Introduction
Lecanora Ach., a cosmopolitan genus with over 500 species (Kirk et al. Reference Kirk, Cannon, Minter and Stalpers2008), is one of the largest genera of lichenized ascomycetes, widely distributed from polar to tropical regions (Brodo Reference Brodo1984; Miyawaki Reference Miyawaki1988; Lumbsch Reference Lumbsch1994; Dickhäuser et al. Reference Dickhäuser, Lumbsch and Feige1995; Lumbsch et al. Reference Lumbsch, Feige and Elix1995, Reference Lumbsch, Guderley and Elix1996; Lumbsch & Elix Reference Lumbsch, Elix, McCarthy and Mallett2004; Ryan et al. Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004). The genus is characterized by asci of the Lecanora-type, simple, colourless ascospores, amphithecia often containing algal cells and crustose to placodioid growth forms (Lumbsch & Elix Reference Lumbsch, Elix, McCarthy and Mallett2004; Ryan et al. Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004). The morphological and anatomical characters within this genus show remarkable variability and, in addition, the chemistry is also complex. Not surprisingly, molecular data suggest the genus to be non-monophyletic (Arup & Grube Reference Arup and Grube1998, Reference Arup and Grube2000; Grube et al. Reference Grube, Baloch and Arup2004). The genus has received considerable attention during the past few decades which has resulted in numerous regional revisions and the description of numerous new species (Brodo Reference Brodo1984; Miyawaki Reference Miyawaki1988; Brodo et al. Reference Brodo, Owe-Larsson and Lumbsch1994; Lumbsch et al. Reference Lumbsch, Guderley and Elix1996; Guderley Reference Guderley1999; Guderley & Lumbsch Reference Guderley and Lumbsch1999; Lumbsch & Elix Reference Lumbsch, Elix, McCarthy and Mallett2004; Ryan et al. Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004; Nayaka Reference Nayaka2005; Han et al. Reference Han, Zhao and Guo2009; Knudsen & Lendemer Reference Knudsen and Lendemer2009). However, species in East Asia have been poorly studied, with the exception of Japanese species of the core group of Lecanora that were studied recently (Miyawaki Reference Miyawaki1988, Reference Miyawaki1994; Harada et al. Reference Harada, Miyawaki and Kawana1999). It was the French lichenologist Auguste-Marie Hue (Reference Hue1909), who recognized the first three species of Lecanora from Korea: L. hueana (Vain.) Hue, L. membranifera Hue and L. oreina (Ach.) Ach., all of which are now synonymized into other genera (Dimelaena, Rinodina). Subsequently Hue (Reference Hue1915) described 5 more species of Lecanora from Korea; all except, one, L. subrubra Hue (= L. japonica Müll. Arg.), are now transferred to Caloplaca. A recent checklist of Korean lichens (Hur et al. Reference Hur, Koh and Harada2005) listed only 12 Lecanora species. Thus we have recently turned our attention to the diversity of this genus in Korea and undertook field work in various parts of South Korea. This led to the discovery of two new species of Lecanora, together with eight new records for the country, and these are described below.
Materials and Methods
The study is based chiefly upon c. 200 specimens in KoLRI from various regions of South Korea. Additional specimens were studied from the following herbaria: BP and SDNU.
Apothecial anatomy was studied using hand-cut sections mounted in water. The morphological and anatomical characters of the specimens were examined using a stereo microscope (NIKON C-PS 1068908) and a microscope (OLYMPUS BX50). With the exception of L. loekoesii, L. nipponica and L. plumosa, for which only one specimen was available, ascospores measurements are based on at least 50 observations, and are presented in the format: (min.–) [x−SD]–[x+SD] (–max.), where min. and max. are the extreme values, x the arithmetic mean, and SD the corresponding standard deviation. Photographs of the thallus were taken using a SONY DSC-F717 and the photo of ascospores was taken using NIKON ECLIPSE 80i.
The terms hypothecium, amphithecium (consisting of medulla and cortex) and parathecium (or exciple) are used in the sense of Brodo (Reference Brodo1984) and Miyawaki (Reference Miyawaki1988), respectively.
Chemical analysis using spot tests and thin layer chromatography followed standard methods (Orange et al. Reference Orange, James and White2010), mainly using solvent system C.
