Four new monotypic genera of the caloplacoid lichens (Teloschistaceae, lichen-forming ascomycetes)

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1 Four new monotypic genera of the caloplacoid lichens (Teloschistaceae, lichen-forming ascomycetes) Sergii Y. Kondratyuk 1, *, László Lőkös 2, Jung A. Kim 3, Anna S. Kondratiuk 3,4, Min Hye Jeong 3, Seol Hwa Jang 3, Soon-Ok Oh 3 and Jae-Seoun Hur 3 1 M. H. Kholodny Institute of Botany, Tereshchenkivska str. 2, Kiev, Ukraine 2 Department of Botany, Hungarian Natural History Museum, H-1476 Budapest, Pf. 222, Hungary 3 Korean Lichen Research Institute, Sunchon National University, Sunchon , Korea; E- mail: jshur1@sunchon.ac.kr 4 Institute of Biology, Scientific Educational Centre, Taras Shevchenko National University of Kiev, Volodymyrska str. 64/13, Kiev, Ukraine Running title: Four new genera of caloplacoid lichens *Corresponding author: ksya_net@ukr.net

2 Abstract Four robust monophyletic branches having the highest level of bootstrap support in the phylogenetic tree of the Teloschistaceae based on combines data set of ITS and LSU nr DNA and 12S SSU mt DNA sequences, which do not belong to any other earlier proposed genera of the subfamily Caloplacoideae, are described as new monotypic genera, i.e. the genus Fauriea S. Y. Kondr., L. Lőkös et J.-S. Hur gen. nov. for South Korean Caloplaca chujaensis, the genus Jasonhuria S. Y. Kondr., L. Lőkös et S.-O. Oh gen. nov. for Eastern Asian Caloplaca bogilana, the genus Loekoesia S. Y. Kondr., S.-O. Oh et J.-S. Hur gen. nov. for South Korean Caloplaca austrocoreana, as well as the genus Olegblumia S. Y. Kondr., L. Lőkös et J.- S. Hur gen. nov. for European and North American Caloplaca demissa. Legal descriptions, comparison with closely related taxa and discussion of their position are provided. Four new consequent combinations for type species of these genera are proposed. Fauriea chejuensis is for the first time recorded for China. Keywords Caloplacoideae, Fauriea, Jasonhuria, Loekoesia, Olegblumia Taxonomy of the Teloschistaceae has dramatically changed since 2012 based on molecular phylogeny [1 7]. Three subphyla, i.e. subfamilies Teloschistoideae, Caloplacoideae and Xanthorioideae proposed in years [3, 8] are recently added by new one [9]. Number of accepted genera of the Teloschistaceae evolved from 10 in Kärnefelt ([10]) to 29 [1] and to 67 [5 7]. New genera of the Teloschistoid and Xanthorioid lichens are also recently described [11 12]. The aim of this paper is to provide legal description for four robust monophyletic groups of the subfamily Caloplacoideae, which do not belong to any other earlier described genera of this subfamily. The new four genera described below are the following: genus Fauriea for the Eastern Asian species Caloplaca chujaensis; genus Jasonhuria for the Eastern Asian Caloplaca bogilana; genus Loekoesia for the South Korean Caloplaca austrocoreana; as well as genus

3 Olegblumia for the European and North American Caloplaca demissa. MATERUAIL AND METHODS Specimens were examined using standard microscopical techniques, i.e. hand-sectioned under Nikon SMZ-645 dissecting microscope (Nikon Corp., Tokyo, Japan), sections were observed under Nikon E-200 and Olympus BX-51 microscope (same as above). Spot test reactions were performed on thalli under a compound microscope. Chemicals were extracted in analytical grade acetone in a 1 ml Eppendorf tube. Thin layer chromatography (TLC) was performed using a glass plate coated with TLC Silica gel 60, in solvent system A (toluene : dioxin : acetic acid = 180:45:5) [13]. Total DNA was extracted directly from the thalli according to Ekman ([14]) and was purified with DNeasy Plant Mini Kit (QIAGEN, Germany). The nuclear ribosomal RNA gene region including the internal transcribed spacers 1 and 2 and the 5.8S subunit (ITS) was amplified using the primers ITS1F [15] and ITS4 [16], the 28S LSU using the primer LR5 [17], and the 12S mtssu using the primers mtssu1-mtssu3r and mtssu2r [2, 18]. The amplification was done using a Takara JP/TP600 PCR machine (Takara Bio Inc., Japan). One initial cycle of 5 min at 94 ºC was followed by 30 cycles of the following steps: 30 seconds at 94 ºC, 39 seconds at 57 ºC and 1 min at 72 ºC. Amplifications were ended with a final cycle at 72 ºC for 10 min. PCR products were then sent to the sequencing facilities of the Genotech Cooperation, Seoul, South Korea for cleaning and sequencing. The sequencing was carried out using the fluorescent marker BigDye and an ABI 3730xl sequencing machine (Applied Biosystems, Carlsbad, CA, USA). The consensus sequence was aligned with all related species sequences retrieved from the GenBank database. The consensus sequences were then deposited into GenBank under the accession numbers XXXXXXXX XXXXXXXX. Phylogenetic analysis was performed using

