Abstract
Fungi that are barely lichenized or non-lichenized and closely related to lichenized taxa, the so-called borderline fungi, are an important element in reconstructing the evolutionary history of lichenized lineages. Arthoniaceae is a prime example including non-lichenized, saprotrophic lineages which potentially were precursors to lichenized taxa. In this study, we focused on saprotrophic species of Arthonia sensu lato, including new sequence data for Arthonia pinastri. We obtained fresh material of this taxon from a living branch of Fraxinus ornus in Italy to assess its taxonomic status and to elucidate its phylogenetic relationships within Arthonia. Thin sections of the thallus and ascomata of A. pinastri confirmed the absence of a photobiont. Maximum likelihood and Bayesian analyses of combined mtSSU, nuLSU and RPB2 sequence data placed the species close to A. dispersa (barely lichenized or non-lichenized) and A. punctiformis (non-lichenized) in a clade closely related to Arthonia sensu stricto, and the A. pinastri clade is here resurrected under the name Naevia. Ancestral character state analysis within a broader context of Arthoniales does not support the saprotrophic lifestyle to be a plesiomorphic feature, but suggests loss of lichenization in Naevia, as well as loss and possible regain in a second clade containing saprotrophic species and including taxa resembling Mycoporum, underlining the evolutionary plasticity of Arthoniales. These two clades constitute model taxa to further investigate the evolution of alternative biological lifestyles within the context of chiefly lichenized taxa.
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References
Acharius E (1803) Methodus qua omnes detectos lichenes. Marquard, Stockholm
Acharius E (1806) Arthonia, novum genus Lichenum, quod descripsit Erik Acharius, M.D. et Profess. etc. Neues J Bot 1:1–23
Acharius E (1808) Förteckning på de i Sverige växande arter af lafvarnas familj. Kongl Vetensk Acad Nya Handl 29:125–132
Almquist S (1880) Monographia Arthoniarum Scandinaviae. Kongl Svenska Vetensk-Akad Handl 17:1–69
Ariyawansa HA, Hyde KD, Jayasiri SC, Buyck B, Chethana KWT, Dai DQ, Dai YC, Daranagama DA, Jayawardena RS, Lücking R, Ghobad-Nejhad M, Niskanen T, Thambugala KM, Voigt K, Zhao RL, Li GJ, Doilom M, Boonmee S, Yang ZL, Cai Q, Cui YY, Bahkali AH, Chen J, Cui BK, Chen JJ, Dayarathne MC, Dissanayake AJ, Ekanayaka AH, Hashimoto A, Hongsanan S, Jones EBG, Larsson E, Li WJ, Li QR, Liu JK, Luo ZL, Maharachchikumbura SSN, Mapook A, McKenzie EHC, Norphanphoun C, Konta S, Pang KL, Perera RH, Phookamsak R, Phukhamsakda C, Pinruan U, Randrianjohany E, Singtripop C, Tanaka K, Tian CM, Tibpromma S, Abdel-Wahab MA, Wanasinghe DN, Wijayawardene NN, Zhang JF, Zhang H, Abdel-Aziz FA, Wedin M, Westberg M, Ammirat JF, Bulgakov TS, Lima DX, Callaghan TM, Callac P, Chang CH, Coca LF, Dal Forno M, Dollhofer V, Fliegerová K, Greiner K, Griffith GW, Ho HM, Hofstetter V, Jeewon R, Kang JC, Wen TC, Kirk PM, Kytövuori I, Lawrey JD, Xing J, Li H, Liu ZY, Liu XZ, Liimatainen K, Lumbsch HT, Matsumura M, Moncada B, Nuankaew S, Parnmen S, Santiago ALCMA, Sommai S, Song Y, Souza CAF, Souza-Motta CM, Su HY, Suetrong S, Wang Y, Wei SF, Yuan HS, Zhou LW, Réblová M, Fournier J, Camporesi E, Luangsa-ard JJ, Tasanathai K, Khonsanit A, Thanakitpipattana D, Somrithipol S, Diederich P, Millanes AM, Common RS, Stadler M, Yan JY, Li XH, Lee HW, Nguyen TTT, Lee HB, Battistin E, Marsico O, Vizzini A, Vila J, Ercole E, Eberhardt U, Simonini G, Wen HA, Chen ZH (2015) Fungal diversity notes 111–252: taxonomic and phylogenetic contributions to fungal taxa. Fungal Divers 75:27–274
