Biodiversity assessment of ascomycetes inhabiting Lobariella lichens in Andean cloud forests led to one new family, three new genera and 13 new species of lichenicolous fungi
More details
Hide details
Department of Lichenology, W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Kraków, Poland
Navarro Villoslada 16, 3º dcha., E-31003 Pamplona, Navarra, Spain
Department of Biology, Duke University, Durham, NC 27708, USA
Department of Plant Taxonomy and Nature Conservation, Faculty of Biology, University of Gdańsk, Wita Stwosza 59, 80-308 Gdańsk, Poland
Laboratory of Molecular Analyses, W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Kraków, Poland
Publication date: 2019-12-16
Plant and Fungal Systematics 2019; 64(2): 283–344
Neotropical mountain forests are characterized by having hyperdiverse and unusual fungi inhabiting lichens. The great majority of these lichenicolous fungi (i.e., detectable by light microscopy) remain undescribed and their phylogenetic relationships are mostly unknown. This study focuses on lichenicolous fungi inhabiting the genus Lobariella (Peltigerales), one of the most important lichen hosts in the Andean cloud forests. Based on molecular and morphological data, three new genera are introduced: Lawreyella gen. nov. (Cordieritidaceae, for Unguiculariopsis lobariella), Neobaryopsis gen. nov. (Cordycipitaceae), and Pseudodidymocyrtis gen. nov. (Didymosphaeriaceae). Nine additional new species are described (Abrothallus subhalei sp. nov., Atronectria lobariellae sp. nov., Corticifraga microspora sp. nov., Epithamnolia rugosopycnidiata sp. nov., Lichenotubeufia cryptica sp. nov., Neobaryopsis andensis sp. nov., Pseudodidymocyrtis lobariellae sp. nov., Rhagadostomella hypolobariella sp. nov., and Xylaria lichenicola sp. nov.). Phylogenetic placements of 13 lichenicolous species are reported here for Abrothallus, Arthonia, Globonectria, Lawreyella, Monodictys, Neobaryopsis, Pseudodidymocyrtis, Sclerococcum, Trichonectria and Xylaria. The name Sclerococcum ricasoliae comb. nov. is reestablished for the neotropical populations formerly named S. lobariellum (Sclerococcales). A key to sexual and asexual states of 40 species of lobariellicolous ascomycetous fungi is provided. Teleomorph-anamorph connections were established for several species using molecular methods and/or visual observations in nature. Additionally, we found that the anamorphic species Cornutispora ophiurospora inhabiting Lobariella was often accompanied by ascomata of Spirographa. Results of phylogenetic analyses, including newly generated sequences of several Cornutispora and Spirographa species inhabiting various host lichens, support the conclusion that Cornutispora is a synonym of Spirographa. Our Maximum Likelihood inference based on multiple loci show that all studied Spirographa (including Cornutispora) belong to a new lineage within Ostropales. Based on these highly supported phylogenetic placements and the distinct character states of their conidiomata, in comparison with other Lecanoromycetes, a new family is proposed – Spirographaceae fam. nov. This new lineage includes broadly distributed mycoparasites, inhabiting various lichen and fungal hosts, and representing an early diversification event preceding the lichen-forming clade of Fissurinaceae, Gomphillaceae and Graphidaceae. Two lichenicolous species, Asteroglobulus giselae and Pleoscutula arsenii, were found to be nested within the Spirographa clade, and their teleomorph-anamorph connections were confirmed based on genotypic and phenotypic data. This phylogenetic result is corroborated by their highly similar ascomata anatomy. Together these results strongly indicate that both species are congeneric with Spirographa. As a result, four new species (S. aggregata sp. nov., S. galligena sp. nov., S. maroneae sp. nov., and S. parmotrematis sp. nov.) and 15 new combinations are proposed (Spirographa ascaridiella comb. nov., S. arsenii comb. nov., S. ciliata comb. nov., S. giselae comb. nov., S. herteliana comb. nov., S. hypotrachynae comb. nov., S. intermedia comb. nov., S. lichenicola comb. nov., S. limaciformis comb. nov., S. ophiurospora comb. nov., S. pittii comb. nov., S. pyramidalis comb. nov., S. triangularis comb. nov., S. tricupulata comb. nov., and S. vermiformis comb. nov.). Species of the genus Spirographa, as outlined here, are strongly host-specific, mainly at the generic level of their host. Some host genera can harbour more than one Spirographa species.
Alstrup, V. & Hawksworth, D. L. 1990. The lichenicolous fungi of Greenland. Meddelelser om Gronland, Bioscience 31: 1–90.
Altschul, S. F., Gish, W., Miller, W., Myers, E. W. & Lipman, D. J. 1990. Basic local alignment search tool. Journal of Molecular Biology 215: 403–410.
Ariyawansa, H.A., Tanaka, K., Thambugala, K. M., Phookamsak, R., Tian, Q., Camporesi, E., Hongsanan, S., Monkai, J., Wanasinghe, D. N., Mapook, A., Chukeatirote, E., Kang, J.-C., Xu, J.-C., McKenzie, E. H. C., Jones, E. B. G. & Hyde, K. D. 2014. A molecular phylogenetic reappraisal of the Didymosphaeriaceae (= Montagnulaceae). Fungal Diversity 68: 69–104.
Ashrafi, S., Helaly, S., Schroers, H. J., Stadler, M., Richert-Poeggeler, K. R., Dababat, A. A. & Maier, W. 2017. Ijuhya vitellina sp. nov., a novel source for chaetoglobosin A, is a destructive parasite of the cereal cyst nematode Heterodera filipjevi. PLoS One 12: e0180032.
Aptroot, A. 2004. Two new ascomycetes with long gelatinous appendages collected from monocots in the tropics. Studies in Mycology 50: 307–312.
Baloch, E., Gilenstam, G. & Wedin, M. 2013 The relationships of Odontotrema (Odontotremataceae) and the resurrected Sphaeropezia (Stictidaceae) – new combinations and three new Sphaeropezia species. Mycologia 105: 384–397.
