zhang, X.G., Zhang, L.H., Zhang, X.P., Xu, Z.H. & Ma, J. (2025) Six Novel species of Distoseptispora (Distoseptisporaceae, Distoseptisporales) and Helminthosporium (Massarinaceae, Pleosporales) isolated from terrestrial habitats in Southern China. Journal of Fungi 11: 494. https://doi.org/10.3390/jof11070494
- Link, H.F. (1809) Observationes in ordines plantarum naturales. Dissertatio Ima. Gesellschaft Naturforschender Freunde zu Berlin, Magazin 3 (1): 3–42. [https://BiotaNZ.landcareresearch.co.nz/references/1cb0f202-36b9-11d5-9548-00d0592d548c]
- Liu, Y.J., Whelen, S. & Hall, B.D. (1999) Phylogenetic relationships among ascomycetes: evidence from an RNA polymerase II subunit. Molecular Biology and Evolution 16: 1799–1808. https://doi.org/10.1093/oxfordjournals.molbev.a026092
- Liu, J., Hu, Y., Luo, X., Castañeda-Ruíz, R.F. & Ma, J. (2022) Three novel species of Helminthosporium (Massarinaceae, Pleosporales) from China. MycoKeys 94: 73–89. https://doi.org/10.3897/mycokeys.94.95888
- Louangphan, J., Perera, R.H. & Gomes De Farias, A.R. (2024) A new addition to Kirschsteiniotheliaceae: Kirschsteiniothelia chiangmaiensis sp. nov. from Northern Thailand. Phytotaxa 634 (1): 49–62. https://doi.org/10.11646/phytotaxa.634.1.4
- Lu, L., Karunarathna, S.C., Rajeshkumar, K.C., Elgorban, A.M., Jayawardena, R.S., Hongsanan, S., Suwannarach, N., Kumla, J., Xiong, Y.-R., Hyde, K.D., Han, M.Y., Zheng, D.G., Li, Q., Dai, D.Q. & Tibpromma, S. (2025) Unveiling fungal diversity associated with coffee trees in China using a polyphasic approach and a global review of coffee saprobic fungi. IMA Fungus 16: e144874. https://doi.org/10.3897/imafungus.16.144874.suppl1
- Luo, X.X., Liao, M.G., Hu, Y.F., Zhang, X.G., Xu, Z.H. & Ma, J. (2025) Identification of three novel species and one new record of Kirschsteiniothelia (Kirschsteiniotheliaceae, Kirschsteiniotheliales) from Jiangxi, China. MycoKeys 112: 277–306. https://doi.org/10.3897/mycokeys.112.142028
- Luttrell, E.S. (1963) Taxonomic criteria in Helminthosporium. Mycologia 55: 643–674. https://doi.org/10.2307/3756441
- Luttrell, E.S. (1964) Systematics of Helminthosporium and related genera. Mycologia 56 (1): 119–132. https://doi.org/10.1080/00275514.1964.12018090
- Matsushima, T. (1971) Microfungi of the Solomon Islands and Papua-New Guinea. Matsushima Fungus Collection, Kobe, Japan, 78 pp.
- Matsushima, T. (1993) Matsushima Mycological Memoirs No. 7. Matsushima Fungus Collection, Kobe, Japan, 141 pp.
