Research Paper

Noteworthy foliicolous lichens and a lichenicolous fungus collected from Okinawa Island, southern Japan

Kento Miyazawa, Yoshihito Ohmura

Published on: 06 February 2026

Page: 159 - 184

DOI: 10.6165/tai.2026.71.159

Abstract

Based on field surveys conducted from 2020 to 2024 on Okinawa Island, southern Japan, a total of 26 genera, 51 species, and one subspecies of foliicolous lichens, together with one lichenicolous fungus, were identified. Among them, seven foliicolous lichens are newly recorded from Japan, including Badimia polillensis, Byssoloma brunneodiscum, B. melanodiscocarpum (= B. discordans var. flavescens, syn. nov.), Gyalectidium imperfectum, Porina perminuta, Strigula pseudosubtilissima, and S. smaragdula subsp. multiformis (comb. & stat. nov.). An additional 39 species are newly recorded from Okinawa Island. A lichenicolous fungus, Hemigrapha strigulae, is reported from Asia for the first time. A total of 169 mtSSU sequences representing 48 species and eight nuITS sequences representing three species were newly generated. For 15 species, mtSSU sequences have been deposited in GenBank for the first time. These data provide the first molecular evidence for Microtheliopsis uleana, allowing a preliminary evaluation of its phylogenetic placement within Chaetothyriales. Strigula smaragdula subsp. novae-zelandiae (comb. & stat. nov.) is also proposed.

Keyword: Asia, Ascomycota, lichenized fungi, mtSSU, nuITS, phylogeny, subspecies, taxonomy

Literature Cited

Andersen, H.L., Ekman, S. 2005 Disintegration of the Micareaceae (lichenized Ascomycota): a molecular phylogeny based on mitochondrial rDNA sequences. Mycol. Res. 109(1): 21–30.
DOI: 10.1017/S0953756204001625View Article Google Scholar

Aptroot, A., Ferraro, L.I., Lai, M.J., Sipman, H.J.M., Sparrius, L.B. 2003 Foliicolous lichens and their lichenicolous ascomycetes from Yunnan and Taiwan. Mycotaxon 88(1): 41–47.

Aptroot, A., Sparrius, L.B. 2006 Additions to the lichen flora of Vietnam, with an annotated checklist and bibliography. Bryologist 109(3): 358–371.
DOI: 10.1639/0007-2745(2006)109[358:ATTLFO]2.0.CO;2View Article Google Scholar

Aptroot, A., Sparrius, L.B., Alvarado, P. 2018 Aquacidia, a new genus to accommodate a group of skiophilous temperate Bacidia species that belong in the Pilocarpaceae (lichenized ascomycetes). Gorteria 40: 11–14. (see supplement)

Asahina, Y. 1936 Mikrochemischer Nachweis der Flechtenstoffe (I). J. Jpn. Bot. 12(7): 516–525.

Culberson, C.F., Johnson, A. 1982 Substitution of methyl tert-butyl ether for diethyl ether in the standardized thin-layer chromatographic method for lichen products. J. Chromatogr. 238(2): 483–487.
DOI: 10.1016/S0021-9673(00)81336-9View Article Google Scholar

Dai, D.Q., Tang, L.Z., Liu, C., Wang, H.B., Hyde, K.D. 2018 Studies on Parmulariaceae I. A phylogeny based on available sequence data, introducing Parmulariales ord. nov., and Hemigraphaceae, Melaspileellaceae and Stictographaceae fam. nov. Phytotaxa 396(2): 69–79.
DOI: 10.11646/phytotaxa.369.2.1View Article Google Scholar

del Prado, R., Schmitt, I., Kautz, S., Palice, Z., L?cking, R., Lumbsch, H.T. 2006 Molecular data place Trypetheliaceae in Dothideomycetes. Mycol. Res. 110(5): 511–520. (see supplement)
DOI: 10.1016/j.mycres.2005.08.013View Article Google Scholar

Diederich, P., Ertz, D., Lawrey, J.D., Sikaroodi, M., Untereiner, W.A. 2013 Molecular data place the hyphomycetous lichenicolous genus Sclerococcum close to Dactylospora (Eurotiomycetes) and S. parmeliae in Cladophialophora (Chaetothyriales). Fungal Divers. 58(1):61–72. (see supplement)
DOI: 10.1007/s13225-012-0179-4View Article Google Scholar