Booth, C. 1971. Fusarium: Laboratory Guide to the Identification of the Major Species. The Common Wealth Mycological Institute, Kew.
Brackel, W. von. 2011a. Lichenicolous fungi and lichens from Puglia and Basilicata (southern Italy). Herzogia 24: 65–101.
Brackel, W. von. 2011b. Lichenopeltella rangiferinae sp. nov. and some other lichenicolous fungi from Iceland. Acta Botanica Islandica 15: 51–60.
Brackel, W. von. 2014. Kommentierter Katalog der flechtenbewohnenden Pilze Bayerns. Bibliotheca Lichenologica 109: 1–476.
Candoussau, F., Boqueras, M., Gómez-Bolea, A., Læssøe, T., Lowen, R., Rogers, J. D., Rossman, A. Y. & Samuels, G. J. 2007. Observations on Neobarya, including new species and new combinations. Sydowia 59: 179–215.
Carbone, I., White, J. B., Miadlikowska, J., Arnold, A. E., Miller, M. A., Kauff, F., U’Ren, J. M., May, G. & Lutzoni, F. 2017. T-BAS: Tree-Based Alignment Selector toolkit for phylogenetic-based placement, alignment downloads, and metadata visualization: an example with the Pezizomycotina tree of life. Bioinformatics 33:1160–1168.
Carbone, I., White, J. B., Miadlikowska, J., Arnold, A. E., Miller, M. A., Magain, N., U’Ren, J. M. & Lutzoni, F. 2019. T-BAS version 2.1: Tree-Based Alignment Selector toolkit for evolutionary placement and viewing of alignments and metadata on curated and custom trees. Microbiology Resource Announcements 8: e00328-19.
Castresana, J. 2000. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution 17: 540–552.
Castlebury, L. A., Rossman, A. Y., Sung, G., Hyten, A. S., & Spatafora, J. W. 2004. Multigene phylogeny reveals new lineage for Stachybotrys chartarum, the indoor air fungus. Mycological Research 108: 864–872.
Chaverri, P., Bischoff, J. F., Evans, H. C. & Hodge, K. T. 2005. Regiocrella, a new entomopathogenic genus with a pycnidial anamorph and its phylogenetic placement in the Clavicipitaceae. Mycologia 97: 1225–1237.
Chaverri, P., Salgado, C., Hirooka, Y., Rossman, A. Y. & Samuels, G. J. 2011. Delimitation of Neonectria and Cylindrocarpon (Nectriaceae, Hypocreales, Ascomycota) and related genera with Cylindrocarpon-­like anamorphs. Studies in Mycology 68: 57–78.
Cole, M. S. & Hawksworth, D. L. 2001. Lichenicolous fungi, mainly from the USA, including Patriciomyces gen. nov. Mycotaxon 77: 305–338.
Crous, P. W., Mohammed, C., Glen, M., Verkley, G. J. M. & Groenewald, J. Z. 2007. Eucalyptus microfungi known from culture. 3. Eucasphaeria and Sympoventuria genera nova, and new species of Furcaspora, Harknessia, Heteroconium and Phacidiella. Fungal Diversity 25: 19–36.
Crous, P. W., Wingfield, M. J., Burgess, T. I., Hardy, G. E., Crane, C., Barrett, S., Cano-Lira, J. F., Le Roux, J. J., Thangavel, R., Guarro, J., Stchigel, A. M., Martín, M. P., Alfredo, D. S., Barber, P. A., Barreto, R. W., Baseia, I. G., Cano-Canals, J., Cheewangkoon, R., Ferreira, R. J., Gené, J., Lechat, C., Moreno, G., Roets, F., Shivas, R. G., Sousa, J. O., Tan, Y. P., Wiederhold, N. P., Abell, S. E., Accioly, T., Albizu, J. L., Alves, J. L., Antoniolli, Z. I., Aplin, N., Araújo, J., Arzanlou, M., Bezerra, J. D., Bouchara, J. P., Carlavilla, J. R., Castillo, A., Castroagudín, V. L., Ceresini, P. C., Claridge, G. F., Coelho, G., Coimbra, V. R., Costa, L. A., da Cunha, K. C., da Silva, S. S., Daniel, R., de Beer, Z. W., Dueñas, M., Edwards, J., Enwistle, P., Fiuza, P. O., Fournier, J., García, D., Gibertoni, T. B., Giraud, S., Guevara-Suarez, M., Gusmão, L. F., Haituk, S., Heykoop, M., Hirooka, Y., Hofmann, T. A., Houbraken, J., Hughes, D. P., Kautmanová, I., Koppel, O., Koukol, O., Larsson, E., Latha, K. P., Lee, D. H., Lisboa, D. O., Lisboa, W. S., López-Villalba, Á., Maciel, J. L., Manimohan, P., Manjón, J. L., Marincowitz, S., Marney, T. S., Meijer, M., Miller, A. N., Olariaga, I., Paiva, L. M., Piepenbring, M., Poveda-Molero, J. C., Raj, K. N., Raja, H. A., Rougeron, A., Salcedo, I., Samadi, R., Santos, T. A., Scarlett, K., Seifert, K. A., Shuttleworth, L. A., Silva, G. A., Silva, M., Siqueira, J. P., Souza-Motta, C. M., Stephenson, S. L., Sutton, D. A., Tamakeaw, N., Telleria, M. T., Valenzuela-Lopez, N., Viljoen, A., Visagie, C. M., Vizzini, A., Wartchow, F., Wingfield, B. D., Yurchenko, E., Zamora, J. C. & Groenewald, J. Z. 2016. Fungal Planet description sheets: 469–557. Persoonia 37: 218–403.
Culberson, C. F. & Kristinsson, H. 1970. A standardized method for the identification of lichen products. Journal of Chromatography 46: 85–93.
Currie, C. R., Wong, B., Stuart, A. E., Schultz, T. R., Rehner, S. A., Mueller, U. G., Sung, G., Spatofora, J. W. & Straus, N. A. 2003 Ancient tripartite coevolution in the attine ant-microbe symbiosis. Science 299: 386–388.