Beltramini de Casati F (1858) Lichenografia Bassanese enumerati e descriti. Roberti, Bassano
de Lamarck JB, de Candolle AP (1805) Flore française, 3rd edn, Tome 2. de l’Imprimerie de Stouppe, Paris, 600 pp
Diederich P, Lawrey JD, Ertz D (2018) The 2018 classification and checklist of lichenicolous fungi, with 2000 non-lichenized, obligately lichenicolous taxa. Bryologist 121:340–426
Divakar PK, Kauff F, Crespo A, Leavitt SD, Lumbsch HT (2013) Understanding phenotypical character evolution in parmelioid lichenized fungi (Parmeliaceae, Ascomycota). PLoS One 8(11):e83115
Drummond AJ, Suchard MA, Xie D, Rambaut A (2012) Bayesian phylogenetics with BEAUti and the BEAST 1.7. Mol Biol Evol 29:1969–1973
Dufour L (1818) Revision du genre Opégraphe de la Flora Francaise. J. Phys Chim Hist Nat 87:200–225
Engler A (1903) Syllabus der Pflanzenfamilien. Borntraeger, Berlin
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 focusing on the genus Opegrapha. Mycol Res 113:141–152
Ertz D, Sanderson N, Łubek A, Kukwa M (2018) Two new species of Arthoniaceae from old-growth European forests, Arthonia thoriana and Inoderma sorediatum, and a new genus for Schismatomma niveum. Lichenologist 50:161–172
Flotow J (1849) Dr. Rabenhorst’s Lichenes Italici. Die Lichenen, welche der Dr. L. Rabenhorst auf seiner Reise durch die östlichen und südlichen Provinzen Italiens im Jahr 1847 gesammelt hat. Linnaea 6:353–382
Fries EM (1824) Schedulae Criticae de Lichenibus Exsiccatis Sueciae. Berlingiana, Londini Gothorum
Fries EM (1825) Systema Orbis Vegetabilis. Typographia Academica, Lund
Fries EM (1849) Summa Vegetabilium Scandinaviae. Holmiae et Lipsiae, Berlin
Frisch A, Holien H (2018) Arthonia toensbergii, a new lichenicolous fungus on Mycoblastus affinis from the boreal rainforests in Norway. Graph Scr 30:34–43
Frisch A, Thor G, Ertz D, Grube M (2014) The Arthonialean challenge: restructuring Arthoniaceae. Taxon 63:727–744
Frisch A, Ohmura Y, Ertz D, Thor G (2015) Inoderma and related genera in Arthoniaceae with elevated white pruinose pycnidia or sporodochia. Lichenologist 47:233–256
Gargas A, DePriest PT, Grube M, Tehler A (1995) Multiple origins of lichen symbioses in fungi suggested by SSU rDNA phylogeny. Science 268:1492–1495
Gerstmans C, Ertz D (2016) Arthonia phlyctiformis Nyl. (Arthoniaceae) retrouvé en France après plus d’un siècle. Bull Inform Ass Fr Lichénologie 41:10–12
Glez-Peña D, Gómez-Blanco D, Reboiro-Jato M, Fdez-Riverola F, Posada D (2010) ALTER: program-oriented conversion of DNA and protein alignments. Nucleic Acids Res 38:14–18
Grube M (1998) Classification and phylogeny in the Arthoniales (lichenized Ascomycetes). Bryologist 101:377–391
Grube M (2001) Coniarthonia, a new genus of arthonioid lichens. Lichenologist 33:491–502
Grube M (2007) Arthonia. In: Nash TH III, Gries C, Bungartz F (eds) Lichen Flora of the greater sonoran desert region, vol 3. Lichens Unlimited, Arizona State University, Tempe, pp 39–61