- Nelsen, M.P., Lücking, R., Grube, M., Mbatchou, J.S., Muggia, L., Rivas-Plata, E. & Lumbsch, H.T. (2009) Unravelling the phylogenetic relationships of lichenised fungi in Dothideomyceta. Studies in Mycology 64: 135–144. https://doi.org/10.3114/sim.2009.64.07
- Olivier, C., Berbee, M.L., Shoemaker, R.A. & Loria, R. (2000) Molecular phylogenetic support from ribosomal DNA sequences for origin of Helminthosporium from Leptosphaeria-like loculoascomycete ancestors. Mycologia 92 (4): 736–746. https://doi.org/10.1080/00275514.2000.12061213
- Quaedvlieg, W., Verkley, G.J.M., Shin, H.D., Barreto, R.W., Alfenas, A.C., Swart, W.J., Groenewald, J.Z. & Crous, P.W. (2013) Sizing up septoria. Studies in Mycology 75: 307–390. https://doi.org/10.3114/sim0017
- Rehner, S.A. & Buckley, E. (2005) Beauveria phylogeny inferred from nuclear ITS and EF1-α sequences: evidence for cryptic diversification and links to Cordyceps teleomorphs. Mycologia 97: 84–98. https://doi.org/10.1080/15572536.2006.11832842
- Rodríguez-Andrade, E., Stchigel, A.M., Guarro, J. & Cano-Lira, J.F. (2020) Fungal diversity of deteriorated sparkling wine and cork stoppers in Catalonia, Spain. Microorganisms 8 (1): 1–12. https://doi.org/10.3390/microorganisms8010012
- Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
- Saitou, N. & Nei, M. (1987) The neighbor-joining method: A new method for reconstructing phylogenetic trees. Molecular Biology and Evolution 4: 406–425. https://doi.org/10.1093/oxfordjournals.molbev.a040454
- Sambrook, J. & Russell, D.W. (2001) Molecular cloning: a laboratory manual. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York, 2344 pp.
- Schoch, C.L., Shoemaker, R.A., Seifert, K.A., Hambleton, S., Spatafora, J.W. & Crous, P.W. (2006) A multigene phylogeny of the Dothideomycetes using four nuclear loci. Mycologia 98 (6): 1041–1052. https://doi.org/10.3852/mycologia.98.6.1041
- Schoch, C.L., Crous, P.W., Groenewald, J.Z., Boehm, E.W., Burgess, T.I., de Gruyter, J., de Hoog, G.S., Dixon, L.J., Grube, M., Gueidan, C., Harada, Y., Hatakeyama, S., Hirayama, K., Hosoya, T., Huhndorf, S.M., Hyde, K.D., Jones, E.B.G., Kohlmeyer, J., Kruys, A., Li, Y.M., Lucking, R., Lumbsch, H.T., Marvanova, L., Mbatchou, J.S., McVay, A.H., Miller, A.N., Mugambi, G.K., Muggia, L., Nelsen, M.P., Nelson, P., Owensby, C.A., Phillips, A.J., Phongpaichit, S., Pointing, S.B., Pujade-Renaud, V., Raja, H.A., Plata, E.R., Robbertse, B., Ruibal, C., Sakayaroj, J., Sano, T., Selbmann, L., Shearer, C.A., Shirouzu, T., Slippers, B., Suetrong, S., Tanaka, K., Volkmann-Kohlmeyer, B., Wingfield, M.J., Wood, A.R., Woudenberg, J.H., Yonezawa, H., Zhang, Y. & Spatafora, J.W. (2009) A class-wide phylogenetic assessment of Dothideomycetes. Studies in Mycology 64: 1–15. https://doi.org/10.3114/sim.2009.64.01
- Shoemaker, R.A. & Hambleton, S. (2001) " Helminthosporium" asterinum, Polydesmus elegans, Imimyces, and allies. Canadian Journal of Botany 79 (5): 592–599. https://doi.org/10.1139/b01-038
- Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30: 1312–1313. https://doi.org/10.1093/bioinformatics/btu033
- Stenroos, S., Laukka, T., Huhtinen, S., Döbbeler, P., Myllys, L., Syrjänen, K. & Hyvönen, J. (2010) Multiple origins of symbioses between ascomycetes and bryophytes suggested by a five‐gene phylogeny. Cladistics 26 (3): 281–300. https://doi.org/10.1111/j.1096-0031.2009.00284.x