Diederich, P. 1997. Lichenicolous fungi in the tropics. Bibliotheca Lichenologica 68: 215–221.
Diederich, P. 2003. New species and new records of American lichenicolous fungi. Herzogia 16: 41–90.
Diederich, P. 2004. Spirographa. In: Nash III, T.H., Ryan, B. D., Diederich, P., Gries, C. & Bungartz, F. (eds): Lichen Flora of the Greater Sonoran Desert Region, Vol. 2, pp. 702–703. Lichens Unlimited, Arizona State University, Tempe, Arizona.
Diederich, P. & Coppins, B. J. 2014. Diplolaeviopsis symmictae (Helotiales, Ascomycota), a new lichenicolous fungus on Lecanora symmicta. Bulletin de la Société des Naturalistes Luxembourgeois 115: 151–155.
Diederich, P. & Etayo, J. 2000. A synopsis of the genera Skyttea, Llimoniella and Rhymbocarpus (lichenicolous Ascomycota, Leotiales). The Lichenologist 32: 423–485.
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 Diversity 58: 61–72.
Diederich, P., Lawrey, J. D., Capdet, M., Pereira, S., Romero, A. I., Etayo, J., Flakus, A., Sikaroodi, M.& Ertz, D. 2014. New lichen-associated bulbil-forming species of Cantharellales (Basidiomycetes). The Lichenologist 46: 333–347.
Diederich, P., Lawrey, J. D. & Ertz, D. 2018. The 2018 classification and checklist of lichenicolous fungi, with 2000 non-lichenized, obligately lichenicolous taxa. The Bryologist 12: 340–425.
Diederich, P., Common, R. S., Braun, U., Heuchert, B., Millanes, A., Suija, A. & Ertz. D. 2019. Lichenicolous fungi from Florida growing on Graphidales. Plant and Fungal Systematics 64: 249–282.
Ekanayaka, A. H., Jones, G. E. B., Hyde, K. D. & Zhao, Q. 2019. A stable phylogeny for Dactylosporaceae. Cryptogamie Mycologie 40: 23–44.
Ertz, D., Miadlikowska, J., Lutzoni, F., Dessein, S., Raspé, O., Vigneron, N., Hofstetter, V. & Diederich, P. 2009. Towards a new classification of the Arthoniales (Ascomycota) based on a three-gene phylogeny focussing on the genus Opegrapha. Mycological Research 113: 141–152.
Ertz, D., Diederich, P., Lawrey, J. D., Berger, F., Freebury, C. E., Coppins, B., Gardiennet, A. & Hafellner, J. 2015. Phylogenetic insights resolve Dacampiaceae (Pleosporales) as polyphyletic: Didymocyrtis (Pleosporales, Phaeosphaeriaceae) with Phoma-like anamorphs resurrected and segregated from Polycoccum (Trypetheliales, Polycoccaceae fam. nov.). Fungal Diversity 74: 53–89.
Etayo, J. 2002. Aportaciόn al conocimiento de los hongos liquenícolas de Colombia. Bibliotheca Lichenologica 84: 1–154.
Etayo, J. 2010a. Lichenicolous fungi from the western Pyrenees. V. Three new ascomycetes. Opuscula Philolichenum 8: 131–139.
Etayo, J. 2010b. Hongos liquenícolas de Perú. Bulletin de la Société linnéenne de Provence 61: 1–46.
Etayo, J. 2017. Hongos liquenícolas de Ecuador. Opera Lilloana 50: 1–535.
Etayo, J. & Diederich, P. 1995. Lichenicolous fungi from the western Pyrenees, France and Spain. I. New species of Deuteromycetes, pp. 205–221. In: Daniels, F. J. A., Schulz, M.& Peine, J. (eds), Flechten Follmann. Contributions to Lichenology in Honour of Gerhard Follmann. Geobotanical and Phytotaxonomical Study Group, Botanical Institute, University of Cologne, Cologne.
Etayo, J. & Diederich, P. 1996a. Lichenicolous fungi from the western Pyrenees, France and Spain. II. More Deuteromycetes. Mycotaxon 60: 415–428.
Etayo, J. & Diederich, P. 1996b. Lichenicolous fungi from the western Pyrenees, France and Spain. III. Species on Lobaria pulmonaria. Bulletin de la Société des Naturalistes Luxembourgeois 97: 93–118.
Etayo, J. & Sancho, L. G. 2008. Hongos liquenícolas del Sur de Sudamérica, especialmente de Isla Navarino (Chile). Bibliotheca Lichenologica 98: 1–302.
Etayo, J., Flakus, A., Suija, A. & Kukwa, M. 2015. Macroskyttea parmotrematis gen. et sp. nov. (Helotiales, Leotiomycetes, Ascomycota), a new lichenicolous fungus from Bolivia. Phytotaxa 224: 247–257.
Flakus, A. & Kukwa, M. 2012a. New records of lichenicolous fungi from Bolivia. Opuscula Philolichenum 11: 36–48.
Flakus, A. & Kukwa, M. 2012b. New species of lichenicolous fungi from Bolivia. The Lichenologist 44: 469–477.
Flakus, A., Kukwa, M.& Czarnota, P. 2006. Some interesting records of lichenized and lichenicolous Ascomycota from South America. Polish Botanical Journal 51: 209–215.
Flakus, A., Etayo, J., Pérez-Ortega, S., Kukwa, M., Palice, Z. & Rodriguez-Flakus, P. 2019. A new genus, Zhurbenkoa, and a novel nutritional mode revealed in the family Malmideaceae (Lecanoromycetes, Ascomycota). Mycologia 111: 593–611.
Frisch, A., Thor, G., Ertz, D. & Grube, M. 2014. The arthonialean challenge: restructuring Arthoniaceae. Taxon 63: 727–744.
Frisch, A., Grube, M., Kashiwadani, H. & Ohmura, Y. 2018. Arthoniaceae with reddish, K+ purple ascomata in Japan. Phytotaxa 356: 19–33.