Grube M, Giralt M (1996) Studies on some species of Arthothelium occurring in the western Mediterranean. Lichenologist 28(1):15–36
Grube M, Matzer M, Hafellner J (1995) A preliminary account of the lichenicolous Arthonia species with reddish, K + reactive pigments. Lichenologist 27:25–42
Hall TA (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucl Acids Symp Ser 41:95–98
Harris RC (1995) More Florida Lichens. Including the 10¢ Tour of the Pyrenolichens. R.C. Harris, Bronx
Hein P (1976) Revision der Gattung Laetinaevia Nannf (Ascomycetes) und Neuordnung der Naevioideae. Willdenowia Beiheft 9(3):5–136
Henssen A, Jahns H (1973) (“1974”) Lichenes. Georg Thieme, Stuttgart
Henssen A, Thor G (1994) Developmental morphology of the “Zwischengruppe” between ascohymeniales and ascoloculares. In: Hawksworth DL (ed) Ascomycete systematics. Problems and perspectives in the nineties. Plenum Press, New York, pp 43–56
Huelsenbeck JP, Ronquist F (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17:754–755
Hyde KD, Jones EBG, Liu JK, Ariyawansa H, Boehm E, Boonmee S, Braun U, Chomnunti P, Crous PW, Dai DQ, Diederich P, Dissanayake A, Doilom M, Doveri F, Hongsanan S, Jayawardena R, Lawrey JD, Li YM, Liu YX, Lücking R, Monka J, Muggia L, Nelsen MP, Pang KL, Phookamsak R, Senanayake IC, Shearer CA, Suetrong S, Tanaka K, Thambugala KM, Wijayawardene NN, Wikee S, Wu HX, Zhang Y, Begoña AH, Alias SA, Aptroot A, Bahkali AH, Bezerra JL, Bhat DJ, Camporesi E, Chukea E, Gueidan C, Hawksworth DL, Hirayama K, Hoog SD, Kang JK, Knudsen K, Li WJ, Li XH, Liu ZY, Mapook A, Mckenzie EHC, Miller AN, Mortimer PE, Phillips AJL, Raja HA, Scheuer C, Schumm F, Taylor JE, Tian Q, Tibpromma S, Wanasinghe DN, Wang Y, Xu JC, Yacharoen S, Yan JY, Zang M (2013) Families of Dothideomycetes. Fungal Divers 63:1–313
Index Fungorum (2020) http://www.indexfungorum.org/Names/Names.asp. Accessed 2 Jan 2020
Ismail SI, Batzer JC, Harrington TC, Crous PW, Lavrov DV, Li H, Gleason ML (2016) Ancestral state reconstruction infers phytopathogenic origins of sooty blotch and flyspeck fungi on apple. Mycologia 108(2):292–302
Kantvilas G, Wedin M (2015) Lichenicolous species of the Ascomycete genus Arthonia Ach. from Kangaroo Island. J Adelaide Bot Gard 29:1–6
Jayasiri SC, Hyde KD, Abd-Elsalam KA, Abdel-Wahab MA, Ariyawansa HA, Bhat J, Buyck B, Dai YC, Ertz D, Hidayat I, Jeewon R, Jones EBG, Karunarathna SC, Kirk P, Lei C, Liu JK, Maharachchikumbura SSN, McKenzie E, Ghobad-Nejhad M, Nilsson H, Pang KL, Phookamsak R, Rollins AW, Romero AI, Stephenson S, Suetrong S, Tsui CKM, Vizzini A, Wen TC, de Silva NI, Promputtha I, Kang JC (2015) The Facesoffungi database: fungal names linked with morphology, molecular and human attributes. Fungal Divers 74:3–18
Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Mol Biol Evol 30:772–780
Kirschstein W (1935) Neue und seltene Ascomyceten. Ann Mycol 33:202–229
Lee BG, Hur JS (2016) Three new species and nine new records in the genus Arthonia from South Korea. Mycobiology 44:202–216
Lendemer JC, Ray D (2017) Two new pinicolous Arthonia (Arthoniaceae; Arthoniomycetes) from the Delmarva Peninsula of the Atlantic Coastal Plain in eastern North America. The Bryologist 120(1):11–18
Lendemer JC, Harris RC, Tripp EA (2013) The lichens and allied fungi of Great Smoky Mountains National Park: an annotated checklist with comprehensive keys. Mem New York Bot Gard 104:1–152