- Su, H.Y., Hyde, K.D., Maharachchikumbura, S.S., Ariyawansa, H.A., Luo, Z.L., Promputtha, I., Tian, Q., Lin, C.G., Shang, Q.J., Zhao, Y.C., Chai, H.M., Liu, X.Y., Bahkali, A.H., Bhat, J.D., McKenzie, E.H.C. & Zhou, D.Q. (2016) The families Distoseptisporaceae fam. nov., Kirschsteiniotheliaceae, Sporormiaceae and Torulaceae, with new species from freshwater in Yunnan Province, China. Fungal Diversity 80 (1): 375–409. https://doi.org/10.1007/s13225-016-0362-0
- Sun, Y.R., Hyde, K.D., Liu, N.G., Jayawardena, R.S., Wijayawardene, N.N., Ma, J., Zhang, Q., Al-Otibi, F. & Wang, Y. (2025) Micro-fungi in southern China and northern Thailand: Emphasis on medicinal plants. Fungal Diversity 131: 99–299. https://doi.org/10.1007/s13225-024-00549-4
- Tamura, K. & Nei, M. (1993) Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Molecular Biology and Evolution 10: 512–526. https://doi.org/10.1093/oxfordjournals.molbev.a040023
- Tanaka, K., Hirayama, K., Yonezawa, H., Sato, G., Toriyabe, A., Kudo, H., Hashimoto, A., Matsumura, M., Harada, Y., Kurihara, Y., Shirouzu, T. & Hosoya, T. (2015) Revision of the massarineae (Pleosporales, Dothideomycetes). Studies in Mycology 82: 75–136. https://doi.org/10.1016/j.simyco.2015.10.002
- Tang, X., Jeewon, R., Jayawardena, R.S., Gomdola, D., Lu, Y.Z., Xu, R.J., Alrefaei, A.F., Alotibi, F., Hyde, K.D. & Kang, J.C. (2024) Additions to the genus Kirschsteiniothelia (Dothideomycetes); three novel species and a new host record, based on morphology and phylogeny. MycoKeys 110: 35–66. https://doi.org/10.3897/mycokeys.110.133450
- Tangthirasunun, N., Silar, P., Bhat, D.J., Chukeatirote, E., Wijayawardene, D.N.N., Maharachchikumbura, S.S.N. & Hyde, K.D. (2014) Morphology and phylogeny of Pseudorobillarda eucalypti sp. nov., from Thailand. Phytotaxa 176: 251–259. https://doi.org/10.11646/phytotaxa.176.1.24
- Thambugala, K.M., Chunfang, Y., Camporesi, E., Bahkali, A.H., Liu, Z.Y. & Hyde, K.D. (2015) Pseudodidymosphaeria gen. nov. in Massarinaceae. Phytotaxa 231 (3): 271–282. https://doi.org/10.11646/phytotaxa.231.3.5
- Thambugala, K.M., Wanasinghe, D.N., Phillips, A.J.L., Camporesi, E., Bulgakov, T.S., Phukhamsakda, C., Dissanayake, A., Tennakoon, D.S., Tibpromma, S., Chen, Y.Y., Liu, Z.Y. & Hyde, K.D. (2017) Mycosphere notes 1–50: grass (Poaceae) inhabiting Dothideomycetes. Mycosphere 8: 697–796. https://doi.org/10.5943/mycosphere/8/4/13
- Tian, Q., Li, W.J., Hyde, K.D., Camporesi, E., Bhat, D.J., Chomnunti, P. & Xu, J.C. (2018) Molecular taxonomy of five species of microfungi on Alnus spp. from Italy. Mycological Progress 17: 255–274. https://doi.org/10.1007/s11557-017-1336-7
- Tian, W.H., Jin, Y., Liao, Y.C., Faraj, T.K.H., Guo, X.Y. & Maharachchikumbura, S.S.N. (2024) Phylogenetic insights reveal new taxa in Thyridariaceae and Massarinaceae. Journal of Fungi 10: 542. https://doi.org/10.3390/jof10080542
- Tibpromma, S., Hyde, K.D., McKenzie, E.H., Bhat, D.J., Phillips, A.J., Wanasinghe, D.N., Samarakoon, M.C., Jayawardena, R.S., Dissanayake, A.J., Tennakoon, D.S., Doilom, M., Phookamsak, R., Tang, A.M.C., Xu, J.C., Mortimer, P.E., Promputtha, I., Maharachchikumbura, S.S.N., Khan, S. & Karunarathna, S.C. (2018) Fungal diversity notes 840–928: micro-fungi associated with Pandanaceae. Fungal Diversity 93: 1–160. https://doi.org/10.1007/s13225-018-0408-6
