Introduction
Caloplaca Th. Fr., a cosmopolitan genus occurring mostly in xeric and mesic habitats, includes perhaps more than 1000 species (Kärnefelt Reference Kärnefelt1989; Søchting & Lutzoni Reference Søchting and Lutzoni2003; Arup Reference Arup2006). It comprises a group of lichens with various growth forms (crustose, squamulose, lobate, subfruticose and even endosubstratic), with hyaline polarilocular spores (occasionally plurilocular) and usually with anthraquinone pigments (occasionally also with depsides, depsidones, xanthones and dibenzofurans) (Santesson Reference Santesson1970; Søchting & Frödén Reference Søchting and Frödén2002; Elix Reference Elix2008).
The genus has not been well worked out in South Korea and the literature pertaining to this genus is either scanty or scattered in various floristic accounts (Hue Reference Hue1913; Zahlbruckner Reference Zahlbruckner1930; Sato Reference Sato1943; Kim Reference Kim1981; Moon Reference Moon1999). Hur et al. (Reference Hur, Young and Harada2005) reported the occurrence of 14 species of Caloplaca from South Korea, tentatively including five species of Blastenia (B. atramentaria (Hue) Zahlbr., B. granuligera (Hue) Zahlbr., B. hexaspora (Hue) Zahlbr., B. injucunda (Hue) Zahlbr. and B. multicolor (Hue) Zahlbr.); these Blastenia species have not been studied by any taxonomists until now. Also, no recent contribution to this genus for any region of South Korea exists. Our paper presents descriptions of two new species, along with collection data for nine taxa new to South Korea. Our contribution increases the number of known South Korean Caloplaca species to twenty-five. Compared with Caloplaca biodiversity known in European territories of similar latitude and area, for example Greece (Abbott Reference Abbott2009), the number of known South Korean Caloplaca species is a small fraction of a total Caloplaca biodiversity in South Korea. Extensive exploration and further investigations on this genus in the near future will definitely increase its tally.
Materials and Methods
The present study is based on lichen specimens lodged at the herbarium of Lichen & Allied Bioresource Center, Korean Lichen Research Institute (KoLRI), Sunchon National University, Korea, as well as fresh collections from Jeju and Bogil-Do (Do means Island). Morphological characters of the thallus and reproductive structures, colour, size and shape were examined using standard microscopical techniques under a NIKON C-PS 1068908 dissecting microscope. Hand-cut sections prepared for studying the anatomy of thalli and fruiting bodies were examined under an OLYMPUS BX 50 compound microscope. All measurements were made on material mounted in water and lactophenol cotton blue (LCB) was used as a stain. For characters such as size of thallus, ascomata and thickness of the hymenium, hypothecium, exciple, ascospores and conidia dimensions, ten measurements were obtained per specimen. Only free ascospores lying outside the asci were measured. The dimensions are generally presented as (smallest value recorded–) mean of smallest recorded – mean of largest recorded (–largest value recorded). K, and C reactions were carried out on hand sections of thalli and apothecia under the microscope. Chemical analyses were made by TLC (White & James Reference White and James1985; Elix et al. Reference Elix, Johnston and Parker1987) using solvent system C (toluene : acetic acid; 85:15) and HPLC chromatography (Yoshimura et al. Reference Yoshimura, Kinoshita, Yamamoto, Huneck and Yamada1994) using YMC-Pack ODS-A column and eluent solvent of MeOH : H2O : H3PO4 (80:20:1). Terminology for apothecial tissues generally follows Wetmore (Reference Wetmore1994), while that for thallus follows Nash & Gries (Reference Nash, Gries, Nash, Ryan, Gries and Bungartz2002).
New Species
Caloplaca bogilana Y. Joshi & Hur sp. nov
Similis C. leptozona sed differt in substantiis secondariis dissimilibus, sporis grandioribus et distributione maritima.