Fryar, S. C., Haelewaters, D. & Catcheside, D. E. A. 2019. Annabella australiensis gen. & sp. nov. (Helotiales, Cordieritidaceae) from South Australian mangroves. Mycological Progress 18: 973–981.
Galloway, D. J. & Quilhot, W. 1998. Checklist of Chilean lichen-forming and lichenicolous fungi. Gayana Botanica 55: 111–185.
Gams, W., Stielow, B. Gräfenhan, T.& Schroerset, H.-J. 2019. The ascomycete genus Niesslia and associated monocillium-like anamorphs. Mycological Progress 18: 5–76.
Gerlach, W., Nirenberg, H. I. 1982. The genus Fusarium – A pictorial atlas Mitteilungen aus der Biologischen Bundesanstalt für Land-und Forstwirtschaft Berlin-Dahlem. Germany, Kommissionsverlag Paul Parey.
Gierl, C. & Kalb, K. 1993. Die Flechtengattung Dibaeis. Eine Übersicht über die rosafrüchtigen Arten von Baeomyces sens. lat. nebst Anmerkungen zu Phyllobaeis gen. nov. Herzogia 9: 593–645.
Giralt, M. & Hawksworth, D. L. 1991. Diplolaeviopsis ranula, a new genus and species of lichenicolous coelomycetes growing on the Lecanora strobilina group in Spain. Mycological Research 95: 759–761.
Grum-Grzhimaylo, A. A., Debets, A. J. M., van Diepeningen, A. D., Georgieva, M. L. & Bilanenko, E. N. 2013a. Sodiomyces alkalinus, a new holomorphicalkaliphilic ascomycete within the Plectosphaerellaceae. Persoonia 31: 147–158.
Grum-Grzhimaylo, A. A., Georgieva, M. L., Debets, A. J. M. & Bilanenko, E. N. 2013b. Are alkalitolerant fungi of the Emericellopsis lineage (Bionectriaceae) of marine origin? IMA Fungus 4: 213–228.
Hafellner, J. 1979. Karschia. Revision einer Sammelgattung an der Grenze von lichenisierten und nichlichenisier ten Ascomyceten. Beiheft zur Nova Hedwigia 62: 1–248.
Hafellner, J. 1982. Studien über lichenicole Pilze und Flechten I. Die Gattung Pleoscutula Vouaux. Herzogia 6: 289–298.
Halleen, F., Schroers, H. J., Groenewald, J. Z. & Crous, P. W. 2004. Novel species of Cylindrocarpon (Neonectria) and Campylocarpon gen. nov. associated with black foot disease of grapevines (Vitis spp.). Studies in Mycology 50: 431–456.
Hawksworth, D. L. 1976. New and interesting microfungi from Slapton, South Devonshire: Deuteromycotina III. Transactions of the British Mycological Society 67: 51–59.
Hawksworth, D. L. 1979. The lichenicolous hyphomycetes. Bulletin of the British Museum (Natural History), Botany Series 6: 183–300.
Hawksworth, D. L. 1980. Notes on some fungi occurring on Peltigera, with a key to accepted species. Transactions of the British Mycological Society 74: 363–386.
Hawksworth, D. L. & Santesson, R. 1990. A revision of the lichenicolous fungi previously referred to Phragmonaevia. Bibliotheca Lichenologica 38: 121–143.
Hawksworth, D. L. & Sherwood M. A. 1982. Two new families in the Ascomycotina. Mycotaxon 16: 262–264.
Hawksworth, D. L., Millanes, A. M. & Wedin, M. 2010. Roselliniella revealed as an overlooked genus of Hypocreales, with the description of a second species on parmelioid lichens. Persoonia 24: 12–17.
Heuchert, B. & Braun, U. 2006. On some dematiaceous lichenicolous hyphomycetes. Herzogia 19: 11–21.
Hijikawa, Y., Matsuzaki, M., Suzuki, S., Inaoka, D. K., Tatsumi, R., Kido, Y., & Kita, K. 2016. Re-identification of the ascofuranone-producing fungus Ascochyta viciae as Acremonium sclerotigenum. The Journal of Antibiotics 70: 304–307.
Hirooka, Y., Rossman, A. Y. & Chaverri, P. 2011. A morphological and phylogenetic revision of the Nectria cinnabarina species complex. Studies in Mycology 68: 35–56.
Hirooka, Y., Kobayashi, T., Ono, T., Rossman, A. Y. & Chaverri, P. 2010. Verrucostoma, a new genus in the Bionectriaceae from the Bonin Islands, Japan. Mycologia 102: 418–429.
Holien, H. & Triebel, D. 1996. Spirographa vinosa, a new odontotremoid fungus on Ochrolechia and Pertusaria. The Lichenologist 28: 307–313.
Hsieh, W. H., Chen, C. Y. & Sivanesan, A. 1997. Iodosphaeria polygoni sp. nov., a new pyrenomycete from Taiwan. Mycological Research 101: 841–842.
Hsieh, H. M., Lin, C. R., Fang, M. J., Rogers, J. D., Fournier, J., Lechat, C. & Ju Y. M. 2010 Phylogenetic status of Xylaria subgenus Pseudoxylaria among taxa of the subfamily Xylarioideae (Xylariaceae) and phylogeny of the taxa involved in the subfamily. Molecular Phylogenetics Evolutin 54: 957–969.
Huanraluek, N., Ertz, D., Phukhamsakda, C., Hongsanan, S., Jayawardena, R. S. & Hyde, K. D. 2019. The family Pyrenidiaceae resurrected. Mycosphere 10: 634–654.
Huhtinen, S., Hawksworth, D. L. & Ihlen, P. G. 2008. Observations on two glassy-haired lichenicolous discomycetes. The Lichenologist 40: 549–557.
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.
Ihlen, P. G. & Wedin, M. 2008. An annotated key to the lichenicolous Ascomycota (including mitosporic morphs) of Sweden. Nova Hedwigia 86: 275–365.
Jaklitsch, W. M. 2009. European species of Hypocrea part I. The greenspored species. Studies in Mycology 63: 1–91.