4 the ITS region and LSU gene of nrdna and 12S SSU mtdna sequences of the treated fungi retrieved from the GenBank database and the 5 lichen-forming fungi investigated in this study. Sequence alignment was conducted in BioEdit and a phylogenetic tree was generated by the maximum parsimony (MP), minimum evolution (ME), and maximum likelihood (ML) analysis methods performed in Mega 5.0 [19] with the number of bootstrap trials set to 1, sequences on nrdna and mtdna are for the first time submitted to GenBank for the following taxa: Fauriea chujaensis, Jasonhuria bogilana, and Loekoesia austrocoreana. RESULTS AND DISCUSSION Description of taxa Fauriea S.Y. Kondr., L. Lőkös et J.-S. Hur gen. nov. MycoBank No. MB XXXXX Thallus crustose, continuous; grey to whitish grey or lead-grey in places; apothecia seem to be lecanorine, immersed into thallus to semi-immersed, disc dark brown, true exciple scleroplectenchymatous to mesodermatous paraplectenchymatous, cortical layer paraplectenchymatous; ascospores bipolar. Conidiomata not seen. Thallus and epihymenium K ; anthraquinones absent. Type species: Fauriea chujaensis (S. Y. Kondr., L. Lőkös et Hur) S. Y. Kondr., L. Lőkös, J. Kim, A. S. Kondratiuk, S.-O. Oh et J.-S. Hur. Thallus crustose, continuous/entire (not areolate); grey to whitish grey or lead-grey in places; with of Lecanora type apothecia with brown or dark brown disc. Hypothallus black. Apothecia at first immersed into thallus, later semi-immersed or seen in thalline warts, seem to be lecanorine, thalline margin whitish grey, disc dark brown, true margin if developed entire, light brown or transparent brown; in section true exciple scleroplectenchymatous to mesodermatous paraplectenchymatous; cortical layer of thalline exciple paraplectenchymatous;

5 paraphyses broom-like branched, the uppermost portions becoming brownish; asci 8-spored; bipolar ascospores narrowly ellipsoid, slightly widened at the septum; ascospore septum of medium width. Conidiomata not seen. Chemistry: thallus and epihymenium K ; anthraquinones absent. Ecology: On siliceous rocks in the coastal zone, often growing together with other members of the genus Caloplaca (C. yeosuensis S. Y. Kondr et J.-S. Hur, C. aff. diffluens (Hue) Zahlbr., C. sideritis (Tuck.) Zahlbr., as well as members of the genera Lecanora, Ramalina, Phaeophyscia, etc. Distribution: So far it is known from several collections from islands and coastal part of mainland of South Korea and China. It is for the first time recorded from China here. Etymology: It is named after the known French botanist Urbain Jean Faurie ( ), who provided important collections of lichens, mosses and vascular plants of the Eastern Asian region. Taxonomic notes: The genus Fauriea is positioned in one clade with the genus Rufoplaca Arup, Søchting et Frödén and the Caloplaca furax group. However, this clade has very weak support, while the two mentioned genera and the Caloplaca furax group have shown very high level of bootstrap support, as after ITS nrdna, mtdna sequences, as well as after combined data set (Fig. 1). Macroscopically the representatives of the genus Fauriea is similar to some species of the genus Pyrenodesmia, i.e. P. variabilis (Pers.) A. Massal., a Northern Hemisphere lichen growing on calcareous rocks. However, Fauriea chujaensis differs from Pyrenodesmia variabilis in having continuous/entire thallus, in having smaller apothecia, in having narrower ascospores, and in having much wider ascospore septum, as well as in the lack of the white pruina on apothecium discs and in the lack of K+ and C+ reactions of thallus and epihymenium (see also [20]). Unfortunately there are hitherto no molecular data on Caloplaca atroalba (Tuck.)

6 Zahlbr., a North American lichen growing on non-calcareous or calcareous rocks in having crustose grey thallus without asexual propagules and brown apothecium discs, to which Fauriea chujaensis is similar. However, Fauriea chujaensis differs from Caloplaca atroalba in having only grey and continuous/entire crustose thallus, in having smaller apothecia, and in having smaller and especially much narrower ascospores, and in having much wider ascospore septum, as well as in the lack of K+ and C+ reactions of thallus and epihymenium. After having mainly immersed or semi-immersed apothecia Fauriea chujaensis may resemble some species of the genus Aspicilia A. Massal., but microscopical study of ascospores easily confirms its position in the family Teloschistaceae. Fauriea chujaensis is similar to Caloplaca sideritis with which it sometimes grows side by side, in having similar greyish crustose thallus and lecanorine apothecia with dark grey or grey thalline margins, but it differs from C. sideritis in the lack of yellowish apothecium disc and in the lack of K+ and C+ reaction of epithecium and thalline cortical layer. From species of the genus Lecanora Ach. with which Fauriea chujaensis often growing side by side it differs in having immersed to semi-immersed (not constricted at the basis when uplifted in thalline warts) apothecia, in having black hypothallus, as well as in having bipolar ascospores. Jasonhuria S. Y. Kondr., L. Lőkös et S.-O. Oh gen. nov. MycoBank No. MB XXXXX Thallus saxicolous, crustose, grey to greyish white; cortex paraplectenchymatous. Apothecia biatorine to lecanorine; disc orange-brown to brownish red or rust-red; thalline margin concolorous with the thallus; proper margin black, true exciple paraplectenchymatous, outer region aeruginose pigmented. Conidia ellipsoid. Constituents: atranorin, gyrophoric and lecanoric acids (major compounds), parietin (traces). Type species: Jasonhuria bogilana (Y. Joshi et Hur) S. Y. Kondr., L. Lőkös, J. Kim, A. S. Kondratiuk et S.-O. Oh.