Diederich, P., Wedin, M. 2000 The species of Hemigrapha (lichenicolous Ascomycetes, Dothideales) on Peltigerales. Nord. J. Bot. 20(2): 203–214.
DOI: 10.1111/j.1756-1051.2000.tb01571.xView Article Google Scholar

Dong, W., Hyde, K.D., Jeewon, R., Karunarathna, S.C., Zhang, H., Rossi, W., Leonardi, M., Kezo, K., Kaliyaperumal, M., Shu, Y.-X., Yang, C.-L., Wang, F.-H., Liu, F., Ma, J., Zhang, J.-Y., Lu, Y.-Z., Abdel-Wahab, M.A., Le?o, A.F., Cust?dio, F.A., Cond?, T.O., Pereira, O.L., Liao, C.-F., Zhao, H.-J., Xu, R.-J., Zhao, Q., Du, T.-Y., Tibpromma, S., Monkai, J., Lumyong, S., He, S.-C., Bundhun, D., Yang, Y., Xiao, Y., Tennakoon, D.S., Han, L.-S., Dai, D.-Q., Li, H., Yang, Y., Manawasinghe, I.S., de Silva, N.I., Calabon, M.S., Ren, G.-C., Wei, D.-P., Wen, T.-C., Tang, X., Armand, A., Jayawardena, R.S., Singh, R., Rajwar, S., Kumar, S., Yang, Y.-Y., Tarafder, E., Acharya, K., Shen, H.-W., Luo, Z.-L., Liu, J.-W., Yu, F.-Q., Li, Y.-X., Chen, Y., Maharachchikumbura, S.S.N., Mahadevakumar, S., Sarma, P.V.S.R.N., Danteswari, C., Podile, A.R., Chandranayaka, S., Abdollahzadeh, J., Amirashayeri, P., Piri Kakihai, S., Bashiri, S., Karpowicz, F., Glejdura, S., Kunca, V., Ronikier, A., Ronikier, M., Rutkowski, R., Mleczko, P., Suwannarach, N., Kumla, J., Senwanna, C., Gafforov, Y., Sanna, M., Mua, A., Porcu, G., Casula, M., Rinaldi, A.C., Mohammadi Hamidi, L., Ahmadpour, A., Ghosta, Y., Oset, M., Kukwa, M., Guzow-Krzemi?ska, B., Usman, M., Khalid, A.N., Ossowska, E.A., Kosecka, M., Miller, S.L., Henkel, T.W., Aime, M.C., Kaygusuz, O., Bandini, D., Piepenbring, M., Kabdraisova, A., Svantesson, S., Ferreira, R.J., Mart?n, M.P., Baseia, I.G., Kiss, L., Tan, Y.P., Vaghefi, N., Shivas, R.G., Chen, L.-J., Cheng, S.-Q., Liang, Y.-S., Chen, L.-H., Chellapan, N., Arumugam, E., Vasan, V., Subramani, P., Murugadoss, R., Afshari, N., Gomdola, D., Sun, Y.-R., Abeywickrama, P.D., Gao, Y., Gomes de Farias, A.R., Gui, H., Suduri, L., Javan-Nikkhah, M., Hashemlou, E., Tian, X.-G., Velez, P., Hernandez-Monroy, A., Gasca-Pineda, J., Dissanayake, A.J., Chaiwan, N., Wu, N., Senanayake, I.C., Doilom, M. 2025 Fungal diversity notes 2017–2122: taxonomic and phylogenetic contributions to freshwater fungi and other fungal taxa. Fungal Divers. 134(1): 185–459.
DOI: 10.1007/s13225-025-00560-3View Article Google Scholar

Elix, J.A., McCarthy, P.M. 2018 Ten new lichen species (Ascomycota) from Australia. Australas. Lichenol. 82: 20–59.

Eriksson, O.E. 1981 The families of bitunicate ascomycetes. Opera Bot. 60: 1–209.