Jaklitsch, W. M. & Voglmayr, H. 2011. Nectria eustromatica sp. nov., an exceptional species with a hypocreaceous stroma. Mycologia 103: 209–218.
Johnson, D., Sung, G. H., Hywel-Jones, N. L., Luangsa-ard, J., Bischoff, J. F., Kepler, R. M. & Spatafora, J. W. 2009. Systematics and evolution of the genus Torrubiella (Hypocreales, Ascomycota). Mycological Research 113: 279–289.
Katoh, K., Kuma, K., Toh, H. & Miyata, T. 2005. MAFFT version 5: improvement in accuracy of multiple sequence alignment. Nucleic Acids Research 33: 511–518.
Kepler, R. M., Luangsa-ard, J. J., Hywel-Jones, N. L., Quandt, C. A., Sung, G.-H., Rehner, S. A., Aime, M. C., Henkel, T. W., Sanjuan, T., Zare, R., Chen, M., Li, Z., Rossman, A. Y., Spatafora, J. W. & Shrestha, B. 2017. A phylogenetically-based nomenclature for Cordycipitaceae (Hypocreales). IMA Fungus 8: 335–353.
Knoph, J.-G. 2004. Cornutispora herteliana, a new lichenicolous coelomycete from Japan. Bibliotheca Lichenologica 88: 345–348.
Kocourková, J. 2000 [1999]. Lichenicolous fungi of the Czech Republic (the first commented checklist). Acta Musei Nationalis Pragae, Series S, Historia Naturalis 55: 59–169.
Kondratyuk, S. Y. & Galloway, D. J. 1995a. Some new species of lichenicolous fungi. Bibliotheca Lichenologica 58: 235–244.
Kondratyuk, S. Y. & Galloway, D. J. 1995b. Two new lichenicolous fungi from Lobaria and Sticta (Stictaceae), pp. 255–261. In: Daniels, F. J. A., Schulz, M. & Peine, J. (eds), Flechten Follmann. Contributions to Lichenology in Honour of Gerhard Follmann. Geobotanical and Phytotaxonomical Study Group, Botanical Institute, University of Cologne, Cologne.
Kraichak, E., Huang, J.-P., Nelsen, M., Leavitt, S. D. & Lumbsch, H. T. 2018. A revised classification of orders and families in the two major subclasses of Lecanoromycetes (Ascomycota) based on a temporal approach. Botanical Journal of the Linnean Society 188:233–249.
Lanfear, R., Calcott, B., Ho, S.Y. & Guindon, S. 2012. PartitionFinder: Combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution 29: 1695–1701.
Lanfear, R., Frandsen, P. B., Wright, A. M., Senfeld, T. & Calcott, B. 2016. PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution 34: 772–773.
Lawrey, J. D., Rossman, A. Y. & Lowen, R. 1994. Inhibition of selected hypocrealean fungi by lichen secondary metabolites. Mycologia 86: 502–506.
Lawrey, J. D., Etayo, J., Dal-Forno, M., Driscoll, K. E. & Diederich, P. 2015. Molecular data support establishment of a new genus for the lichenicolous species Neobarya usneae (Hypocreales). The Bryologist 118: 83–92.
Lechat, C. & Fournier, J. 2016. Lasionectriella, a new genus in the Bionectriaceae, with two new species from France and Spain, L. herbicola and L. rubioi. 8: 59–65.
Lechat, C., Lesage-Meessen, L. & Favel, A. 2015. A new species of Ijuhya, I. fournieri, from French Guiana. 7: 101–104.
Lechat, C., Fournier, J., Vega, M. & Priou, J-P. 2018. Geonectria, a new genus in the Bionectriaceae from France. 10: 81–55.
Leighton, W. A. 1875 Lichenological researches in North and South Wales in 1874. Grevillea 3: 113–116.
Lombard, L., Houbraken, J., Decock, C., Samson, R. A., Meijer, M., Réblová, M., Groenewald, J. Z. & Crous, P. W. 2016. Generic hyper-diversity in Stachybotriaceae. Persoonia 36: 156–246.
Lücking, R., Stuart, B. L. & Lumbsch, H. T. 2004. Phylogenetic relationships of Gomphillaceae and Asterothyriaceae: evidence from a combined Bayesian analysis of nuclear and mitochondrial sequences. Mycologia 96: 283–294.
Lumbsch, H. T., Kraichak, E., Parnmen, S., Rivas Plata, E., Aptroot, A., Cáceres, M. E. S., Ertz, D., Feuerstein, S. C., Mercado-Díaz, J. A., Staiger, B., van den Broeck, D. & Lücking, R. 2014. New higher taxa in the lichen family Graphidaceae (lichenized Ascomycota: Ostropales) based on a three-gene skeleton phylogeny. Phytotaxa 189: 39–51.
Lutzoni, F., Wagner, P., Reeb, V. & Zoller, S. 2000. Integrating ambiguously aligned regions of DNA sequences in phylogenetic analyses without violating positional homology. Systematic Biology 49: 628–651.
Maddison, W. P. & Maddison, D.R. 2015. Mesquite: A modular system for evolutionary analysis, version 3.11. https://www.
Miadlikowska, J., Kauff, F., Högnabba, F., Oliver, J. C., Molnár, K., Fraker, E., Gaya, E., Hafellner, J., Hofstetter, V., Gueidan, C., Otálora, M. A. G., Hodkinson, B., Kukwa, M., Lücking, R., Björk, C., Sipman, H. J. M., Burgaz, A. R., Thell, A., Passo, A., Myllys, L., Goward, T., Fernández-Brime, S., Hestmark, G., Lendemer, J., Lumbsch, H. T., Schmull, M., Schoch, C. L., Sérusiaux, E., Maddison, D. R., Arnold, A. E., Stenroos, S. & Lutzoni, F. 2014. Multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 312 genera and 66 families. Molecular Phylogenetics and Evolution 79: 132–168.