7 Thallus saxicolous, crustose, areolate to cracked areolate, grey to greyish white. Cortex paraplectenchymatous, necral layer absent. Apothecia biatorine to lecanorine, adnate to sessile; disc orange-brown to brownish red to rust-red, plane to convex, epruinose; thalline margin concolorous with the thallus; proper margin black, Hymenium hyaline, hypothecium hyaline, without oil-droplets; true exciple paraplectenchymatous, outer region ±aeruginose pigmented. Paraphyses thin, with a few swollen cells at the top. Asci 8-spored, ascospores polarilocular, ellipsoid, ascospore septum of medium width. Pycnidia present, ostiole black. Conidia ellipsoid. Chemistry: Thallus and medulla K+ yellow, C, Pd, UV. Apothecial discs K+ red, C, Pd. Ostiolar tissue of pycnidia and aeruginose region of proper exciple K. Constituents: atranorin, gyrophoric and lecanoric acids (major compounds), parietin (traces). Ecology: It is so far known from the coastal regions where it grows abundantly on large siliceous boulders (rocks) both on subvertical and horizontal faces exposed to the sun along with Caloplaca cinnabarina (Ach.) Zahlbr., Buellia spp., Lecanora spp., Heterodermia diademata (Taylor) D. D. Awasthi, Physcia spp., Endocarpon petrolepideum Ach., Phylliscum spp., Aspicilia spp., Xanthoparmelia saxeti (Stizenb.) Amo de Paz, A. Crespo, Elix et Lumbsch, Xanthoparmelia spp., Ramalina spp., Verrucaria spp. Species diversity: So far it is monophyletic genus, while we expect that at least one more species closely related to Jasonhuria bogilana will be described based on Eastern Asian material. Distribution: This species was described only from the Bogil Island, southern part of South Korea, but now it is known from numerous coastal localities as continental as various islands of this country. Etymology: It is named after a known South Korean lichenologist, our friend prof. Jae- Seoun Hur (Sunchon, South Korea), the founder of the Korean Lichen Research Institute of Sunchon National University, who has played an important role in study of the Korean Lichen Flora and the whole genome investigations of lichen-forming fungi and their practical

8 application, as well as who described the type species of this genus. Taxonomic notes: The genus Jasonhuria is characterized by crustose, cracked areolate to areolate, greyish, K+ yellow thallus, a rust-red apothecial disc, a black proper margin, a grey thalline margin and maritime distribution, as well as atranorin, gyrophoric and lecanoric acids as major compounds. Unfortunately molecular data on the following taxa Caloplaca agrata (Vain.) Zahlbr., C. leptozona (Nyl.) Zahlbr., C. subleptozona Y. Joshi et Upreti, C. poliotera (Nyl.) J. Steiner and C. subpoliotera Y. Joshi et Upreti, mentioned as similar to type species in original description [21] are hitherto absent. Possibly some of them will be member of the new genus after checking of their molecular characters. Jasonhuria is similar to Usnochroma Søchting, Arup et Frödén in having gyrophoric acid, but differs in having different colour of thallus and apothecia, in having anthraquinones of parietin chemosyndrome, as well as in the lack of usnic acid. Loekoesia S. Y. Kondr., S.-O. Oh et J.-S. Hur gen. nov. MycoBank No. MB XXXXX Thallus crustose, entire to areolate; grey; soralia rounded, stipitate, aggregated in irregular groups, bright white; soredious mass bluish to whitish; soredia powdery. Hypothallus bluish black. Apothecia black, biatorine, true exciple paraplectenchymatous with well-developed matrix. Thallus K+ yellow, then greenish yellow, Pd+ slowly becoming pale yellow; probably contains atranorin and other compounds. Type species: Loekoesia austrocoreana (S. Y. Kondr., L. Lőkös et J.-S. Hur) S. Y. Kondr., J. Kim, A. S. Kondratiuk, S.-O. Oh et J.-S. Hur. Thallus crustose, entire to areolate; plumbeus grey to greyish white with brighter white soralia, sometimes coalescing in places; soralia rounded, stipitate, often aggregated in irregular groups; soredious mass bluish or becoming whitish. Soredia powdery, bluish. Hypothallus bluish black. Apothecia black, biatorine; true exciple paraplectenchymatous with well-developed