Eriksson, O.E., Reynolds D.R. 1994 Discussion 9: Arthoniales, Dothideales, Opegraphales, Pyrenulales, and Verrucariales. In: Hawksworth, D.L. (ed.), Ascomycete systematics: Problems and perspectives in the nineties, pp. 411–417. Plenum Press, New York.
DOI: 10.1007/978-1-4757-9290-4_41View Article Google Scholar

Ertz, D., Diederich, P. 2015 Dismantling Melaspileaceae: a first phylogenetic study of Buelliella, Hemigrapha, Karschia, Labrocarpon and Melaspilea. Fungal Divers. 71(1): 141–164.
DOI: 10.1007/s13225-015-0321-1View Article Google Scholar

Ertz, D., Diederich, P. 2022 Unravelling the diversity of the lichen genus Porina (Porinaceae) in Mauritius. Plant Ecol. Evol. 155(1): 123–152.
DOI: 10.5091/plecevo.84545View Article Google Scholar

Ertz, D., Heuchert, B., Braun, U., Freebury, C.E., Common, R.S., Diederich, P. 2016 Contribution to the phylogeny and taxonomy of the genus Taeniolella, with a focus on lichenicolous taxa. Fungal Biol. 120(11): 1416–1447.
DOI: 10.1016/j.funbio.2016.05.008View Article Google Scholar

Ferraro, L.I., L?cking, R., S?rusiaux, E. 2001 A world monograph of the lichen genus Gyalectidium (Gomphillaceae). Bot. J. Linn. Soc. 137(3): 311–345.
DOI: 10.1111/j.1095-8339.2001.tb01126.xView Article Google Scholar

Ford, M., Blanchon, D.J., Veale, A., Doyle, E.J., Rolfe, J.R., de Lange, P.J. 2019 Hidden in plain sight—a new Strigula species segregated from Strigula novae-zelandiae (lichenized Ascomycota: Strigulaceae). Phytotaxa 424(5): 267–281.
DOI: 10.11646/phytotaxa.424.5.1View Article Google Scholar

Frisch, A., Thor, G., Ertz, D., Grube, M. 2014 The Arthonialean challenge: restructuring Arthoniaceae. Taxon 63(4): 727–744.
DOI: 10.12705/634.20View Article Google Scholar

Gardes, M., Bruns, T.D. 1993 ITS primers with enhanced specificity for basidiomycetes - application to the identification of mycorrhizae and rusts. Mol. Ecol. 2(2): 113–118.
DOI: 10.1111/j.1365-294X.1993.tb00005.xView Article Google Scholar

Gueidan, C., Villase?or, C.R., de Hoog, G.S., Gorbushina, A.A., Untereiner, W.A., Lutzoni, F. 2008 A rock-inhabiting ancestor for mutualistic and pathogen-rich fungal lineages. Stud. Mycol. 61(1):111–119. (see supplement)
DOI: 10.3114/sim.2008.61.11View Article Google Scholar

Hafellner, J. 1984 Studien in Richtung einer naturlicheren Gliederung der Sammelfamilien Lecanoraceae und Lecideaceae. Nova Hedwig. Beih. 79: 241–371.

Halda, J.P., Woo, J.-J., Liu, D., Oh, S.-O., Joshi, Y., Hur, J.-S. 2022 Jejulea byssolomoides gen. et sp. nov., a remarkable Pilocarpaceae (lichen-forming Ascomycetes) from Jeju Island, South Korea. Mycobiology 50(3): 172–180.
DOI: 10.1080/12298093.2022.2081407View Article Google Scholar

Harada, H., Kawana, T., Suenaga, M. 1999 Lichens of Chiba-ken, central Japan (7). Three noteworthy species. Bulletin of the Biological Society of Chiba 49(2): 77–82.

Izumitsu, K., Hatoh, K., Sumita, T., Kitade, Y., Morita, A., Gafur, A., Ohta, A., Kawai, M., Yamanaka, T., Neda, H., Ohta, Y., Tanaka, C. 2012 Rapid and simple preparation of mushroom DNA directly from colonies and fruiting bodies for PCR. Mycoscience 53(5): 396–401.
DOI: 10.1007/S10267-012-0182-3View Article Google Scholar

Jiang, S.-H., L?cking, R., Liu, H.-J., Wei, X.-L., Xavier-Leite, A.B., Portilla, C.V., Ren, Q., Wei, J.-C. 2022(2021) Twelve new species reveal cryptic diversification in foliicolous lichens of Strigula s.lat. (Strigulales, Ascomycota). J. Fungi 8(1): 2.
DOI: 10.3390/jof8010002View Article Google Scholar