The New Species
Lecanora hafelliana L. Lü, Y. Joshi & Hur sp. nov
MycoBank No.: MB 519214
Thallus epiphloeodes, crustaceus, continuus ad verrucosus, candidus ad griseus ad cineraceus. Apothecia sessilia, 0·5–1·6 mm in diametro. Disci glandacei vel rufi, ±pruinosi. Margo thallinus albidus, integer. Cortex distinctus. Amphithecium crystalla minuta continens. Epihymenium brunneum ad fuscum, granulosum. Hymenium hyalinum, 40–50 µm altum. Hypothecium hyalinum. Asci clavati, 8 spori. Ascosporae hyalinae, simplices, (10–)10·5–15(–15·9) × (5·7–)6·2–7·8(–8·3) µm. Pycnidia non visa. Thallus atranorinum, zeorinum, acida sticticum, hafellicum et usnicum continens.
Typus: South Korea, Jeollanam-do, Gwangyang-si, Okryong-myeon, Mt Baekwoon, 35°06′52·5″N, 127°36′43·5″E, alt. 1037 m, on bark, 27 June 2006, Hur 060423 (KoLRI—holotypus; SDNU—isotypus).
(Fig. 1A)
Fig. 1. Lecanora hafelliana. A, habit; B, example of a TLC analysis of L. hafelliana samples; C, distribution in South Korea. Scale: A = 1 mm. This figure appears in colour online.
Thallus corticolous, crustose, dull whitish to grey to ash grey, well-developed, continuous, rough to verruculose, margin definite. Prothallus absent.
Apothecia sessile, 0·5–1·6 mm diam.; discs yellowish brown to dark reddish brown, ± with white pruina, plane to slightly concave; margins usually paler than thallus, thick and prominent, smooth, entire, occasionally flexuose. Amphithecium with numerous small crystals (allophana-type) soluble in K, 75–90 µm thick laterally, 100–125 µm thick basally; cortex hyaline, gelatinous, distinct, 62·5–75 µm thick laterally, 85–100 µm basally. Epihymenium dark brown to blackish brown, 5–10 µm high, with coarse granules (chlarotera-type) soluble in K. Hymenium hyaline, not inspersed with oil droplets, 40–50 µm high; subhymenium hyaline, 17–25 µm high. Hypothecium hyaline, 22–30 µm high. Paraphyses simple, neither thickened apically nor pigmented. Asci clavate, 8-spored. Ascospores simple, hyaline, ellipsoid to narrowly ellipsoid, (10–)10·5–15(–15·9) × (5·7–)6·2–7·8(–8·3) µm.
Pycnidia not observed.
Chemistry. Spot tests: thallus K+ yellow, C−, KC+ yellow, P+ pale yellow. Secondary metabolites: stictic acid complex, hafellic acid, zeorin, usnic acid, atranorin (Fig. 1B).
Etymology. The specific epithet ‘hafelliana’ refers to the presence of hafellic acid.
Ecology and distribution. Lecanora hafelliana is a common species in South Korea found growing on bark of various deciduous trees including Acer, Betula, Cornus and Quercus, at elevations of 732–1663 m (Fig. 1C). The species was also collected from the bark of Pinus koraiensis in China at an elevation of 682 m.
Notes. This species is characterized by the rough to verruculose thallus, thick and smooth apothecial margins, coarse granules in the epihymenium, allophana-type amphithecium, and by the presence of usnic acid, zeorin, hafellic acid and the stictic acid complex, in addition to atranorin. It can be readily distinguished from other species of Lecanora by the presence of hafellic acid as a major substance. This acid can be detected as a yellow-orange spot at Rf value 0·33 in solvent C, and characterized by sky blue fluorescence under long wavelength UV light after sulfuric acid treatment and charring. It was previously known only from Buellia subtropica (Elix) Elix (Elix Reference Elix and McCarthy2009).
Some specimens with rather thick apothecial margins may be confused with Ochrolechia spp., but can be distinguished by having simple, unbranched paraphyses and smaller ascospores. Morphologically it is similar to L. megalocheila (Hue) H. Miyaw. and L. imshaugii Brodo. Lecanora megalocheila differs in having fine granules in the epihymenium, and in lacking usnic acid and hafellic acid; L. imshaugii differs in having flexuose apothecial margins, a clear, red-brown epihymenium, and lacks hafellic acid.