Typus: South Korea, Jeonnam Prov., Wando Co., Bogil Island, 34°09′14.7″ N, 126°37′33.2″ E, alt. 5 m, on rock, 31 December 2004, Jae-Seoun Hur 041679 (KoLRI—holotypus; KNH—isotypus).
(Fig. 1A)
Caloplaca bogilana. A, habitus (holotype in KoLRI); B, distribution in South Korea. Scale: A = 2 mm.
Thallus saxicolous, crustose, determinate, usually growing in distinct irregular patches, areolate to cracked areolate, coalescing withother thalli to cover large areas, 2–9 cm diam., (110–)125–135(–150) µm thick, areoles plane to ± subconvex, 0·3–1 mm diam., grey to greyish white, dull, chalky. Cortex 22–25 µm high, paraplectenchymatous, made up of thin-walled cells, necral layer absent. Algal layer of even and continuous algal cells. Medulla macroscopically white, very thin, of compactly arranged hyaline hyphae. Vegetative diaspores absent. Prothallus present, black, often surrounding the thallus or where the thallus meets another thallus.
Apothecia biatorine to lecanorine, few to numerous, scattered to ± aggregated (1–3), adnate to sessile, round to ± angular owing to pressure, 0·3–0·6(–1·0) mm diam. Disc orange-brown to brownish red to rust-red, plane to convex, epruinose. Proper margin smooth, entire, thin, 20–40 µm, flush, persistent to evanescent, black. Thalline margin smooth, entire, thin to ± thick, 75–130(–210) µm, flush, persistent to evanescent, concolorous with the thallus. Epihymenium brown, with granular epipsamma, (7·5–)12·5–15·5(–16·3) µm high. Hymenium hyaline, (58·4–)62·5–87·5(–90) µm high. Hypothecium hyaline, (50–)75–100(–175) µm high, of isodiametric cells, without oil-droplets; proper margin annular, paraplectenchymatous, outer region ± aeruginose pigmented; thalline margin with algae. Paraphyses thin, simple to furcate at the top, with 1–2(–3) swollen cells at the top. Asci 57·5–62·5 × 10–12·5 µm, 8-spored, ascospores polarilocular, ellipsoid, (10–)12·5–15(–17)× 5·2–7·5 µm, isthmus (2·8–)3·3–5·8(–6·2) µm.
Pycnidia present, few to numerous, often 1 per areole, ostiole black. Conidia ellipsoid, 2·5–3·7 × 1·7–2 µm.
Chemistry. Spot tests: thallus and medulla K+ yellow, C−, Pd−, UV−. Apothecial discs K+ red, C−, Pd−. Ostiolar tissue and aeruginose region of proper exciple are K−. Secondary metabolites: atranorin, gyrophoric and lecanoric acids are the major compounds, while parietin is in traces.
Etymology. The specific epithet derives its name from the type locality (Bogil Island).
Ecology and distribution. The new taxon is so far known only from the coastal regions of Bogil Island (Fig. 1B) where it grows abundantly on large siliceous boulders (rocks) both on sub-vertical and horizontal faces exposed to the sun along with Caloplaca cinnabarina (Ach.) Zahlbr., C. stantonii W. A. Elder ex Arup, Buellia spp., Lecanora spp., Heterodermia diademata (Taylor) D.D. Awasthi, Physcia spp., Endocarpon petrolepideum Ach., Phylliscum spp., Aspicilia spp.,Karoowia saxeti (Stizenb.) Hale, Xanthoparmelia spp., Ramalina spp., Verrucaria spp.