Jiang, S.-H., L?cking, R., Xavier-Leite, A.B., C?ceres, M.E.S., Aptroot, A., Portilla, C.V., Wei, J.-C. 2020 Reallocation of foliicolous species of the genus Strigula into six genera (lichenized Ascomycota, Dothideomycetes, Strigulaceae). Fungal Divers. 102(1): 257–291.
DOI: 10.1007/s13225-020-00445-7View Article Google Scholar

Jiang, S.-H., Wei, X.-L., Wei, J.-C. 2017a A new species and two new records of Strigula (lichenized Ascomycota) from China. Mycoscience 58(6): 391–397. (see supplement)
DOI: 10.1016/j.myc.2017.05.003View Article Google Scholar

Jiang, S.-H., Wei, X.-L., Wei, J.-C. 2017b Two new species of Strigula (lichenized Dothideomycetes, Ascomycota) from China, with a key to the Chinese foliicolous species. MycoKeys 19(1): 31–42. (see supplement)
DOI: 10.3897/mycokeys.19.11174View Article Google Scholar

J?rgensen, P.M. 2019 The troublesome genus Thamnolia (lichenized Ascomycota). Lichenologist 51(3): 221–226.
DOI: 10.1017/S0024282919000203View Article Google Scholar

Kalb, K., L?cking, R., S?rusiaux, E. 2000 Studies in Bacidia sensu lato (lichenized Ascomycetes: Lecanorales). I. The genus Bapalmuia. Mycotaxon 75(1): 281–309.
DOI: 10.5962/p.414798View Article Google Scholar

Katoh, K., Rozwicki, J., Yamada, K.D. 2019 MAFFT online service: multiple sequence alignment, interactive sequence choice and visualization. Brief. Bioinform. 20(4): 1160–1166.
DOI: 10.1093/bib/bbx108View Article Google Scholar

Kistenich, S., Timdal, E., Bendiksby, M., Ekman, S. 2018 Molecular systematics and character evolution in the lichen family Ramalinaceae (Ascomycota: Lecanorales). Taxon 67(5): 871–904. (see supplement)
DOI: 10.12705/675.1.SView Article Google Scholar

Kumar, S., Stecher, G., Li, M., Knyaz, C., Tamura, K. 2018 MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Mol. Biol. Evol. 35(6):1547–1549.
DOI: 10.1093/molbev/msy096View Article Google Scholar

Launis, A., Pyk?l?, J., van den Boom, P., S?rusiaux, E., Myllys, L. 2019 Four new epiphytic species in the Micarea prasina group from Europe. Lichenologist 51(1): 7–25. (see supplement)
DOI: 10.1017/S0024282918000555View Article Google Scholar

Lebreton, E., Ertz, D., L?cking, R., Aptroot, A., Carriconde, F., Ah-Peng, C., Huang, J.-P., Chen, K.-H., Stenger, P.-L., C?ceres, M.E.S., van den Boom, P., S?rusiaux, E., Magain, N. 2025 Global phylogeny of the family Gomphillaceae (Ascomycota, Graphidales) sheds light on the origin, diversification and endemism in foliicolous lineages. IMA Fungus 16: e144194.
DOI: 10.3897/imafungus.16.144194View Article Google Scholar

L?cking, R. 1997 Additions and corrections to the knowledge of the foliicolous lichen flora of Costa Rica. The genus Fellhanera, with notes on Bacidia pauciseptata. Trop. Bryol. 13(1): 141–173.
DOI: 10.11646/bde.13.1.16View Article Google Scholar

L?cking, R. 2008 Foliicolous lichenized Fungi. Flora Neotropica Monograph 103. Organization for Flora Neotropica and The New York Botanical Garden Press, Bronx, New York. 866 pp.