9 matrix; ascospores bipolar hyaline, elongated ellipsoid with rounded ends, ascospore septum of medium width. Chemistry: Thallus K+ yellow, then greenish yellow; Pd+ slowly becoming pale yellow; epihymenium K+ purple and becoming lighter to/or hyaline or dull crimson; probably contains atranorin and other compounds. Ecology: In coastal zone on rock surface growing together with Pyxine endochrysina Nyl., Physcia adscendens (Fr.) H. Olivier, Caloplaca squamosa (B. de Lesd.) Zahlbr. and species of the genera Aspicilia, Myelochroa, Buellia, Dimelaena, Verrucaria, and Lecanora. Etymology: It is named after a known Hungarian lichenologist, our friend and colleague Dr László Lőkös (Budapest, Hungary), who has made important contribution in the study of North and South Korean lichen-forming fungi, as well as who was co-author of the type species of this genus. Distribution: It is hitherto known from Eastern Asia (South Korea). Taxonomic notes: Loekoesia austrocoreana is similar to Caloplaca albovariegata (B. de Lesd.) Wetmore, a western North American species growing on calcareous and non-calcareous rocks having blue-grey thallus colour, stipitate areoles, thick irregular thallus cortex with epinecral layer, and the clumps of algae causing variegated areole surface. However, Loekoesia austrocoreana differs from the latter in having regular cortical layer of thallus, in having lower hymenium, in having shorter and narrower ascospores and wider ascospore septum, in having distinctly bluish epihymenium and bluish lateral true exciple and in absence of reaction with K in thalline cortex and lateral outermost part of true exciple, as well as in the lack of epinecral layer and clumps of algae [4, 22]. A number of other members of the genus Pyrenodesmia A. Massal., i.e. P. variabilis (Pers.) A. Massal., Caloplaca conversa (Kremp.) Jatta, Caloplaca atroalba (Tuck.) Zahlbr., Caloplaca peliophylla (Tuck.) Zahlbr., differs from Loekoesia austrocoreana in having much wider ascospores and in the lack of soredia.

10 Caloplaca oblongula (H. Magn.) Wetmore differs from Loekoesia austrocoreana in having light purplish brown epihymenium, in having non-septate or one-septate larger (both longer and wider) ascospores ( μm vs μm), and narrower ascospore septum (0 1.5 μm vs. 4 6 μm wide), as well as in having apothecial margin K+ purple [22]. The genus Loekoesia is similar to some representatives of the Australian genus Marchantiana S. Y. Kondr., Kärnefelt, Elix, A. Thell et J.-S. Hur of the Teloschistoideae, i.e. M. kalbiorum (S. Y. Kondr. et Kärnefelt) S. Y. Kondr., Kärnefelt, A. Thell, Elix, J. Kim, A. S. Kondratiuk et J.-S. Hur, but differs in its distribution and in its position in the subfamily Caloplacoideae after phylogenetic analysis based on combined set of ITS and LSU nrdna and 12S SSU mtdna sequences. Olegblumia S. Y. Kondr., L. Lőkös et J.-S. Hur gen. nov. MycoBank No. MB XXXXX Thallus lobate, distinctly rosette-like, upper surface brown to brownish grey; lobes flat to subconvex, very narrow; soralia laminal with convex, highly uplifted brownish soredious mass; soredia irregularly rounded, brown to brownish green; chemistry: constituents: vicanicin and caloploicin. Type species: Olegblumia demissa (Flot.) S. Y. Kondr., L. Lőkös, J. Kim, A. S. Kondratiuk, S.-O. Oh et J.-S. Hur. Thallus lobate, distinctly rosette-like, 5 8 mm diam., but often in much larger aggregations; upper surface brown, dark brown to greenish brown in peripheral portion and greyish brown or whitish greyish in the centre, in shade conditions thallus grey or whitish grey to whitish brown or brownish grey (without magnifying lens it seems to be with whitish pruina). Lobes to 1 1.5( 2) mm long, flat to subconvex, very narrow to ( 0.3) mm wide, towards the tips branched or divided into 2 3( 4) secondary lobules almost the same width; total width of terminal portion of single lobe with all secondary lobules to (0.3 )0.4 1 mm wide. Soralia