Miller, M.A., Pfeiffer, W. & Schwartz, T. 2010. Creating the CIPRES Science Gateway for inference of large phylogenetic trees, pp. 45–52. In: Proceedings of the Gateway Computing Environments Workshop (GCE), New Orleans, Louisiana, 14 Nov 2010. Piscataway, IEEE.
Moncada, B., Lücking R. & Betancourt-Macuase L. 2013. Phylogeny of the Lobariaceae (lichenized Ascomycota: Peltigerales), with a reappraisal of the genus Lobariella. The Lichenologist 45: 203–263.
Müller, J. 1880. Lichenologische Beiträge. X. Flora (Regensburg) 63: 40–45.
Navarro-Rosines, P., Roux C. & Etayo J. (eds). Aproximación a la revisión del género Pyrenidium (Dacampiaceae, Dothideomycetes, hongos liquenícolas no liquenizados). Association Française de Lichénologie. Societat Catalana de Micologia (in press).
Nelson, P. E., Toussoun, T. A., Marasas, W. F. O. 1983. Fusarium species: An illustrated manual for identification. Pennsylvania, USA: Pennsylvania State University Press.
Nylander, W. 1866. Collectio lichenum ex insula Cuba. Flora (Regensburg) 49: 289–295.
Nylander, W. 1868. Addenda nova ad lichenographiam Europaeam. Contin. VII. Flora (Regensburg) 51: 161–165.
Olariaga, I., Teres, J., Martín, J., Prieto, M., & Baral, H.-O. 2019. Pseudosclerococcum golindoi gen. et sp. nov., a new taxon with apothecial ascomata and a Chalara-like anamorph within the Sclerococcales (Eurotiomycetes). Mycological Progress 18: 895–905.
Orange, A., James, P. W. & White, F. J. 2001. Microchemical Methods for the Identification of Lichens. British Lichen Society.
Pärtel, K., Baral, H.-O., Tamm, H. & Põldmaa, K. 2016. Evidence for the polyphyly of Encoelia and Encoelioideae with reconsideration of respective families in Leotiomycetes. Fungal Diversity 82: 183–219.
Pérez-Ortega, S., Suija, A., Crespo, A. & de los Ríos, A. 2014. Lichenicolous fungi of the genus Abrothallus (Dothideomycetes: Abrothallales ordo nov.) are sister to the predominantly aquatic Janhulales. Fungal Diversity 64: 295–304.
Peterson, K. R. & Pfister, D. H. 2010. Phylogeny of Cyttaria inferred from nuclear and mitochondrial sequence and morphological data. Mycologia 102: 1398–1416.
Pino-Bodas, R., Zhurbenko, M. P. & Stenroos, S. 2017. Phylogenetic placement within Lecanoromycetes of lichenicolous fungi associated with Cladonia and some other genera. Persoonia 39: 91–117.
Pirozynski, K.A. 1973. Three hyperparasites of ascomycetes. Mycologia 65: 761–767.
Pitt, W. M., Urbez-Torres, J. R. & Trouillas, F. P. 2014. Munkovalsaria donacina from grapevines and Desert Ash in Australia. Mycosphere 5: 656–661.
Põldmaa, K. 2011. Tropical species of Cladobotryum and Hypomyces producing red pigments. Studies in Mycology 68: 1–34.
Põldmaa K., Larsson E. & Köljalg U. 1999. Phylogenetic relationships in Hypomyces and allied genera, with emphasis on species growing on wood-decaying homobasidiomycetes. Canadian Journal of Botany 77: 1756–1768.
Punithalingam, E. 2003. Nuclei, micronuclei and appendages in tri- and tetraradiate conidia of Cornutispora and four other coelomycete genera. Mycological Research 107: 917–948.
Rambaut, A. 2009. FigTree version 1.3.1 [computer program]
Réblová, M., Untereiner, W. A., Štěpánek, V. & Gams, W. 2016. Disentangling Phialophora section Catenulatae: disposition of taxa with pigmented conidiophores and recognition of a new subclass, Sclerococcomycetidae (Eurotiomycetes). Mycological Progress 16: 27–46.
Rehner, S. A. & Samuels, G. J. 1995. Molecular systematics of the Hypocreales: a teleomorph gene phylogeny and the status of their anamorphs. Canadian Journal of Botany 73: 816–823.
Rehner, S. A., Minnis, A. M., Sung, G.-H., Luangsa-ard, J. J., Devotto, L. & Humber, R. A. 2011. Phylogeny and systematic of the anamorphic, entomopathogenic genus Beauveria. Mycologia 103: 1055–1073.
Resl, P., Fernández-Mendoza, F., Mayrhofer, H., & Spribille, T. 2010. The evolution of fungal substrate specificity in a widespread group of crustose lichens. Proceedings of the Royal Society B: Biological Sciences 285: 20180640.
Rodriguez, F., Oliver, J. L., Marin, A. & Medina, J. R. 1990. The general stochastic model of nucleotide substitution. Journal of Theoretical Biology 142: 485–501.
Rodriguez-Flakus, P & Printzen, C. 2014. Palicella, a new genus of lichenized fungi and its phylogenetic position within Lecanoraceae. The Lichenologist 46: 535–552.
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M. A., Huelsenbeck, J. P. 2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542.
Rossman, A.Y. 1979. A preliminary account of the taxa described in Calonectria. Mycotaxon 8: 485–558.
Rossman, A. Y. 1983. The phragmosporous species of Nectria and related genera. Mycological Papers 150: 1–164.
Rossman, A. Y., Samuels, G. J., Rogerson, C. T. & Lowen, R. 1999. Genera of Bionectriaceae, Hypocreaceae and Nectriaceae (Hypocreales, Ascomycetes). Studies in Mycology 42: 1–248.
Rossman, A.Y., Mckemy, J. M., Pardo-Schultheiss, R. A. & Schroers, H.-J. 2001. Molecular studies of the Bionectriaceae using large subunit rDNA sequences. Mycologia 93: 100–110.