Liu Y, Whelen S, Hall B (1999) Phylogenetic relationships among ascomycetes: evidence from an RNA polymerase II subunit. Mol Biol Evol 16:1799–1808
Liu JK, Hyde KD, Jeewon R, Phillips AJ, Maharachchikumbura SS, Ryberg M, Liu ZY, Zhao Q (2017) Ranking higher taxa using divergence times: a case study in Dothideomycetes. Fungal Divers 84:75–99
Lücking R, Nelsen MP (2018) On Ediacarans, protolichens, and lichen-derived Penicillium: a critical reassessment of the evolution of lichenization in fungi. In: Krings M, Harper CJ, Cuneo NR, Rothwell GW (eds) Transformative paleobotany. Academic Press, San Diego, pp 551–590
Lücking R, Hodkinson BP, Leavitt SD (2017) The 2016 classification of lichenized fungi in the Ascomycota and Basidiomycota—approaching one thousand genera. Bryologist 119:361–416
Lumbsch HT (1999) The ascoma development in Mycoporum elabens (Mycoporaceae, Dothideales). Plant Biol 1:321–326
Lutzoni F, Pagel M, Reeb V (2001) Major fungal lineages are derived from lichen symbiotic ancestors. Nature 411(6840):937–940
Lutzoni F, Kauff F, Cox CJ, McLaughlin D, Celio G, Dentinger B, Padamsee M, Hibbett D, James TY, Baloch E, Grube M (2004) Assembling the fungal tree of life: progress, classification, and evolution of subcellular traits. Am J Bot 91:1446–1480
Marchand NL (1896) Énumération méthodique et raisonnée des familles et des genres de la classe des mycophytes (champignons et lichens). Société d’Éditions Scientifiques, Paris
Massalongo AB (1852) Ricerche sull’autonomia dei licheni crostosi e materiali pella loro naturale ordinazione. Frizerio, Verona
Massalongo AB (1855) Frammenti Lichenografici. Ramanzini, Verona
Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES science gateway for inference of large phylogenetic trees. In: Proceedings of the gateway computing environments workshop (GCE), 14 Nov. 2010, New Orleans, LA, pp 1–8
Miranda-González R, Aptroot A, Lücking R, Flakus A, Barcenas-Peña A, de los Ángeles Herrera-Campos M (2020) The identity, ecology and distribution of Polypyrenula (Ascomycota: Dothideomycetes): a new member of Trypetheliaceae revealed by molecular and anatomical data. Lichenologist 52(1):27–35
Moen VS (2019) Molecular systematics and species delimitation in Coniocarpon and Arthonia punctiformis s.lat. in Norway. Masters thesis, Norwegian University of Science and Technology
Muggia L, Baloch E, Stabentheiner E, Grube M, Wedin M (2011) Photobiont association and genetic diversity of the optionally lichenized fungus Schizoxylon albescens. FEMS Microbiol Ecol 75:255–272
Müller J (1895) Sertum Australiense s. species novae Australienses Thelotremearum, Graphidearum et Pyrenocarpearum. Bull Herb Boissier 3:313–327
Nelsen MP, Lücking R, Grube M, Mbatchou JS, Muggia L, Plata ER, Lumbsch HT (2009) Unravelling the phylogenetic relationships of lichenised fungi in Dothideomyceta. Stud Mycol 64:135–144
Nelsen MP, Lücking R, Mbatchou JS, Andrew CJ, Spielmann AA, Lumbsch HT (2011) New insights into relationships of lichen-forming Dothideomycetes. Fungal Divers 51:155–162
Nimis PL (2016) The Lichens of Italy—a second annotated catalogue. EUT Edizioni Università di Trieste, Trieste
Nylander JAA (2004) MrModeltest v2.2 program distributed by the author. Evolutionary Biology Centre, Uppsala University, Uppsala
Nylander W (1858) Synopsis Methodica Lichenum, vol 1. Martinet, Paris