Remarks. Caloplaca bogilana is characterized by a cracked areolate to areolate, matt, greyish, K+ yellow thallus surrounded by a black prothallus, a rust-red apothecial disc, a black proper margin, a grey thalline margin and maritime distribution. The new taxon is affiliated to the Caloplaca sideritis group (Wetmore Reference Wetmore1996) as anthraquinone pigments are present only in the apothecial discs and are absent from the thallus. The new taxon may be confused with Caloplaca agrata (Vain.) Zahlbr., C. leptozona (Nyl.) Zahlbr., C. subleptozona Y. Joshi & Upreti, C. poliotera (Nyl.) J. Steiner and C. subpoliotera Y. Joshi & Upreti. Caloplaca agrata differs in having granules in the thallus cortex and apothecia without thalline margins; C. leptozona differs in having smaller spores (10–11 × 4–5·5 µm), paraphyses without swollen tips, different chemistry (lacking gyrophoric and lecanoric acid) and an inland distribution; C. subleptozona differs in having cream to yellowish white thallus, biatorine apothecia with yellow pruinose discs and a temperate distribution; C. subpoliotera differs in having rimose-areolate thallus, numerous soredia, biatorine apothecia, smaller spores 6–8(–11) × (2–)3–5 µm and an inland distribution, while C. poliotera differs in having a rimose-areolate thallus, smaller spores (9–12 × 4–6 µm) and an inland distribution.
Additional specimens examined. South Korea: Jeonnam Prov.: Wando Co., Bogil Island, Bogil-myeon, Tong-ri, near Tong-ri beach, 34°09′640″ N, 126°35′115″ E, alt. 15 m, on rock, 2010, Y. Joshi et al. 100162 (KoLRI); Jung-ri, near Jung-ri beach, 34°09′714″ N, 126°35′527″ E, alt. 13 m, on rock, 2010, Y. Joshi et al. 100226 (KoLRI); Yesong-ri, near Yesong-ri beach, 34°08′293″ N, 126°33′566″ E, on rock, 2010, Y. Joshi et al. 100236 (KoLRI).
Caloplaca subflavorubescens Y. Joshi & Hur sp. nov
Similis C. flavorubescens et C. gordejevi sed differt in thallo squamoso vel subfolioso, margine crenato, substantiis secondariis dissimilibus.
Typus: South Korea, Kangwon Prov., Mt. Seokbyung, 37°34′31.0″ N, 128°52′02.7″ E, alt. 860 m, on bark, 24 May 2008, Jae-Seoun Hur 080212 (KoLRI—holotypus; KNH—isotypus).
(Fig. 2A)
Caloplaca flavorubescens. A, habitus (holotype in KoLRI); B, distribution in South Korea. Scale: A = 2 mm.
Thallus corticolous, subsquamulose to squamulose to subfoliose to ± crustose, covering a large area, up to 7 cm diam., (172–)190–250(–265) µm thick, yellowish green, greenish yellow, greenish grey to grey; upper surface with small lobules, 0·2–0·3(–0·7)× 0·1–0·2(–0·7) mm diam., uplifted at the margins, crowded to ± overlapping. Cortex (10–)12·5–15·5(–17) µm high, paraplectenchymatous, made up of thin walled cells, necral layer absent. Algal layer of even and continuous algal cells. Medulla macroscopically white, of compactly arranged hyaline hyphae, prosoplectenchymatous. Vegetative diaspores absent. Prothallus rarely present, black.
Apothecia zeorine, numerous, scattered to ± aggregated (1–4), sessile to ± constricted at the base, round to ± angular, 0·3–1(–2·2) mm diam., disc orange-red to orange-brown, plane to subconvex, glossy, proper margin smooth, entire, persistent, thin, 30–100(–180) µm, flush, concolorous or paler than disc, thalline margin present, persistent to ± evanescent in later stages, crenate to ± smooth, flush, (80–)100–170(–200) µm, concolorous to disc. Epihymenium golden brown, (10·2–)12·5–15(–25·3) µm high; hymenium hyaline, (57·7–)65–100(–110) µm high; hypothecium hyaline, of indistinct cells, with numerous oil-droplets, (103·8–)120–175(–180) µm high; proper margin cupular, prosoplectenchymatous, of compactly interwoven hyphae with narrow lumina; thalline margin with algae that covers the proper margin entirely. Paraphyses septate, with 1–3 swollen cells at the tip. Asci 8-spored, ascospores polarilocular, ellipsoid to broadly ellipsoid, (12–)15–17·5(–19) ×(5·5–)6–7(–7·5) µm, isthmus 7·5–10 µm.