Roux, C. & Triebel, D. 1994. Révision des espèces de Stigmidium et de Sphaerellothecium (champignons lichénicoles non lichénisés, Ascomycetes) correspondant à Pharcidia epicymatia sensu Keissler ou à Stigmidium schaereri auct. Bulletin de la Société Linéenne de Provence 45: 451–542.
Samuels, G. J., Müller, E. & Petrini, O. 1987. Studies in the Arnphisphaeriaceae (sensu lato) 3. New species of Monographella and Pestalosphaeria, and two new genera. Mycotaxon 28: 473–499.
Santesson, R., Moberg, R., Nordin, A., Tønsberg, T. & Vitikainen, O. 2004. Lichen-forming and Lichenicolous Fungi of Fennoscandia. Museum of Evolution, Uppsala University, Uppsala, Sweden.
Schoch, C. L., Seifert, K. A., Huhndorf, S., Robert, V., Spouge, J. L., Levesque, C. A., Chenb, W. & Fungal Barcoding Consortium. 2012. Nuclear ribosomal internal transcribed spacer (ITS) region as a universal DNA barcode marker for Fungi. Proceedings of the National Academy of Sciences 109: 6241–6246.
Schroers, H.J. 2001. A monograph of Bionectria (Ascomycota, Hypocreales, Bionectriaceae) and its Clonostachys anamorphs. Studies in Mycology 46: 1–214.
Sherwood-Pike, M. A. 1987. The ostropalean fungi III: The Odontotremataceae. Mycotaxon 28: 137–177.
Shoemaker, R. A. & Hambleton, S. 2001. ‘Helminthosporium’ asterinum, Polydesmus elegans, Imimyces, and allies. Canadian Journal of Botany 79: 592–599.
Seifert, K. A., McMullen, C. R., Yee, D., Reeleder, R. D. & Dobinson, K. F. 2003. Molecular differentiation and detection of ginseng adapted isolates of the root rot fungus Cylindrocarpon destructans. Phytopathology 93:1533–1542.
Spatafora, J. W. & Blackwell, M. 1993. Molecular systematics of unitunicate perithecial ascomycetes: the Clavicipitales-Hypocreales connection. Mycologia 85: 912–922.
Spooner, B. M. & Kirk, P. M. 1990. Observations on some genera of Trichothyriaceae. Mycological Research 94: 223–230.
Spribille, T., Pérez-Ortega, S., Tønsberg, T. & Schirokauer, D. 2010. Lichens and lichenicolous fungi of the Klondike Gold Rush National Historic Park, Alaska, in a global biodiversity context. The Bryologist 113: 439–515.
Spribille, T., Resl, P., Ahti, T., Pérez-Ortega, S., Mayrhofer, H. & Lumbsch, H.T. 2014. Molecular systematics of the wood-inhabiting, lichen-forming genus Xylographa (Baeomycetales, Ostropomycetidae) with eight new species. Symbolae Botanicae Upsalienses 37: 1–87.
Stamatakis, A. 2006. RAxML-VI-HPC: Maximum likelihoodbased phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22: 2688–2690.
Stamatakis, A., Hoover, P. & Rougemont, J. 2008. A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology 57: 758–771.
Stenroos, S., Laukka, T., Huhtinen, S., Dobbeler, P., Myllys L., Syrjänen, K. & Hyvönen, J. 2010. Multiple origins of symbioses between ascomycetes and bryophytes suggested by a five-gene phylogeny. Cladistics 26: 281–300.
Suh, S-O. & Blackwell, M. 1999. Molecular phylogeny of the cleistothecial fungi placed in Cephalothecaceae and Pseudeurotiaceae. Mycologia 91: 836–848.
Suija, A., Pérez-Ortega, S. & Hawksworth, D. L. 2011. Abrothallus halei (Ascomycota, incertae sedis), a new lichenicolous fungus on Lobaria species in Europe and North America. The Lichenologist 43: 51–55.
Suija, A., Ertz, D., Lawrey, J. D. & Diederich, P. 2015a. Multiple origin of the lichenicolous life habit in Helotiales, based on nuclear ribosomal sequences. Fungal Diversity 70: 55–72.
Suija, A., de los Ríos, A. & Pérez-Ortega, S. 2015b. A molecular reappraisal of Abrothallus species growing on lichens of the order Peltigerales. Phytotaxa 195: 201–226.
Suija, A, van den Boom, P., Zimmermann, E., Zhurbenko, M. P., Diederich, P. 2017 Lichenicolous species of Hainesia belong to Phacidiales (Leotiomycetes) and are included in an extended concept of Epithamnolia. Mycologia 109: 882–899.
Suija, A., Kaasalainen, U., Kirika, P. M. & Rikkinen, J. 2018.Taitaia, a novel lichenicolous fungus in tropical montane forests in Kenya (East Africa). The Lichenologist 50: 173–184.
Summerbell, R. C., Gueidan, C., Schroers, H-J., de Hoog, G. S., Starink, M., Arocha Rosete, Y., Guarro, J. & Scott, J. A. 2011. Acremonium phylogenetic overview and revision of Gliomastix, Sarocladium, and Trichothecium. Studies in Mycology 68: 139–162.
Sung, G. H., Hywell-Jones, N. L., Sung, J. M., Luangsa-ard, J. J., Shrestha, B. & Spatafora, J. W. 2007. Phylogenetic classification of Cordyceps and the clavicipitaceous fungi. Studies in Mycology 57: 5–59.
Thomas, D. C., Vandegrift, R., Ludden, A., Carroll, G. C. & Roy, B. A. 2016 Spatial ecology of the fungal genus Xylaria in a tropical cloud forest. Biotropica 48: 381–393.
Tretiach, M. 2002. Niesslia robusta, a new lichenicolous fungus on Tephromela grumosa from Tuscany, Italy. Nova Hedwigia 75: 357–365.
U’Ren, J. M., Miadlikowska, J., Zimmerman, N. B., Lutzoni, F., Stajich, J. E. & Arnold, A. E. 2016. Contributions of North American endophytes to the phylogeny, ecology, and taxonomy of Xylariaceae (Sordariomycetes, Ascomycota). Molecular Phylogenetics and Evolution 98: 210–232.