Nylander W (1861) Lichenes Scandinaviae (sive Prodromus Lichenographiae Scandinaviae). Not Sällsk Fauna Fl Fenn Förhandl 5:1–313
Park JS, Park SY, Park CH, Kondratyuk SY, Oh SO, Hur JS (2017) Taxonomic revision of the lichen genera Pertusaria, Varicellaria, and Variolaria (Pertusariales, Ascomycota) in South Korea. Mycobiology 45:270–285
Petrak F (1941) Beiträge zur Kenntnis der orientalischen Pilzflora. Ann Naturhist Mus Wien 52:301–396
Petrak F (1953) Neue Beitrage zur Pilzflora der Türkei. Sydowia 7:14–44
Rabenhorst L (1870) Kryptogamen-Flora von Sachsen, der Oberlausitz, Thüringen und Nordböhmen mit Berücksichtigung der benachbarten Länder, vol 2. Abt. Die Flechten. Kummer, Leipzig
Rambaut A (2014) FigTree v1.4: tree figure drawing tool. http://treebio.ed.ac.uk/software/figtree
Rambaut A, Drummond AJ (2009) Tracer. MCMC trace analysis tool version v1.5.0. http://tree.bio.ed.ac.uk/software/tracer/
Rehm H (1896) Die Pilze Deutschlands, Oesterreichs und der Schweiz. III. Abteilung: Ascomyceten: Hysteriaceen und Discomyceten. Dr. L. Rabenhorst’s Kleine Kryptogamenflora von Deutschland, Oesterreich und der Schweiz, Erster Band: Pilze, Zweite Auflage, Zweite edn. Kummer, Leipzig
Reinke J (1895) Abhandlungen über Flechten III. Jahrb wiss Bot 28:49–160
Roychowdhury KN (1979) Lichens of the Indian Botanic Garden, Howrah. Nelumbo 21:163–167
Schneider A (1897) Reinke’s discussions of lichenology.IV. Bull Torrey. Bot Club 24:237–243
Schrader HA (1797) Nova genera plantarum. Auctore Henrico Adolpho Schrader. Pars prima. Apud Siegfried Lebrecht Crusium, Lipsiae
Sernander R (1907) Om nagra former för art- och varietetsbildning hos lafvarna. Svensk Bot Tidskr 1(97–115):135–186
Smith CW, Aptroot A, Coppins BJ, Fletcher A, Gilbert OL, James PW, Wolseley PA (eds) (2009) The Lichens of Great Britain and Ireland. British Lichen Society, London
Stamatakis A (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30:1312–1313
Stamatakis A, Hoover P, Rougemont J (2008) A rapid bootstrap algorithm for the RAxML web servers. Syst Biol 57:758–771
Sundin R (1999) The genus Arthonia sect. Arthonia in Europe, Northern Africa and North America–A revision and phylogenetic analysis. Phylogenetic and taxonomic studies within Arthonia Ach. (Ascomycetes, Arthoniales). Botaniska Instiutionen, Stockholms Universitet, Stockholm
Sundin R, Tehler A (1998) Phylogenetic studies of the genus Arthonia. Lichenologist 30:381–413
Sundin R, Thor G, Frisch A (2012) A literature review of Arthonia s.lat. Biblioth. Lichenologica 108:257–290
Vainio EA (1901) Lichenes. In: Catalogue of the African Plants collected by Dr. Friedrich Welwitsch in 1853–61, Vol. 2, Part 2. Longmans & Co., London, pp 396–463
Van den Broeck D, Frisch A, Razafindrahaja T, Van de Vijver B, Ertz D (2018) Phylogenetic position of Synarthonia (lichenized Ascomycota, Arthoniaceae), with the description of six new species. Pl Ecol Evol 151:327–351
Vilgalys R, Hester M (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. J Bacteriol 172:4238–4246
von Arx JA (1954) Revision einiger Gattungen der Ascomyceten. Acta Bot Neerl 3:83–93