Pycnidia rare, ostiole orange-red. Conidia bacilliform, 2·5–3·7 × 0·7–1·2 µm.
Chemistry. Spot tests: thallus K+ red (in yellowish regions), + yellow (in greyish regions), C−, Pd−, UV−. Apothecial disc and epihymenium K+ red, C+ red, Pd−. Medulla K−, C−, Pd−. Secondary metabolites: parietin, 7-chloroemodin, emodin, fragilin, atranorin, gyrophoric acid and triterpenes between Rf 6–7.
Two chemotypes are reported here for the new species: the most common chemotype (1) comprises 7-chloroemodin, emodin, fragilin, parietin and atranorin; while the rare one (2) contains gyrophoric acid as an additional substance.
Etymology. The specific epithet derives its name from C. flavorubescens (Huds.) J. R. Laundon, with which it is most confused in the field.
Ecology and distribution. Currently C. subflavorubescens is known from four localities in Kangwon Province. The species is found growing over bark between elevations of 860–1305 m in temperate regions of Mt. Seokbyung, Mt. Galiwang, Mt. Odae and Mt. Gaebang (Fig. 2B).
Remarks. The new taxon can be recognized by its subsquamulose to subfoliose greenish yellow to greenish grey thallus, with very small uplifted lobules, orange-red apothecia, crenate thalline margin, bacilliform conidia and corticolous habitat. Caloplaca flavorubescens (Huds.) J. R. Laundon and C. gordejevi (Tomin) Oxner ex Khodosovtsev with deep reddish apothecia are difficult to separate from the new taxon, but differ in having no lobules on the thallus surface. Also C. flavorubescens and C. gordejevi never have 7-chloroemodin or bacilliform conidia. Caloplaca himalayana Y. Joshi & Upreti, another related species known from temperate regions of the Himalaya, is often confused with this new taxon in thallus colour and ± similar chemistry (7-chloroemodin) but differs in having a crustose thallus, biatorine ferrugineous red apothecia with smooth margins, and in lacking pycnidia and some secondary metabolites (parietin and atranorin). Caloplaca juniperi Poelt & Hinter., another related species having a yellowish thallus and ferrugineous coloured apothecial discs, differs from the new taxon in being substratum in specific (only on Juniperus bark) and in its distribution in the Himalaya.
Additional specimens examined. South Korea: Kangwon Prov.: Mt. Odae, 37°46′06.7″ N, 128°36′10.7″ E,alt. 1305 m, on bark, 2008, Jae-Seoun Hur 080515 (KoLRI); Mt. Galiwang, 37°27′31.0″ N, 128°32′17.9″ E, alt. 1098 m, on bark, 2008, Jae-Seoun Hur 080082(KoLRI); Mt. Seokbyung, 37°34′57.3″ N, 128°52′27.1″ E,alt. 870 m, on bark, 2008, Jae-Seoun Hur 080227 (KoLRI).
New Records
C. cinnabarina (Ach.) Zahlbr
In Engler & Prantl., Nat. Pflanzenfam., Teil. I (Leipzig) 1: 228 (1908).—Lecanora cinnabarina Ach., Lich. univ.: 402 (1810).
For a description see Wetmore & Kärnefelt (Reference Wetmore and Kärnefelt1999).