Additional specimens examined. South Korea: Jeollanam-do: Gurye-gun, Sandong-myeon, Mt Jiri, on bark, 2006, Hur 060164 (KoLRI); Toji-myeon, Mt Jiri, on Cornus bark, 2006, Hur 060905 (KoLRI); Gurye-eup, Mt Jiri, on bark, 2004, Hur 040289, on bark, 2006, Hur 060175, 060202, 060255 (KoLRI); Gwangyang-si, Okryong-myeon, Mt Baekwoon, on bark, 2006, Hur 060386 (KoLRI). Jeollabuk-do: Muju-gun, Seolcheon-myeon, Mt Deogyu, on bark, 2006, Hur 060492, on bark, 2005, Hur 050172 (KoLRI). Gyeongsangnam-do: Hamyang-gun, Seosang-myeon, Mt Baekwoon, on bark, 2006, Hur 060625 (KoLRI); Sancheong-gun, Sicheon-myeon, Jungsan-ri, Mt Jiri, on Quercus bark, 2006, Hur 060643, on Betula bark, 2006, Hur 060698, 060708-1 (KoLRI). Gangwon-do: Pyeongchang-gun, Jinbu-myeon, Mt Odae, on Quercus bark, 2008, Hur 080529 (KoLRI); Taebaek-si, Mungoksodo-dong, Mt Taebaek, on bark, 2005, Lőkös 050707 (KoLRI). Chungcheongbuk-do: Boeun-gun, Songnisan-myeon, Mt Songni, on bark, 2006, Hur 060010-1 (KoLRI); Danyang-gun, Gagok-myeon, Mt Sobaek, on Quercus bark, 2007, Hur 070396, Hur 070446, on Quercus bark, 2007, Hur 070494, on bark, 2007, Hur 070352 (KoLRI).—China: Jilin Prov.: Mt Changbai, on bark, 2006, Hur CH060190 (KoLRI).
Lecanora loekoesii Y. Joshi, L. Lü, & Hur sp. nov
MycoBank No.: MB 519215
Thallus epiphloeodes, tenuis, continuus, cinereus. Apothecia sessilia vel adnata, 0·4–1 mm in diametro. Disci lividi, ±pruinosi. Margo thallinus albidus, integer. Cortex indistinctus. Amphithecium crystalla minuta continens. Epihymenium lividum, granulosum. Hymenium hyalinum, 62·5–70 µm altum. Hypothecium hyalinum. Asci clavati, 12–16 spori. Ascosporae hyalinae, simplices, (12·1–)12·6–15·3(–16·2) × (7–)7·5–8·5(–9) µm. Pycnidia non visa. Thallus atranorinum, acidum usnicum, acidum norsticticum et zeorinum continens.
Typus: South Korea, Gangwon-do, Taebaek-si, Mungoksodo-dong, Mt. Taebaek, 37°06′48·8″N, 128°56′51·0″E, alt. 910 m, on Quercus bark, 13 October 2005, Lőkös 050717 (KoLRI—holotypus; BP—isotypus).
(Fig. 2A)
Fig. 2. Lecanora loekoesii. A, habit; B, asci and ascospores (section in water); C, distribution in South Korea. Scales: A = 1 mm; B = 10 µm. This figure appears in colour online.
Thallus corticolous, crustose, grey, continuous, thin, smooth to rough, epruinose, esorediate. Prothallus absent.
Apothecia numerous, sessile to adnate, 0·4–1 mm diam., lecanorine; discs yellowish brown, with or without white pruina, plane; margins usually paler than the thallus (whitish grey), thin, persistent, smooth, entire to occasionally flexuose. Amphithecium containing numerous small crystals (allophana-type) soluble in K, 45–50 µm thick laterally, basally not thickened; cortex absent or indistinct, hyaline, thin, 2–7 µm thick laterally, 5–11 µm thick basally. Epihymenium yellowish brown to golden brown, 20–25 µm high, with fine granules (pulicaris-type) dissolving in K. Hymenium hyaline, not inspersed with oil droplets, 62·5–70 µm high. Subhymenium hyaline, 15–20 µm high. Hypothecium hyaline, not inspersed, 25–45 µm high. Paraphyses simple, not pigmented. Asci clavate, 12–16-spored (Fig. 2B). Ascospores simple, hyaline, ellipsoid, (12·1–)12·6–15·3(–16·2)× (7–)7·5–8·5(–9) µm, wall less than 0·7 µm thick.