Lumbsch, H.T., Schmitt, I., Palice, Z., Wiklund, E., Ekman, S., Wedin, M. 2004 Supraordinal phylogenetic relationships of Lecanoromycetes based on a Bayesian analysis of combined nuclear and mitochondrial sequences. Mol. Phylogenet. Evol. 31(3): 822–832. (see supplement)
DOI: 10.1016/j.ympev.2003.11.001View Article Google Scholar

Lutzoni, F., Kauff, F., Cox, C.J., McLaughlin, D., Celio, G., Dentinger, B., Padamsee, M., Hibbett, D., James, T.Y., Baloch, E., Grube, M., Reeb, V., Hofstetter, V., Schoch, C., Arnold, A.E., Miadlikowska, J., Spatafora, J., Johnson, D., Hambleton, S., Crockett, M., Shoemaker, R., Sung, G.-H., L?cking, R., Lumbsch, T., O'Donnell, K., Binder, M., Diederich, P., Ertz, D., Gueidan, C., Hansen, K., Harris, R.C., Hosaka, K., Lim, Y.-W., Matheny, B., Nishida, H., Pfister, D., Rogers, J., Rossman, A., Schmitt, I., Sipman, H., Stone, J., Sugiyama, J., Yahr, R., Vilgalys, R. 2004 Assembling the fungal tree of life: progress, classification, and evolution of subcellular traits 1. Am. J. Bot. 91: 1446–1480. (see supplement)
DOI: 10.3732/ajb.91.10.1446View Article Google Scholar

Matzer, M. 1996 Lichenicolous Ascomycetes with Fissitunicate Asci on Foliicolous Lichens. Mycological Papers 171, CAB International, Wallingford. x + 202 pp.

McCarthy, P.M. 2003 Additional lichen records from Oceania 9. Foliicolous Trichotheliaceae from the Solomon Islands and Vanuatu. Australas. Lichenol. 52: 20–24.

Miadlikowska, J., Kauff, F., H?gnabba, F., Oliver, J.C., Moln?r, K., Fraker, E., Gaya, E., Hafellner, J., Hofstetter, V., Gueidan, C., Ot?lora, M.A.G., Hodkinson, B., Kukwa, M., L?cking, R., Bj?rk, C., Sipman, H.J.M., Burgaz, A.R., Thell, A., Passo, A., Myllys, L., Goward, T., Fern?ndez-Brimem, S., Hestmark, G., Lendemer, J., Lumbsch, H.T., Schmull, M., Schoch, C.L., S?rusiaux, E., Maddison, D.R., Arnold, A.E., Lutzoni, F., Stenroos, S. 2014 A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317 genera and 66 families. Mol. Phylogenet. Evol. 79: 132–168. (see supplement)
DOI: 10.1016/j.ympev.2014.04.003View Article Google Scholar

Ministry of the Environment, Japan 2025 Characteristics, Yanbaru National Park. https://www.env.go.jp/en/nature/nps/park/yambaru/point/index.html (Accessed: 12 June 2025).

Miyazawa, K., Ohmura, Y. 2023 Byssoloma orientale (Pilocarpaceae, Ascomycota), a new species from East Asia. Lichenologist 55(6): 1–7.
DOI: 10.1017/S0024282923000543View Article Google Scholar

Miyazawa, K., Ohmura, Y. 2024 Discovery of the first lichenized fungus in the family Chaetothyriaceae (Ascomycota), Ceramothyrium ryukyuense sp. nov. Pl. Fung. Syst. 69(2): 167–176.
DOI: 10.35535/pfsyst-2024-0015View Article Google Scholar

Miyazawa, K., Ohmura, Y., Okane, I. 2023 Gyalectidium setiferum (Gomphillaceae, Ascomycota), a foliicolous lichen, new to East Asia and its molecular phylogenetic position. Taiwania 68(1): 101–105.
DOI: 10.6165/tai.2023.68.101View Article Google Scholar

Miyazawa, K., Ohmura, Y., Yamaoka, Y. 2020 Aulaxina microphana (Graphidaceae, lichenized Ascomycota), new to Japan. J. Jpn. Bot. 95(3): 154–157.

Miyazawa, K., Ohmura, Y., Yamaoka, Y. 2022 Noteworthy foliicolous lichens collected from Iriomote Island, southern Japan. Taiwania 67(1): 155–163.
DOI: 10.6165/tai.2022.67.155View Article Google Scholar

Muggia, L., Fleischhacker, A., Kopun, T., Grub, M. 2016 Extremotolerant fungi from alpine rock lichens and their phylogenetic relationships. Fungal Divers. 76(1): 119–142. (see supplement)
DOI: 10.1007/s13225-015-0343-8View Article Google Scholar

Muggia, L., Kopun, T., Ertz, D. 2015 Phylogenetic placement of the lichenicolous, anamorphic genus Lichenodiplis and its connection to Muellerella-like teleomorphs. Fungal Biol. 119(11): 1115–1128. (see supplement)
DOI: 10.1016/j.funbio.2015.08.011View Article Google Scholar