Specimens examined. South Korea: Gyeongnam Prov.: Namhae-gun, Mt. Geum, 34°46′08.3″ N, 127°59′24″ E, alt. 171 m, on rock, 2004, Jae-Seoun Hur 040022 (KoLRI). Jeonbuk Prov.: Naeso temple, 35°37′02.7″ N, 126°35′01.7″ E, alt. 260 m, on rock, 2004, Jae-Seoun Hur 040167 (KoLRI). Jeonnam Prov.: Geogeum Island, 34°25′20.8″ N, 127°08′43.1″ E, alt. 10 m, on rock, 2005, Jae-Seoun Hur 050208/1 (KoLRI); Wando Co., Bogil-myeon, Bogil Island, Tong-ri, near Tongri beach, 34°09′64″ N, 126°35′11.5″ E, alt. 15 m, on rock, 2010, Y. Joshi, H. S. Jeon & M. H. Jeong 100166, 100194 (KoLRI); Jeongdong-ri seaside, 34°10′90″ N, 126°31′83.6″ E, alt. 11 m, on rock, 2010, Y. Joshi, H. S. Jeon & M. H. Jeong 100256, 100257 (KoLRI); Goheung-gun, Jeomam-myeon, Mt. Palyeong, 34°38′23.9″ N, 127°25′19.5″ E, alt. 85 m, on rock, 2010, Y. Joshi, H. S. Jeon & G. S. Han 100270 (KoLRI).
C. decipiens (Arnold) Blomb. & Forssell
Points Förteckning 4: 69 (1880).—Physcia decipiens Arnold, Flora, Jena 50: 562 (1867).
For a description see Wetmore & Kärnefelt (Reference Wetmore and Kärnefelt1998).
Specimens examined. South Korea: Gyeongbuk Prov.: Bonghwa Co., Mt. Cheongryang, 36°47′21.5″ N, 128°54′49″ E, alt. 490 m, on rock, 2004, Jae-Seoun Hur 040090 (KoLRI); 36°47′06.6″ N, 128°55′33.7″ E, alt. 825 m, on rock, 2004, Jae-Seoun Hur 040116 (KoLRI). Jeonbuk Prov.: Mt. Naejang, 35°29′44.7″ N, 126°53′41.3″ E, alt. 535 m, on rock, 2005, Jae-Seoun Hur 050004 (KoLRI); Gochang Co., Mt. Seonwoon, 35°28′30.4″ N, 126°33′42.8″ E, alt. 283 m, on rock, 2004, Jae-Seoun Hur 040076 (KoLRI); Mt. Juwang, 36°23′58.9″ N, 129°09′53.1″ E, alt. 310 m, on rock, 2005, Jae-Seoun Hur 050622 (KoLRI); Mt. Cheondae, 36°09′14.5″ N, 127°36′59.6″ E, alt. 214 m, on rock, 2006, Jae-Seoun Hur 061160 (KoLRI); 36°09′26.6″ N, 127°36′36″ E, alt. 356 m, on rock, 2006, Jae-Seoun Hur 061176 (KoLRI). Jeonnam Prov.: Goheung Co., Yeongnam-myeon, Ucheon-ri, Yongam village, Yongbawi seaside, 34°35′76.5″ N, 127°30′37.5″ E, alt. 10 m, on rock, 2010, Y. Joshi, H. S. Jeon & G. S. Han 100317, 100322, 100327 (KoLRI).
C. ferruginea (Huds.) Th. Fr
Nova Acta R. Soc. Scient. upsal., Ser. 3, 3: 223 (1861) [1860].—Lichen ferrugineus Huds., Fl. Angl. 2: 444 (1762).
For a description see Magnusson (Reference Magnusson1944).
Specimen examined. South Korea: Jeju Prov.: Jeju Island, alt. 565 m, 33°25′67.8″ N, 126°32′93.8″ E, on bark, 2009, Jae-Seoun Hur 090049 (KoLRI).
C. inconspecta Arup
Bryologist 98: 102 (1995).
For a description see Arup (Reference Arup1995).
Specimens examined. South Korea: Chungnam Prov.: Anmyon Island, 36°30′13.7″ N, 126°20′06.6″ E, alt. 5 m, on cliff rock, 2006, Jae-Seoun Hur 061238 (KoLRI). Jeju Prov.: Jeju Island, 33°34′00.1″ N, 126°45′44.4 E″, on dead wood, 2009, Jae-Seoun Hur 090033 (KoLRI).