Pycnidia not observed.
Chemistry. Spot tests: thallus K+ yellow, C−, KC+ yellow, P+ yellow. Secondary metabolites: atranorin, usnic acid, zeorin and norstictic acid.
Etymology. The species is named in honour of Dr László Lőkös (Budapest, Hungary), the collector of the type specimen.
Ecology and distribution. At present L. loekoesii is known only from the type locality where it is found growing on Quercus bark at an elevation of 910 m (Fig. 2C).
Notes. So far, seven multispored species of Lecanora sensu stricto are known worldwide (Guderley & Lumbsch Reference Guderley and Lumbsch1999; Han et al. Reference Han, Zhao and Guo2009). The most distinct character for recognizing the new taxon is the presence of norstictic acid. Lecanora cateilea (Ach.) A. Massal., L. pleospora Müll. Arg., and L. weii L. F. Han & S. Y. Guo resemble L. loekoesii, but differ in several characters. Lecanoracateilea differs in having heavily pruinose apothecial discs, 8–12-spored asci and in containing psoromic acid and atranorin as secondary metabolites; L. pleospora differs in having amphithecia with large crystals and lacking phenolic compounds in addition to the atranorin chemosyndrome; while L. weii differs in having heavily pruinose apothecial discs, a chlarotera-type epihymenium and lacking norstictic acid and zeorin.
New Records
Lecanora campestris (Schaer.) Hue
Bull. Soc. Bot. France 35: 47 (1888).
(Fig. 3A)
Fig. 3. Lecanora species, new records in South Korea. A, L. campestris; B, L. cenisia; C, L. nipponica; D, L. perplexa; E, L. plumosa; F, L. polytropa; G, L. subrugosa; H, L. sulcata. Scales: A–H = 1 mm. This figure appears in colour online.
A description of this species and discussion of similar taxa can be found elsewhere (Brodo Reference Brodo1984; Ryan et al. Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004).
Ecology and distribution. Saxicolous. Widely distributed in the Northern Hemisphere: previously reported from northern Africa, Asia, western Europe and south-western North America (Upreti Reference Upreti1998; Ryan et al. Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004). Records from the Southern Hemisphere were based on misidentifications (Lumbsch Reference Lumbsch1994, Reference Lumbsch1998).
Specimens examined. South Korea: Jeju Island: on rock, 2004, Hur 040888 (KoLRI); Mt. Halla, on shore rock, 2006, Hur 061017 (KoLRI).
Lecanora cenisia Ach
Lich. Univ.: 361 (1810).
(Fig. 3B)
A description of this species and discussion of similar taxa can be found in Brodo (Reference Brodo1984) and Ryan et al. (Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004).
Ecology and distribution. Saxicolous. Previously reported from northern Africa, Asia, Europe and North America (Ryan et al. Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004).
Specimens examined. South Korea: Gyeongsangnam-do: Mungyeong-si, Mungyeong-eup, Mt Joryeong, on rock, 2006, Hur 061064 (KoLRI); Bonghwa-Gun, Myeongho-myeon, Mt Cheongryang, on rock, 2004, Hur 040101 (KoLRI); Tongyeong-si, Yokgi-myeon, Saryang Island, on rock, 2007, Hur 070019 (KoLRI).
Lecanora nipponica H. Miyaw
J. Hattori Bot. Lab. 64: 296 (1988).
(Fig. 3C)
A description of this species and discussion of similar taxa can be found in Miyawaki (Reference Miyawaki1988).
Ecology and distribution. Corticolous. Previously reported from Japan (Miyawaki Reference Miyawaki1988) and China (Lü et al. Reference Brodo, Owe-Larsson and Lumbsch2009a).
Specimen examined. South Korea: Jeollanam-do: Gurye-gun, Toji-myeon, Mt Jiri, on bark, 2006, Hur 060807-3 (KoLRI).
Lecanora perplexa Brodo
Beih. Nova Hedwigia 79: 148 (1984).
(Fig. 3D)
A description of this species and discussion of similar taxa can be found in Brodo (Reference Brodo1984) and Ryan et al. (Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004).
Ecology and distribution. Corticolous. Previously reported from North America (Brodo Reference Brodo1984) and China (Lü et al. Reference Brodo2009b).