Muggia, L., Kopun, T., Grube, M. 2017 Effects of growth media on the diversity of culturable fungi from lichens. Molecules 22(5): 824. (see supplement)
DOI: 10.3390/molecules22050824View Article Google Scholar

Muggia, L., P?rez-Ortega, S., Ertz, D. 2019 Muellerella, a lichenicolous fungal genus recovered as polyphyletic within Chaetothyriomycetidae (Eurotiomycetes, Ascomycota). Pl. Fung. Syst. 64(2): 367–381.
DOI: 10.2478/pfs-2019-0024View Article Google Scholar

Nag Raj, T.R. 1981 Genera coelomycetum. XIX. Discosiella, a lichenized mycobiont. Can. J. Bot. 59(12): 2519–2533.
DOI: 10.1139/b81-302View Article Google Scholar

Nakajima, A., Miyazawa, K., Ohmura, Y. 2025 Koordersiella insectivora (Lophiotremataceae, Ascomycota), a lichenicolous fungus, new to Japan. Bull. Natl. Mus. Nat. Sci. Ser. B 51(3): 91–96.
DOI: 10.1006/lich.1999.0267View Article Google Scholar

Nei, M., Kumar, S. 2000 Molecular Evolution and Phylogenetics. Oxford University Press, New York.

Ohmura, Y. 2023 Lichenes Minus Cogniti Exsiccati. Fasc. XXIX (Nos. 701–725). National Museum of Nature and Science, Tokyo. 4 pp.

Ohmura, Y., Kawachi, M., Kasai, F., Watanabe, M.M., Takeshita, S. 2006 Genetic combinations of symbionts in a vegetatively reproducing lichen, Parmotrema tinctorum, based on ITS rDNA sequences. Bryologist 109(1): 43–59.
DOI: 10.1639/0007-2745(2006)109[0043:GCOSIA]2.0.CO;2View Article Google Scholar

Okinawa Prefecture Governor's Office, Base Countermeasures Division 2024 U.S. Military and Japan Self-Defense Force Bases in Okinawa (Statistical Report), May 2024. https://www.pref.okinawa.lg.jp/heiwakichi/kichi/1017273/1017274/1025057/1029383.html (in Japanese) (Accessed: 21 September 2024).

Quan, Y., Muggia, L., Moreno, L.F., Wang, M., Al-Hatmi, A.M.S., Menezes, N. da S., Shi, D., Deng., S., Ahmed, S., Hyde, K.D., Vicente, V.A., Kang, Y., Stielow, J. B., de Hoog, S. 2020 A re-evaluation of the Chaetothyriales using criteria of comparative biology. Fungal Divers. 103(1): 47–85.
DOI: 10.1371/journal.pone.0063547View Article Google Scholar

R?blov?, M., Untereiner, W.A., R?blov?, K. 2013 Novel evolutionary lineages revealed in the Chaetothyriales (Fungi) based on multigene phylogenetic analyses and comparison of ITS secondary structure. PLOS ONE 8(5): e63547.
DOI: 10.1371/journal.pone.0063547View Article Google Scholar

Rossman, A.Y., Schoch, C.L., Farr, D.F., Nishijima, K., Keith, L., Goenaga, R. 2010 Dolabra nepheliae on rambutan and lychee represents a novel lineage of phytopathogenic Eurotiomycetes. Mycoscience 51(4): 300–309. (see supplement)
DOI: 10.47371/mycosci.MYC51300View Article Google Scholar

Saitou, N., Nei, M. 1987 The neighbor-joining method: a new method for reconstructing phylogenetic trees. Mol. Biol. Evol. 4(4): 406–425.
DOI: 10.1093/oxfordjournals.molbev.a040454View Article Google Scholar

Santesson, R. 1952 Foliicolous lichens I. A revision of the taxonomy of the obligately foliicolous, lichenized fungi. Symb. Bot. Upsal. 12(2): 1–590.

Santesson, R., L?cking, R. 1999 Additions to the foliicolous lichen flora of the Ivory Coast and Guinea (tropical West Africa). Nord. J. Bot. 19(6): 719–734.
DOI: 10.1111/j.1756-1051.1999.tb00681.xView Article Google Scholar

Schumm, F., Aptroot, A. 2010 Seychelles Lichen Guide. Beck, OHG, 73079 S?ssen, Germany. 404 pp.