C. pellodella (Nyl.) Hasse
Contrib. U. S. Nation. Herbarium 17: 115 (1913).—Lecanora pelodella Nyl., Lich. South California, edit. 2: 10 (1898).
For a description see Wetmore (Reference Wetmore1996).
Specimens examined. South Korea: Jeju Prov.: Jeju Island, 33°22′20.5″ N, 126°52′42.4″ E, alt. 1 m, on rock, 2006, Jae-Seoun Hur 061009 (KoLRI); 33°34′00.1″ N, 126°45′44.4″ E, on rock, 2009, Jae-Seoun Hur 090019 (KoLRI).
C. scopularis (Nyl.) Lettau
Hedwigia 52: 242 (1912).—Lecanora scopularis Nyl., Flora, Jena 56: 195 (1883).
For a description see Arup (Reference Arup1994).
Specimens examined. South Korea: Jeonnam Prov.: Wando Co., Bogil Island, Jung-ri, near Jung-ri beach, 34°09′714″ N, 126°35′527″ E, alt. 13 m, on rock, 2010, Y. Joshi & party 100204, 100221, 100223 (KoLRI).
C. squamosa (B. de Lesd.) Zahlbr
Cat. Lich. Univers. 10: 629 (1940).—Placodium squamosum B. de Lesd., Annal. Cryptog. Exot. 6: 123 (1933).
For a description see Wetmore (Reference Wetmore2003).
Specimen examined. South Korea: Jeju Prov.: Jeju Island, 33°34′00.1″ N, 126°45′44.4″ E, on rock, 2009, Jae-Seoun Hur 090020 (KoLRI).
C. stantonii W. A. Weber ex Arup
Bryologist. 95: 454 (1992).
For a description see Arup (Reference Arup1992).
Specimen examined. South Korea: Jeju Prov.: Jeju Island, 33°30′26.2″ N, 126°54′30.6″ E, on rock, 2004, Jae-Seoun Hur 040900, 040903 (KoLRI); 33°27′42.3″ N, 126°55′22.5″ E, on rock, 2004, Jae-Seoun Hur 040909 (KoLRI); 33°33′26″ N, 126°43′56.9″ E, on rock, 2004, Jae-Seoun Hur 040881 (KoLRI). Gyeongbnuk Prov.: Ulleung Co., Ulleung Island, 37°31′53.8″ N, 130°51′55.9″ E, alt. 11 m, on rock, 2005, Jae-Seoun Hur 050318 (KoLRI). Jeonnam Prov.: Wando Co., Bogil Island, 34°08′08.9″ N, 126°30′23.5″ E, alt. 5 m, on rock, 2004, Jae-Seoun Hur 041647, 041650 (KoLRI); Yeosu city, Geomun Island, 34°00′39.4″ N, 127°19′81.6″ E, alt. 14 m, on rock, 2007, Jae-Seoun Hur 070080 (KoLRI).
C. subsoluta (Nyl.) Zahlbr
Cat. Lich. Univers. 7: 185 (1931).—Lecanora murorum* subsoluta Nyl., Flora 56: 197 (1873).
For a description see Wetmore (Reference Wetmore2003).
Specimens examined. South Korea: Jeju Prov.: Jeju Island, 33°22′20.5″ N, 126°52′42.4″ E, alt. 1 m, on rock, 2006, Jae-Seoun Hur 061009 (KoLRI); 33°34′00.1″ N, 126°45′44.4″ E, on rock, 2009, Jae-Seoun Hur 090019, 090022 (KoLRI).
This work was supported by a grant from Korea National Research Resource Center Program (Grant 20090062634) and Korean Forest Service Program through Korea National Arboretum. We are indebted to Dr Jan Vondrák and an unknown reviewer for linguistic revision and helpful comments on an earlier draft of this paper. The first author also thanks Pavel Hrouzek for analysing the HPLC chromatograms and Miss Jung Ae Ryu, Hae Sook Jeon and Jin Young Hur for their kind help and cooperation during this study.