Wanasinghe DN, Phukhamsakda C, Hyde KD, Jeewon R, Lee HB, Jones EBG, Tibpromma S, Tennakoon DS, Dissanayake AJ, Jayasiri SC, Gafforov Y, Camporesi E, Bulgakov TS, Ekanayake AH, Perera RH, Samarakoon MC, Goonasekara ID, Mapook A, Li WJ, Senanayake IC, Li JF, Norphanphoun C, Doilom M, Bahkali AH, Xu JC, Mortimer PE, Tibell L, Tibell S, Karunarathna SC (2018) Fungal diversity notes 709–839: taxonomic and phylogenetic contributions to fungal taxa with an emphasis on fungi on Rosaceae. Fungal Divers 89:1–236
Wedin M, Hafellner J (1998) Lichenicolous species of Arthonia on Lobariaceae with notes on excluded taxa. Lichenologist 30:59–91
Wedin M, Döring H, Gilenstam G (2004) Saprotrophy and lichenization as options for the same fungal species on different substrata: environmental plasticity and fungal lifestyles in the Stictis-Conotrema complex. New Phytol 4:459–465
Wijayawardene NN, Hyde KD, Al-Ani LKT, Tedersoo L, Haelewaters D, Rajeshkumar KC, Zhao RL, Aptroot A, Leontyev DV, Saxena RK, Tokarev YS, Dai DQ, Letcher PM, Stephenson SL, Ertz D, Lumbsch HT, Kukwa M, Issi IV, Madrid H, Phillips AJL, Selbmann L, Pfliegler WP, Horváth E, Bensch K, Kirk PM, Kolaříková K, Raja HA, Radek R, Papp V, Dima B, Ma J, Malosso E, Takamatsu S, Rambold G, Gannibal PB, Triebel D, Gautam AK, Avasthi S, Suetrong S, Timdal E, Fryar SC, Delgado G, Réblová M, Doilom M, Dolatabadi S, Pawłowska JZ, Humber RA, Kodsueb R, Sánchez-Castro I, Goto BT, Silva DKA, de Souza FA, Oehl F, da Silva GA, Silva IR, Błaszkowski J, Jobim K, Maia LC, Barbosa FR, Fiuza PO, Divakar PK, Shenoy BD, Castañeda-Ruiz RF, Somrithipol S, Lateef AA, Karunarathna SC, Tibpromma S, Mortimer PE, Wanasinghe DN, Phookamsak R, Xu J, Wang Y, Tian F, Alvarado P, Li DW, Kušan I, Matočec N, Mešić A, Tkalčec Z, Maharachchikumbura SSN, Papizadeh M, Heredia G, Wartchow F, Bakhshi M, Boehm E, Youssef N, Hustad VP, Lawrey JD, Santiago ALCMA, Bezerra JDP, Souza-Motta CM, Firmino AL, Tian Q, Houbraken J, Hongsanan S, Tanaka K, Dissanayake AJ, Monteiro JS, Grossart HP, Suija A, Weerakoon G, Etayo J, Tsurykau A, Vázquez V, Mungai P, Damm U, Li QR, Zhang H, Boonmee S, Lu YZ, Becerra AG, Kendrick B, Brearley FQ, Motiejūnaitė J, Sharma B, Khare R, Gaikwad S, Wijesundara DSA, Tang LZ, He MQ, Flakus A, Rodriguez-Flakus P, Zhurbenko MP, McKenzie EHC, Stadler M, Bhat DJ, Liu JK, Raza M, Jeewon R, Nassonova ES, Prieto M, Jayalal RGU, Erdoğdu M, Yurkov A, Schnittler M, Shchepin ON, Novozhilov YK, Silva-Filho AGS, Gentekaki E, Liu P, Cavender JC, Kang Y, Mohammad S, Zhang LF, Xu RF, Li YM, Dayarathne MC, Ekanayaka AH, Wen TC, Deng CY, Pereira OL, Navathe S, Hawksworth DL, Fan XL, Dissanayake LS, Kuhnert E, Grossart HP, Thines M (2020) Outline of Fungi and fungus-like taxa. Mycosphere 11(1):1060–1456
Wijayawardene NN, Hyde KD, Rajeshkumar KC, Hawksworth DL, Madrid H, Kirk PM, Braun U, Singh RV, Crous PW, Kukwa M, Lücking R, Kurtzman CP, Yurkov A, Haelewaters D, Aptroot A, Lumbsch HT, Timdal E, Ertz D, Etayo J, Phillips AJL, Groenewald JZ, Papizadeh M, Selbmann L, Dayarathne MC, Weerakoon G, Jones EBG, Suetrong S, Tian Q, Castanéda-Ruiz RF, Bahkali AH, Pang K-L, Tanaka K, Dai DQ, Sakayaroj J, Hujslová M, Lombard L, Shenoy BD, Suija A, Maharachchikumbura SSN, Thambugala KM, Wanasinghe DN, Sharma BO, Gaikwad S, Pandit G, Zucconi L, Onofri S, Egidi E, Raja HA, Kodsueb R, Cáceres MES, Pérez-Ortega S, Fiuza PO, Monteiro JS, Vasilyeva LN, Shivas RG, Prieto M, Wedin M, Olariaga I, Lateef AA, Agrawal Y, Fazeli SAS, Amoozegar MA, Zhao GZ, Pfliegler WP, Sharma G, Oset M, Abdel-Wahab MA, Takamatsu S, Bensch K, de Silva NI, De Kese A, Karunarathna A, Boonmee S, Pfister DH, Lu Y-Z, Luo Z-L, Boonyuen N, Daranagama DA, Senanayake IC, Jayasiri SC, Samarakoon MC, Zeng X-Y, Doilom M, Quijada L, Rampadarath S, Heredia G, Dissanayake AJ, Jayawardana RS, Perera RH, Tang LZ, Phukhamsakda C, Hernández-Restrepo M, Ma X, Tibpromma S, Gusmao LFP, Weerahewa D, Karunarathna SC (2017) Notes for genera—Ascomycota. Fungal Divers 86:1–594