Schumm, F., Elix, J.A. 2015 Atlas of Images of Thin Layer Chromatograms of Lichen Substances. Books on Demand GmbH, Norderstedt. 584 pp.

S?rusiaux, E. 1984 Three New Species of Tricharia (Lichenes, Asterothyriaceae) from New Guinea. Mycologia 76(1): 108–114.
DOI: 10.1080/00275514.1984.12023814View Article Google Scholar

S?rusiaux, E. 1998 Further observations on the Lichen Genus Strigula in New Zealand. Bryologist 101(1): 147–152.
DOI: 10.2307/3244088View Article Google Scholar

Stenroos, S., Huhtinen, S., Lesonen, A., Palice, Z., Printzen, C. 2009 Puttea, gen. nov., erected for the enigmatic lichen Lecidea margaritella. Bryologist 112(3): 544–557. (see supplement)
DOI: 10.1639/0007-2745-112.3.544View Article Google Scholar

Sun, W., Su, L., Yang, S., Sun, J., Liu, B., Fu, R., Wu, B., Liu, X., Cai, L., Guo, L., Xiang, M. 2020 Unveiling the hidden diversity of rock-inhabiting fungi: Chaetothyriales from china. J. Fungi 6(4): 187. (see supplement)
DOI: 10.3390/jof6040187View Article Google Scholar

Suto, Y., Ohtani, S. 2015 Taxonomic study on five species of foliicolous lichenized Ascomycota from Shimane-ken, western Japan. Lichenology 14(1): 27–36.

Suto, Y., Ohtani, S. 2018 Two new species of foliicolous lichenized Ascomycota, Asterothyrium sasae (Asterothyriaceae) and Gyalectidium shimanense (Gomphillaceae) from Shimane-ken, western Japan. Lichenology 17(2): 35–44.

Tamura, K. 1992 Estimation of the number of nucleotide substitutions when there are strong transition-transversion and G + C-content biases. Mol. Biol. Evol. 9(4): 678–687.
DOI: 10.1093/oxfordjournals.molbev.a040752View Article Google Scholar

Tamura, K., Nei, M. 1993 Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Mol. Biol. Evol. 10(3): 512–526.
DOI: 10.1093/oxfordjournals.molbev.a040023View Article Google Scholar

Thor, G., L?cking, R., Matsumoto, T. 2000 The foliicolous lichens of Japan. Symb. Bot. Upsal. 32(3): 1–72.

Untereiner, W.A., Gueidan, C., Orr, M.-J., Diederich, P. 2011 The phylogenetic position of the lichenicolous ascomycete Capronia peltigerae. Fungal Divers. 49(1): 225–233. (see supplement)
DOI: 10.1007/s13225-011-0097-xView Article Google Scholar

Vain, E.A. 1924 Lichenes a W. A. Setchell et H. E. Parks in insula Tahiti a 1922 collecti. Univ. Calif. Publ. Bot. 12(1): 1–16.

Vasse, M., Voglmayr, H., Mayer, V., Gueidan, C., Nepel, M., Moreno, L., de Hoog, S., Selosse, M.-A., McKey, D., Blatrix, R. 2017 A phylogenetic perspective on the association between ants (Hymenoptera: Formicidae) and black yeasts (Ascomycota: Chaetothyriales) Proc. R. Soc. B 284(1850): 20162519.
DOI: 10.1098/rspb.2016.2519View Article Google Scholar

V?zda, A. 1994 Neue foliicole Flechten Ⅱ. Nova Hedwigia 58(1–2): 123–143.