- 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. https://doi.org/10.1128/jb.172.8.4238-4246.1990
- Voglmayr, H. & Jaklitsch, W.M. (2017) Corynespora, Exosporium and Helminthosporium revisited - New species and generic reclassification. Studies in Mycology 87 (1): 43–76. https://doi.org/10.1016/j.simyco.2017.05.001
- Vu, D., Groenewald, M., De Vries, M., Gehrmann, T., Stielow, B., Eberhardt, U., Verkley, G.J.M., Groenewald, J.Z., Cardinali, G. & Houbraken, J. (2019) Large-scale generation and analysis of filamentous fungal DNA barcodes boosts coverage for kingdom fungi and reveals thresholds for fungal species and higher taxon delimitation. Studies in Mycology 92: 135–154. https://doi.org/10.1016/j.simyco.2018.05.001
- Wanasinghe, D.N., Phukhamsakda, C., Hyde, K.D., Jeewon, R., Lee, H.B., Jones, E.B.G., Tibpromma, S., Tennakoon, D.S., Dissanayake, A.J., Jayasiri, S.C., Gafforov, Y., Camporesi, E., Bulgakov, T.S., Ekanayake, A.H., Perera, R.H., Samarakoon, M.C., Goonasekara, I.D., Mapook, A., Li, W.-J., Senanayake, I.C., Li, J., Norphanphoun, C., Doilom, M., Bahkali, A.H., Xu, J., Mortimer, P.E., Tibell, L., Tibell, S. & Karunarathna, S.C. (2018) Fungal diversity notes 709–839: Taxonomic and phylogenetic contributions to fungal taxa with an emphasis on fungi on Rosaceae. Fungal Diversity 89: 1–236. https://doi.org/10.1007/s13225-018-0395-7
- Wang, X.J., Wu, H.Y. & Zhang, M. (2014) A new species of Helminthosporium from Jiangsu, China. Mycotaxon 127 (1): 1–4. https://doi.org/10.5248/127.1
- White, T.J., Bruns, T., Lee, S. & Taylor, J. (1990) 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, New York, pp. 315–322. https://doi.org/10.1016/B978-0-12-372180-8.50042-1
- Wijayawardene, N.N., Hyde, K.D., Bhat, D.J., Goonasekara, I.D., Nadeeshan, D., Camporesi, E., Schumacher, R.K. & Wang, Y. (2015) Additions to brown spored coelomycetous taxa in Massarinae, Pleosporales: introducing Phragmocamarosporium gen. nov. and Suttonomyces gen. nov. Cryptogamie Mycologie 36: 213–224. https://doi.org/10.7872/crym/v36.iss2.2015.213
- Zhang, Y. & Fournier, J. (2015) Kirschsteiniothelia thujina (Peck) D. Hawksw. (Kirschsteiniotheliaceae), reported from Europe for the first time. Ascomycete.org 7 (1): 31–37.
- Zhang, K., Guo, W., Sosa, D., Magdama, F., Serrano, L., Malosso, E., Li, D.W. & Castañeda-Ruiz, R.F. (2020) Phylogeny and morphology of Ellismarsporium parvum and the new combination E. varium. Mycotaxon 135 (2): 443–452. https://doi.org/10.5248/135.443
- Zhao, N.A., Luo, Z.L., Bhat, D.J., Liu, J.K., Bao, D.F., Hao, Y.E., Su, H.Y. & Hyde, K.D. (2018) Helminthosporium submersum sp. nov. (Massarinaceae) from submerged wood in north-western Yunnan Province, China. Phytotaxa 348 (4): 69–278. https://doi.org/10.11646/phytotaxa.348.4.3
- Zimmermann, J., Voss, H., Schwager, C., Stegemann, J. & Ansorge, W. (1988) Automated Sanger dideoxy sequencing reaction protocol. FEBS Letters 233: 432–436. https://doi.org/10.1016/0014-5793(88)80477-0
- Zou, M., Al-Otibi, F., Hyde, K.D., Wang, Y. & Pan, X.-J. (2024) New Helminthosporium (Massarinaceae, Dothideomycetes) and Nigrospora (Incertae sedis, Sordariomycetes) species associated with walnut (Juglans regia L.) in China. MycoKeys 109: 265–284. https://doi.org/10.3897/mycokeys.109.133431