Voglmayr, H. & Jaklitsch, W. M. 2019. Stilbocrea walteri sp. nov., an unusual species of Bionectriaceae. Mycological Progress 18: 91–105.
Vouaux, L. 1913. Synopsis des champignons parasites de lichens. Bulletin de la Société Mycologique de France 29: 447–494.
Wang, Z., Binder, M., Schoch, C. L., Johnston, P. R., Spatafora, J. W. & Hibbett, D. S. 2006. Evolution of helotialean fungi (Leotiomycetes, Pezizomycotina): A nuclear rDNA phylogeny. Molecular Phylogenetics and Evolution 41 : 295–312.
Wei, X. L., Schmitt, I., Hodkinson, B. P., Flakus, A., Kukwa, M., Divakar, P. K., Kirika, P. M., Otte, J., Meiser, A. & Lumbsch, H. T. 2017. Circumscription of the genus Lepra, a recently resurrected genus to accommodate the ‘Variolaria’ – group of Pertusaria sensu lato (Pertusariales, Ascomycota). PloS ONE 12(7): e0180284.
Wu, W.P. & Zhuang, W.Y. 2005. Sporidesmium, Endophragmiella and related genera from China. Fungal Diversity Research Series 15: 1–531.
Zahlbruckner, A. 1907. Ascolichens (Schlauchflechten; Hymenolichens (Basidiomycetenflechten). In: Engler, A. & Prantl, K. (eds), Die Natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten, insbesondere den Nutzpflanzen, unter Mitwirkung zahlreicher hervorragender Fachgelehrten begründet. I Teil, 1Abteilung*: Lichens (Flechten), pp. 49–249. Engelmann, Leipzig.
Zhang, N. & Blackwell, M. 2002. Molecular phylogeny of Melanospora and similar pyrenomycetous fungi. Mycological Research 106: 148–155.
Zhang, Y., Zhang, J., Wang, Z., Fournier, J., Crous, P. W., Zhang, X., Li, W., Ariyawansa, H. A. & Hyde, K. D. 2014. Neotypification and phylogeny of Kalmusia. Phytotaxa 176: 164–173.
Zhurbenko, M. 2007. Corticifraga santessonii and C. chugachiana (Lecanoromycetes, Ascomycota), new species of lichenicolous fungi from the Holarctic. The Lichenologist 39: 221–226.
Zhurbenko, M. P. 2012. Lichenicolous fungi growing on Thamnolia, mainly from the Holarctic, with a worldwide key to the known species. The Lichenologist 44: 147–177.
Zhurbenko, M. P. & Etayo, J. 2012. Stigmidium lobariae, a new lichenicolous fungus from the Holarctic. Mycosphere 3: 62–64.
Zimmermann, E. & Berger, F. 2018. Beitrag zur lichenicolen Mycobiota Österreichs – Funde aus den Tiroler Alpen I. Herzogia 31: 732–762.
Contributions to the knowledge of lichenicolous fungi growing on baeomycetoid lichens and Icmadophila, with a key to the species
Mikhail Zhurbenko, Yoshihito Ohmura
The Lichenologist
Lichenicolous fungi from Vietnam, with the description of four new species
Mikhail Zhurbenko, Paul Diederich, Ludmila Gagarina
New lichenicolous fungi from Brazil, with a checklist of all lichenicolous fungi known from Brazil
Javier Etayo, André Aptroot, Silva da
The Bryologist
One Name – One Fungus: The Influence of Photosynthetic Partners on the Taxonomy and Systematics of Lichenized Fungi
Martin Kukwa, Magdalena Kosecka, Beata Guzow-Krzemińska
Acta Societatis Botanicorum Poloniae
Capronia harrisiana (Ascomycota, Chaetothyriales), a new lichenicolous species on Crocodia aurata from the southern Appalachian Mountains of southeastern North America
Jason Hollinger, James Lendemer
The Bryologist
Annotated Checklist of the Lichenicolous Fungi of Hungary
Nóra Varga, László Lőkös, Edit Farkas
Forecasting the number of species of asexually reproducing fungi (Ascomycota and Basidiomycota)
Nalin Wijayawardene, Alan Phillips, Diana Pereira, Dong-Qin Dai, André Aptroot, Josiane Monteiro, Irina Druzhinina, Feng Cai, Xinlei Fan, Laura Selbmann, Claudia Coleine, Rafael Castañeda-Ruiz, Martin Kukwa, Adam Flakus, Patricia Fiuza, Paul Kirk, Kunhiraman Kumar, Arachchi leperuma, Nakarin Suwannarach, Li-Zhou Tang, Teun Boekhout, Chen Tan, R. Jayasinghe, Marco Thines
Fungal Diversity
Beitrag zur Kenntnis der lichenicolen Mycobiota der Alpen I. – Weitere Funde aus Tirol und der Schweiz
Franz Berger, Erich Zimmermann
Phylogenetic revision of the lichenized family Gomphillaceae (Ascomycota: Graphidales) suggests post-K–Pg boundary diversification and phylogenetic signal in asexual reproductive structures
Xavier-Leite Barreto, Silva da, André Aptroot, Bibiana Moncada, Robert Lücking, Goto Tomio
Molecular Phylogenetics and Evolution
Endophytic fungus Pseudodidymocyrtis lobariellae KL27 promotes taxol biosynthesis and accumulation in Taxus chinensis
Xiaoying Cao, Lingxia Xu, Jingyi Wang, Mengmeng Dong, Chunyan Xu, Guoyin Kai, Wen Wan, Jihong Jiang
BMC Plant Biology
Studies on lichenicolous fungi in the Uppsala (UPS) collection curated by the late Rolf Santesson
Mikhail Zhurbenko
Lichenicole Pilze auf Thamnolia in den Alpen
Erich Zimmermann, Franz Berger
Two new species of Lichenotubeufia (Dothideomycetes, Tubeufiales) from Chile and New Zealand, with a revised key to the genus
Mikhail Zhurbenko
The Lichenologist