Woo JJ, Lőkös L, Farkas E, Park CH, Hur JS (2017) Cryptothecia austrocoreana (Arthoniales, Arthoniaceae), a new species from South Korea. Mycobiol 45:338–343
Yu Y, Harris AJ, Blair C, He X (2015) RASP (Reconstruct Ancestral State in Phylogenies): a tool for historical biogeography. Mol Phyl Evol 87:46–49
Yu Y, Blair C, He X (2019) RASP 4: Ancestral state reconstruction tool for multiple genes and characters. Mol Biol Evol 37:604–606
Zahlbruckner A (1907) Specieller Teil. In: Engler A, Prantl K (eds) Die natürlichen Pflanzenfamilien, Teil 1, Abteilung, vol 1. Engelmann, Leipzig, pp 49–249
Zahlbruckner A (1923) Catalogus Lichenum Universalis 2. Borntraeger, Leipzig
Zhaxybayeva O, Gogarten JP (2002) Bootstrap, Bayesian probability and maximum likelihood mapping: exploring new tools for comparative genome analyses. BMC Genomics 3:4
Zoller S, Scheidegger C, Sperisen C (1999) PCR primers for the amplification of mitochondrial small subunit ribosomal DNA of lichen-forming ascomycetes. Lichenologist 31:511–516
Acknowledgements
We thank the Thailand Research Fund (“The future of specialist fungi in a changing climate: baseline data for generalist and specialist fungi associated with ants, Rhododendron species and Dracaena species DBG6080013” and “Impact of climate change on fungal diversity and biogeography in the Greater Mekong Sub-region RDG6130001”) and “The 2019 high-end foreign expert introduction plan to Kunming Institute of Botany (granted by the Ministry of Science and Technology of the People’s Republic of China (Grant Number G20190139006)) for funding this research. S.C. Karunarathna would like to thank the CAS President’s International Fellowship Initiative (PIFI) under the following grant: 2018PC0006 and the National Science Foundation of China (NSFC, project code 31851110759). We also thank Udeni Jayalal, Nalin Wijayawardene, Ming-Ying Zhang, Mohammed Warris, Lee B.G and Eleni Gentekaki for their support during this research. Dhanushka Wanasinghe would like to thank CAS President’s International Fellowship Initiative (PIFI) for funding his postdoctoral research (Number 2019PC0008), the National Science Foundation of China and the Chinese Academy of Sciences for financial support under the following Grants: 41761144055, 41771063 and Y4ZK111B01. We also extend our gratitude to Teuvo Ahti, David Hawksworth, Pier-Luigi Nimis and Rikard Sundin and Mats Wedin who helped to clarify the nomenclatural status of the genus Naevia.
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Thiyagaraja, V., Lücking, R., Ertz, D. et al. Evolution of non-lichenized, saprotrophic species of Arthonia (Ascomycota, Arthoniales) and resurrection of Naevia, with notes on Mycoporum. Fungal Diversity 102, 205–224 (2020). https://doi.org/10.1007/s13225-020-00451-9
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DOI: https://doi.org/10.1007/s13225-020-00451-9