Vilgalys, R., Hester, M. 1990 Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. J. Bacteriol. 172(8): 4238–4246.
DOI: 10.1128/jb.172.8.4238-4246.1990View Article Google Scholar

Wang, W.-C., Abas, A. Wei, X.-L., Qian, X., Wei, J.-C. 2023 Two new species of the lichenized genus Lasioloma (Byssolomataceae) from Asia. Lichenologist 55(1): 1–7.
DOI: 10.1017/S0024282923000014View Article Google Scholar

Wang, W.-C., Sangvichien, E., Buaruang, K., Jiang, S.-H., Wei, T.-Z., Wei, J.-C. 2021 A preliminary study of Badimia V?zda (Ramalinaceae) in East Asia. Lichenologist 53(4): 327–334.
DOI: 10.1017/S002428292100013XView Article Google Scholar

Wang, W.-C., Sangvichien, E., Wei, T.Z., Wei, J.-C. 2020a A molecular phylogeny of Pilocarpaceae Zahlbr., including a new species of Tapellaria M?ll. Arg. and new records of foliicolous lichenized fungi from Thailand. Lichenologist 52(5): 377–385.
DOI: 10.1017/S0024282920000328View Article Google Scholar

Wang, W.-C., van den Boom, P.P.G., Sangvichien, E., Wei, J.-C. 2020b A molecular study of the lichen genus Byssoloma Trevisan (Pilocarpaceae) with descriptions of three new species from China. Lichenologist 52(5): 387–396.
DOI: 10.1017/S0024282920000390View Article Google Scholar

Weerakoon, G., Aptroot, A., Lumbsch, H.T., Wolseley, P.A., Wijeyaratne, S.C., Gueidan, C. 2012. New molecular data on Pyrenulaceae from Sri Lanka reveal two well-supported groups within this family. Lichenologist 44(5): 639–647. (see supplement)
DOI: 10.1017/S0024282912000333View Article Google Scholar

Woo, J.-J., L?cking, R., Oh, S.-Y., Jeun, Y.-C., Hur, J.-S. 2020 Two new foliicolous species of Strigula (Strigulaceae, Strigulales) in Korea offer insight in phorophyte-dependent variation of thallus morphology. Phytotaxa 443(1): 1–12.
DOI: 10.11646/phytotaxa.443.1.1View Article Google Scholar

Woo, J.-J., L?cking, R., Oh, S.-Y., Jeun, Y.-C., Hur, J.-S. 2025 Uncovering hidden lineages in Korean foliicolous lichens (Strigulaceae, Strigulales): Discovery of a new cryptic genus and species. Mycobiology 53(4): 556–571.
DOI: 10.1080/12298093.2025.2530843View Article Google Scholar

Xavier-Leite, A.B., C?ceres, M.E.S., Aptroot, A., Moncada, B., L?cking, R., Goto, B.T. 2022 Phylogenetic revision of the lichenized family Gomphillaceae (Ascomycota: Graphidales) suggests post-K–Pg boundary diversification and phylogenetic signal in asexual reproductive structures. Mol. Phylogenet. Evol. 168: 107380.
DOI: 10.1016/j.ympev.2021.107380View Article Google Scholar

Yamamoto, Y., Moriguchi, M., Sato, H., Sugimoto, Mas., Sugimoto, May., Tawata, T. 2021 Lichens in Hentona, Kunigami Village, Okinawa Pref., Japan. The Nanki Seibutsu 63(2): 145–149. (In Japanese)

Yang, E.-F., Karunarathna, S.C., Dai, D.-Q., Stephenson, S.L., Elgorban, A.M., Al-Rejaie, S., Xiong, Y.-R., Promputtha, I., Samarakoon, M.C., Tibpromma, S. 2022 Taxonomy and phylogeny of fungi associated with Mangifera indica from Yunnan, China. J. Fungi 8(12): 1249.
DOI: 10.3390/jof8121249View Article Google Scholar

Yang, H., Hyde, K.D., Karunarathna, S.C., Deng, C., Gu, C.-H., Yang, S.A., Zhang, Z.-C. 2018 New species of Camptophora and Cyphellophora from China, and first report of sexual morphs for these genera. Phytotaxa 342(2): 149–159.
DOI: 10.11646/phytotaxa.343.2.5View Article Google Scholar

Yao, Z.-T., Jiang, S.-H., Jia, Z.-F. 2021 Mazosia weii sp. nov. (Roccellaceae) from China, a new species supported by molecular data. Bryologist 124(3): 335–342.
DOI: 10.1639/0007-2745-124.3.335View Article Google Scholar

Zoller, S., Scheidegger, C., Sperise, C. 1999 PCR primers for the amplification of mitochondrial small subunit ribosomal DNA of lichen-forming ascomycetes. Lichenologist 31(5): 511–516.
DOI: 10.1006/lich.1999.0220View Article Google Scholar