11 mainly in the centre of thallus, laminal (in the middle of lobe), at first punctiform or regularly rounded to (0.1 ) mm diam., soon becoming elongated along the lobe, fissure-like, to mm long/across, eroded portions with somewhat uplifted margins of cortical layer with convex, highly uplifted brownish soredious or soredious/isidious mass to confluent often whitish eroded-soredious mass in the centre. Soredia irregularly rounded, ca. (20 )30 50 µm across, seem becoming isidious, brown to dark brown or brownish green well contrasting to light (white) medulla. Apothecia, conidiomata and conidia unknown. Chemistry: Constituents: vicanicin and caloploicin. Ecology: On hard siliceous rocks usually on vertical and inclined surfaces often associated with Aspicilia contorta (Hoffm.) Körb., sometimes richly damaged by species of the genus Lichenostigma Hafellner, and Lichenothelia scopularia (Nyl.) D. Hawksw. (see also [23]), Caloplaca aractina (Fr.) Häyrén, Lecanora aff. frustulosa Stizenb., Lecanora lithophila Oxner and Aspicilia sp., Physcia sp. and Candelariella vitellina (Hoffm.) Müll. Arg. Species diversity and distribution: It is so far monotypic genus, which known from Europe and North America. Etymology: It is named after a known Ukrainian lichenologist Oleg Blum (1937 ) (Kyiv, Ukraine), who has made especially important contribution of lichen ecology of Eurasian lichens, as well as in the usage of lichens for bioindication studies on anthropogenic pollution of environment. Taxonomic notes: This genus Olegblumia is easily distinguished from the other caloplacoid lichens by brownish soredious lobate thallus having negative reaction with K. After morphological and chemical characters can be compared with the genera Elenkiniana S. Y. Kondr., Kärnefelt, Elix, A. Thell et J.-S. Hur, and Leproplaca (Nyl.) Hue of the subfamily Caloplacoideae, while after molecular characters is closely related only to Usnochroma. After having brownish lobate thallus having depsidones vicanicin and caloploicin it can

12 be similar to some species of the genus Elenkiniana, but differs from other taxa of this genus in colour of thallus and in the lack of anthraquinones of the parietin chemosyndrome. After having lobate soredious thallus it can be similar to some species of the genus Leproplaca (especially L. chrysodeta (Vain. ex Räs.) J. Laundon), but differs in having brownish thallus (vs. bright yellow to bright orange or reddish orange), in having brownish soredious mass (vs. bright yellow) and containing depsidones (vs. anthraquinones of the parietin chemosyndrome). This genus is known only from sterile conditions and for long period was considered in the Lecanoraceae (within the genera Lecanora or Placolecanora Räsänen), while after dark brown thallus can be similar to some Parmelias too. The usage of molecular markers allowed us to confirm its position in the family Teloschistaceae [24]. After combined phylogenetic analysis of the subfamily Caloplacoideae (Fig. 1) the genus Olegblumia is a sister group to the genus Usnochroma. New combinations Fauriea chujaensis (S. Y. Kondr., L. Lőkös et Hur) S. Y. Kondr., L. Lőkös, J. Kim, A. S. Kondratiuk, S.-O. Oh et J.-S. Hur. comb. nov. MycoBank No. MB XXXXX Basionym: Caloplaca chujaensis S. Y. Kondr., L. Lőkös et J.-S. Hur in Kondratyuk et al. Acta Bot. Hung. 57 (1 2): XXX (2015). Type: Republic of Korea. Jeju-do Province: Jeju-si, Chuja-do Island, Chuja-myeon, small gorge with steep shaded walls at Muk-ri, along the western coast. Lat.: N; Long.: E; Alt.: ca 10 m a.s.l., on siliceous rock, growing together with Caloplaca siderites. Coll.: Kondratyuk, S. Y. [site 15], L. Lőkös and J. Halda , (KoLRI holotype). Jasonhuria bogilana (Y. Joshi et Hur) S. Y. Kondr., L. Lőkös, J. Kim, A. S. Kondratiuk et S.-O. Oh comb. nov.

13 MycoBank No. MB XXXXX Basionym: Caloplaca bogilana Y. Joshi et Hur, The Lichenologist 42(6): 716 (2010). Type: Republic of Korea, Jeonnam Prov.: Wando Co., Bogil Island, N, E, alt. 5 m, on rock, 31 December 2004, Jae-Seoun Hur (KoLRI holotype). Loekoesia austrocoreana (S. Y. Kondr., L. Lőkös et J.-S. Hur) S. Y. Kondr., J. Kim, A. S. Kondratiuk, S.-O. Oh et J.-S. Hur comb. nov. MycoBank No. XXXXX Basionym: Caloplaca austrocoreana S. Y. Kondr., L. Lőkös et J.-S. Hur, in Kondratyuk et al., Acta Bot. Hung. 55 (1 2): 42 (2013). Type: Republic of Korea: Jeollanam-do Province: Yeosu-si, Nam-myeon, Yusong-ri, Geu-moh-do, on rock. Lat.: N; Long.: E; Alt.: 11 m a.s.l. Coll.: U. Jayalal, J. S. Park and J. A. Ryu (120513), (KoLRI holotype). Olegblumia demissa (Flot.) S. Y. Kondr., L. Lőkös, J. Kim, A. S. Kondratiuk, S.-O. Oh et J.-S. Hur comb. nov. MycoBank No. MB XXXXX Basionym: Imbricaria demissa Flot., Jber. Schles. Ges. Vaterl. Kultur 28: 133 (1850). Synonym: Caloplaca demissa (Flot.) Arup et Grube, Lichenologist 31 (5): 428 (1999). Conclusions The further data on molecular characters of crustose members of the Teloschistaceae will allow clarifying species diversity of four genera described here, as well as their diagnostic characters. Status of a number of phylogenetic branches, which are positioned within such clades as Mikhtomia s. lat., Variospora s. lat., Seirophora s. lat., will be especially discussed in our next paper on taxonomy and molecular phylogeny of the caloplacoid lichens.