Specimens examined. South Korea: Gangwon-do: Inje-gun, Girin-myeon, Mt Jeombong, on bark, 2004, Hur 041349, 041355, 041356 (KoLRI). Jeju Island: on bark, 2004, Hur 040873 (KoLRI). Gyeongsangnam-do: Sancheong-gun, Sicheon-myeon, Jungsan-ri, Mt Jiri, on bark, 2004, Hur 060742 (KoLRI).
Lecanora plumosa Müll. Arg
Flora 65: 484 (1882).
(Fig. 3E)
A description of this species and discussion of similar taxa can be found in Lumbsch (Reference Lumbsch1994), Guderley (Reference Guderley1999) and Lumbsch & Elix (Reference Lumbsch, Elix, McCarthy and Mallett2004).
Ecology and distribution. Saxicolous. A pantropical species extending into the subtropics and warm-temperate regions; previously reported from South Africa, Australasia, North and South America (Ryan et al. Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004). This is the first report from Asia.
Specimen examined. South Korea: Jeollanam-do: Jangheung-gun, Gwansan-eup, Mt Cheongwan, on rock, 2005, Lőkös 050644 (KoLRI).
Lecanora polytropa (Hoffm.) Rabenh
Deutschl. Krypt.-Fl. 2: 37 (1845).
(Fig. 3F)
A description of this well-known species and discussion of similar taxa can be found in Ryan et al. (Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004).
Ecology and distribution. Saxicolous, rarely on wood. It is a cosmopolitan species that is common in all continents, including Antarctica (Ryan et al. Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004).
Specimens examined. South Korea: Jeollabuk-do: Muju-gun, Seolcheon-myeon, Mt Deogyu, on rock, 2005, Hur 050088, on rock, 2006, Hur 060313, on rock, 2006, Hur 060514 (KoLRI). Gangwon-do: Taebaek-si, Mungoksodo-dong, Mt Taebaek, on rock, 2005, Lőkös 050710 (KoLRI).
Lecanora subrugosa Nyl
Flora 58: 15 (1875).
(Fig. 3G)
A description of this species and discussion of similar taxa can be found in Brodo (Reference Brodo1984) and Ryan et al. (Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004). The distinction of this taxon from Lecanora argentata (Ach.) Malme has been disputed (Lumbsch Reference Lumbsch1994) and needs re-examination. Here, we regard it as a gangaleoidin-lacking chemotype of L. argentata.
Ecology and distribution. Corticolous. Previously reported from Africa, Europe, North America (Ryan et al. Reference Ryan, Lumbsch, Messuti, Printzen, Śliwa, Nash, Nash, Ryan, Diederich, Gries and Bungartz2004) and China (Lü et al. Reference Lü, Wang and Zhao2009a).
Specimens examined. South Korea: Gangwon-do: Pyeongchang-gun, Jinbu-myeon, Mt Odae, on Quercus bark, 2008, Hur 080621 (KoLRI); Samcheok-si, Miro-myeon, Mt Duta, on bark, 2008, Hur 080105, Hur 080179 (KoLRI). Chungcheongbuk-do: Gosan-gun, Yeonpung-myeon, Mt Joryeong forest resort, on bark, 2008, Hur 080319 (KoLRI).
Lecanora sulcata (Hue) H. Miyaw
J. Hattori Bot. Lab. 64: 307 (1988).
(Fig. 3H)
A description of this species and discussion of similar taxa can be found in Miyawaki (Reference Miyawaki1988).
Ecology and distribution. Corticolous. Previously only known from Japan (Miyawaki Reference Miyawaki1988).
Specimens examined. South Korea: Gangwon-do: Mt Hambaek, on Quercus bark, 2007, Hur 070657-4 (KoLRI); Gyeongsangnam-do: Yangsan-si, Wondong-myeon, Mt Cheontae, on bark, 2006, Hur 061192 (KoLRI).
We are grateful to Drs L. Lőkös and P. K. Divakar for critically reviewing the manuscript and to Sunchon National University for providing laboratory facilities. This work was supported by a grant from Korea National Research Resource Center Program (2010-0000660) through the National Research Foundation of Korea (NRF) and the Korean Forest Service Program (KNA 2010) through the Korea National Arboretum. One of the authors (LL) would like to thank Irwin M. Brodo for his valuable suggestions and to Hae Sook Jeon and Xin Yu Wang for their kind cooperation during the study.