14 Acknowledgements This work was supported in part by The State Agency on Science, Innovations and Information of Ukraine (M and M ) for SK, and by by the Korean Forest Service Program (KNA 2014) through Korea Forest Research Institute and Korean National Arboretum for JSH. References 1. Arup U, Søchting U, Frödén P. A new taxonomy of the family Teloschistaceae. Nord J Bot 2013a;31(1): Fedorenko NM, Stenroos S, Thell A, Kärnefelt I, Elix JA, Hur JS, Kondratuyk SY. Molecular phylogeny of xanthorioid lichens (Teloschistaceae, Ascomycota), with notes on their morphology. Bibl Lichenol 2012;108: Gaya E, Högnabba F, Holguin Á, Molnár K, Fernández-Brime S, Stenroos S, Arup U, Søchting U, van den Boom P, Lücking R, Sipman HJM, Lutzoni F.. Implementing a cumulative supermatrix approach for a comprehensive phylogentic study of the Teloschistales (Pezizomycotina, Ascomycota). Mol Phylogenet Evol 2012;63: Kondratyuk SY, Lőkös L, Zarei-Darki B, Haji Moniri M, Tchabanenko SI, Galanina I, Yakovchenko L, Hooshmand F, Ezhkin AK, Hur JS. Five new Caloplaca species (Teloschistaceae, Ascomycota) from Asia. Acta Bot Hung 2013a;55(1-2): Kondratyuk S, Jeong MH, Yu NH, Kärnefelt I, Thell A, Elix JA, Kim J, Kondratyuk AS, Hur JS. Four new genera of Teloschistoid lichens (Teloschistaceae, Ascomycota) based on molecular phylogeny. Acta Bot Hung 2013b;55(3-4): Kondratyuk SY, Jeong MH, Yu NN, Kärnefelt I, Thell A, Elix JA, Kim J, Kondratiuk AS, Hur JS. A revised taxonomy for the subfamily Caloplacoideae (Teloschistaceae, Ascomycota) based on molecular phylogeny. Acta Bot Hung 2014a;56(1-2):

15 7. Kondratyuk SY, Kärnefelt I, Thell A, Elix JA, Kim J, Jeong MH, Yu NH, Kondratiuk AS, Hur JS. A revised taxonomy of the subfamily Xanthorioideae (Teloschistaceae, Ascomycota) based on molecular phylogeny. Acta Bot Hung 2014b;56(1-2): Arup U, Søchting U, Frödén P. Addendum to A new taxonomy of the family Teloschistaceae. Nord J Bot 2013b;31: Kondratyuk et al. 2015b New subfamily of the Teloschistaceae (in press). 10. Kärnefelt I. Morphology and phylogeny in the Teloschistales. Cryptogamic Botany 1989;1: Søchting U, Sogaard MZ, Elix JA, Arup U, Elvebakk A, Sancho LG. Catenarina (Teloschistaceae, Ascomycota), a new Southern Hemisphere genus with 7- chlorocatenarin. Lichenologist 2014a;46(2): Søchting U, Garrido-Benavent I, Seppelt R, Castello M, Pérez-Ortega S, De Los Ríos Murillo A, Sancho LG, Frödén P, Arup U. Charcotiana and Amundsenia, two new genera in Teloschistaceae (lichenized Ascomycota, subfamily Xanthorioideae) hosting two new species from continental Antarctica, and Austroplaca frigida, a new name for a continental Antarctic species. Lichenologist 2014b;46(6): Orange A, James PW, White FJ. Microchemical methods for the identification of lichens. 2nd ed. London: British Lichen Society; Ekman S. PCR optimization and troubleshooting, with special reference to the amplification of ribosomal DNA in lichenized fungi. Lichenologist 1999;31(5): Gardes M, Bruns TD. ITS primers with enhanced specificity for basidiomycetes - application to the identification of mycorrhizae and rusts. Mol Ecol 1993;2: White TJ, Bruns T, Lee S, Taylor J. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. PCR protocols: a guide to methods and applications 1990;38:

16 17. Vilgalys R, Hester M. Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. J Bacteriol 1990;172(8): Fedorenko NM, Stenroos S, Thell A, Kärnefelt I, Kondratyuk SY. A phylogenetic analysis of xanthorioid lichens (Teloschistaceae, Ascomycota) based on ITS and mtssu sequences. Bibl Lichenol 2009;100: Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S. MEGA5: molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Mol Biol Evol 2011;28: Kondratyuk SY, Lőkös L, Farkas E, Hur JS. New and noteworthy lichen and lichenicolous fungi 2. Acta Bot Hung 2015a;57(1-2):xxx-xxx. 21. Joshi Y, Wang XY, Yamamoto Y, Koh YJ, Hur JS. A first modern contribution to Caloplaca biodiversity in South Korea: two new species and some new country records. Lichenologist 2010;42(6): Wetmore CM. The lichen genus Caloplaca in North and Central America with brown or black apothecia. Mycologia 1994;86: Mikhailyuk T.I., Kondratyuk S.Y., Nyporko S.O., Darienko T.M., Demchenko E.M., Voytsekhovicj A.O Lichen-forming fungi, bryophytes and terrestrial algae of granitic cayons of Ukraine. Kyiv: Alterpres, pp. 24. Arup U, Grube M. Where does Lecanora demissa (Ascomycota, Lecanorales) belong? Lichenologist 1999;31: Gaya E, Navarro-Rosinés P, Llimona X, Hladun N, Lutzoni F. Phylogenetic reassessment of the Teloschistaceae (lichen-forming Ascomycota, Lecanoromycetes). Mycol Res 2008;112(5): Søchting U, Figueras G. Caloplaca lenae sp. nov. and other Caloplaca species with caloploicin and vicanicin. Lichenologist 2007;39(1): Vondrák et al. (2012)

17 28. Gaya E, Lutzoni F, Zoller S, Navarro-Rosinés P. Phylogenetic study of Fulgensia and allied Caloplaca and Xanthoria species (Teloschistaceae, lichen-forming Ascomycota). Amer J Bot 2003;90(7): Martin MP, Winka K. Alternative methods of extracting and amplifying DNA from lichens. Lichenologist 2000;32:

18 Figure Captions Fig. 1. Phylogenetic tree of the caloplacoid lichens based on combined data set.

19 Table 1. Specimen vouchers used in the phylogenetic analysis with GenBank numbers Specimens included into phylogenetical analysis with GenBank numbers Species name Voucher details / references ITS LSU mt DNA Brigantiaea SK779, Kondratyuk et al. KF KF ferruginea (2013b) Brigantiaea SK780, Kondratyuk et al. KF KF ferruginea (2013b) Blastenia crenularia Gaya et al. (2012) JQ JQ Blastenia ferruginea KC KC KC Blastenia Arup et al. (2013a) KC subochracea Bryoplaca Arup et al. (2013a) KC jungermanniae Bryoplaca Arup et al. (2013a) KC KC sinapisperma Bryoplaca Arup et al. (2013a) KC tetraspora Caloplaca cerina FNM185, Fedorenko et al. EU EU (2009, 2012) Caloplaca cerina Gaya et al. (2012) JQ Caloplaca pelodella SK714, Kondratyuk et al. KF KF (2013b) Caloplaca Gaya et al. (2008) EU stillicidiorum Caloplaca HM thracopontica Caloplaca furax HQ Caloplaca furax Gaya et al. (2012) JQ Caloplaca phaeothamnos JN Eilifdahlia dahlii SK956, Kondratyuk et al. KJ KJ KJ Eilifdahlia dahlii SK959, Kondratyuk et al. Eilifdahlia wirthii SK262, Kondratyuk et al. Elenkiniana Søchting, Figueras (2006) ehrenbergii Elenkiniana gloriae SK750, Kondratyuk et al. Elenkiniana gloriae SK611, Kondratyuk et al. Elenkiniana gloriae SK613, Kondratyuk et al. Fauriea chujaensis SK709, China, CH KoLRI Fauriea chujaensis SK710, China, CH KoLRI KJ KJ KJ KJ KJ KJ DQ KJ KJ KJ KJ KJ KJ SK709 SK709 SK709 SK710 SK710 SK710

20 Fauriea chujaensis SKD07, South Korea, Jejudo, SKD07 SKD07 SKD07 Jeju-si, Chuja-do Island, Chuja-myeon, along the western coast, small gorge with steep shaded walls at Muk-ri, N, E, 10 m alt., on siliceous rock, growing together with Caloplaca aff. pelodella, C. aff. diffluens. Kondratyuk, S. site 15 (141138) KoLRI isotype Franwilsia bastowii SK810, Kondratyuk et al. KJ KJ KJ Franwilsia SK920, Kondratyuk et al. KJ KJ KJ kilcundaensis Franwilsia renatae SK235, Kondratyuk et al. KJ KJ Fulgensia fulgens SK735, Kondratyuk et al. KJ KJ Fulgensia cranfieldii SK983, Kondratyuk et al. KJ KJ KJ Fulgensia poeltii Gaya et al. (2008) EU Fulgogasparrea SK689, Kondratyuk et al. KF KF decipioides (2013b) Gyalolechia Gaya et al. (2008) EU canariensis Gyalolechia canariensis SK583, Kondratyuk et al. KJ Gyalolechia aurea Arup et al. (2013) KC KC KC Huneckia rheinigera SK3204, Kondratyuk et al. KJ Huneckia pollinii SK3206, Kondratyuk et al. KJ KJ KJ Huneckia pollinii SK870, Kondratyuk et al. KJ KJ KJ Jasonhuria bogilana KoLRI , Jeollanamdo, SK SK SK Yeosu-si, Nam-myeon, Geumoh-do, Usil coast side road, N, E 1 m alt., on rock U. Jayalal, J. S. Park & J. A. Ryu (120454) KoLRI Jasonhuria bogilana KoLRI , Jeollanamdo, Goheung-gun, Geumsan-myeon, Eojeon-ri, Geumoh-do, Simpo coast, N, E, 20 m alt., on rock, U. Jayalal, J. S. Park & J. A. SK SK SK120469

21 Ryu (120469) KoLRI Jasonhuria bogilana KoLRI , Jeollanamdo, SK SK SK Yeosu-si, Hwayang- myeon, Imok-ri, Baelga coast, N, E, 12 m alt., on rock, U. Jayalal, J. S. Park & J. A. Ryu (120641) KoLRI Jasonhuria bogilana KoLRI , the same SK SK SK locality U. Jayalal, J. S. Park & J. A. Ryu (120647) KoLRI Josefpoeltia SK991, Kondratyuk et al. KF KF KF sorediosa (2013b) Ioplaca pindarensis Gaya et al. (2012) JQ Kaernefia kaernefeltii SK921, Kondratyuk et al. (2013b) KF KF KF Leproplaca Arup et al. (2013a) KC KC KC xantholyta Leproplaca Gaya et al. (2012) JQ JQ xantholyta Leproplaca obliterans Arup et al. (2013a) KC KC Loekoesia austrocoreana KoLRI South Korea: Jeollanam-do, Yeosu-si, Nam-myeon, Yusong-ri, Geu-moh-do, on rock. Lat.: N; Long.: E; Alt.: 11 m a.s.l. Coll.: U. Jayalal, J. S. Park and J. A. Ryu (120511), SK SK SK Loekoesia austrocoreana Marchantiana maulensis Marchantiana occidentalis Marchantiana occidentalis Mikhtomia gordejevii Mikhtomia gordejevii Mikhtomia oxnerii Mikhtomia oxnerii KoLRI isotype KoLRI , the same locality (120523) KoLRI isotype SK994, Kondratyuk et al. SK981, Kondratyuk et al. SK982, Kondratyuk et al. SK80515, Kondratyuk et al. SK80646, Kondratyuk et al. SK90117, Kondratyuk et al. SK90755, Kondratyuk et al. SK SK SK KJ KJ KJ KJ KJ KJ KJ KJ KJ KJ KJ KJ KJ KJ KJ KJ021312

22 Olegblumia demissa SK C65, Ukraine: Mykolaiv SK C65 SK C65 SK C65 oblast, Arbuzynka district, right bank of Pivdenny Buh River, lower of Konstantinovka village, about 3 5 km lower along the river from Yuzhnoukrainsk town, near stone rapids on river, SE vertical surfaces of granite outcrops, at the plots 22, 23 and 24, Lat.: N; Long.: E; Alt.: ca 18 m a.s.l., Coll.: Kondratyuk, S. Y. (20311), Fedorenko N. M., (KW-L 70478) Olegblumia demissa Arup, Grube (1999) AF Olegblumia demissa Arup et al. (2013a) KC KC Olegblumia demissa Arup, Grube (1999) AF Olegblumia demissa Arup, Grube (1999) AF Oxneria alfredii FNM 152, Fedorenko et al. FNM 152 (2009) Pyrenodesmia SK747, Kondratyuk et al. KJ KJ alociza Pyrenodesmia teicholyta Vondrák et al. (2012) JN Pyrenodesmia teicholyta Arup et al. (2013a) KC Pyrenodesmia Gaya et al. (2003) AY variabilis Rufoplaca Arup et al. (2013a) KC KC KC scotoplaca Rufoplaca Arup et al. (2013a) KC KC KC tristiuscula Seirophora Arup et al. (2013a) KC californica Seirophora lacunosa SK B07, Ukraine: AR Krym, Arabatskaya strelka, on soil at the fortress, 200 m to NW, 10.vi.2003, O. Y. Khodosovsev (KW-L sub Lichenoconium xanthoriae) SK B07 SK B07 SK B07 Seirophora villosa Martin, Winka (2000) AF Teloschistes FNM-139, Fedorenko et al. EU EU flavicans (2009, 2012) Teloschistes Arup et al. (2013a) KC KC KC flavicans Usnochroma Arup et al. (2013a) KC KC KC carphinea Usnochroma EU639595

23 carphinea Usnochroma Gaya et al. (2012) JQ carphinea Usnochroma Gaya et al. (2012) JQ JQ scoriophila Variospora alpigena Arup, Grube (1999) AF Variospora latzelii Vondrák et al. unpubl. JN Variospora velana Arup et al. (2013a) KC KC KC Xanthocarpia SK637, Kondratyuk et al. KJ ochracea (2014b) Xanthoria parietina FNM-177, Fedorenko et al. EU EU (2009, 2012) Xanthoria parietina Gaya et al. (2012) JQ Yoshimuria SK768, Kondratyuk et al. KJ cerussata Yoshimuria SK725, Kondratyuk et al. KJ KJ spodoplaca Yoshimuria galbina SK704, Kondratyuk et al. KJ023197