Introduction
This paper mainly presents the results of an examination of G. Thor's collections of Brianaria and Micarea species in Japan, made between 1994 and 2012, as well as a small number from South Korea in 2001 and 2006, and one from Sri Lanka in 1978. Additional collections housed in the Uppsala herbarium (UPS) were examined during the first author's visit there in January 2011. The latest checklist of Japanese lichens (Ohmura & Kashiwadani Reference Ohmura and Kashiwadani2018) lists only seven species: Brianaria sylvicola (Flot. ex Körb.) S. Ekman & M. Svensson (as Micarea sylvicola), Micarea globulosella (Nyl.) Coppins, M. inquinans (Tul.) Coppins, M. melaena (Nyl.) Hedl., M. peliocarpa (Anzi) Coppins & R. Sant., M. prasina Fr. and M. synotheoides (Nyl.) Coppins, with a further species listed as Catillaria inopinula (Nyl.) Zahlbr. One additional species, M. adnata Coppins, was included in Kashiwadani et al. (Reference Kashiwadani, Kobayashi and Moon2011) but is not included in the checklist (Ohmura & Kashiwadani Reference Ohmura and Kashiwadani2018). Eighty-six species are currently known from Europe (including the four species referred to Brianaria by Ekman & Svensson (Reference Ekman and Svensson2014)). Of these, 63 occur in the British Isles (Coppins (Reference Coppins, Smith, Aptroot, Coppins, Fletcher, Gilbert, James and Wolseley2009), with four species added later). Therefore, more species could be expected to occur in Japan. Seven Micarea species are reported from South Korea; Joshi et al. (Reference Joshi, Nguyen, Wang, Lõkös, Koh and Hur2011) reported M. elachista (Körb.) Coppins & R. Sant. as new to South Korea, Kondratyuk et al. (Reference Kondratyuk, Lőkös, Tschabanenko, Haji Moniri, Farkas, Wang, Oh and Hur2013) described M. coreana Lőkös et al. and reported M. denigrata (Fr.) Hedl. and M. peliocarpa as new to South Korea, and Aptroot & Moon (Reference Aptroot and Moon2015) reported M. lignaria (Ach.) Hedl., M. micrococca (Körb.) Gams ex Coppins and M. misella (Nyl.) Hedl. as new to South Korea. Micarea coreana is discussed below under M. erratica (Körb.) Hertel et al. The species illustrated in Joshi et al. (Reference Joshi, Nguyen, Wang, Lõkös, Koh and Hur2011; fig. 2A, p. 71) as Micarea elachista is not this species and probably not a Micarea at all. The specimen might represent a Mycoblastus sp., as a red pigment is visible in one apothecium where the hymenium is visible.
Materials and Methods
Measurements of ascospores and asci were made in water mounts. For macroscopical investigation a Leica MZ6 stereo microscope with an eyepiece graticule was used. Ascospores, paraphyses, ascus walls, hyphae and conidia were measured with a 0.5 μm precision. For the newly described species, at least 50 ascospores were measured from the type collection. Secondary lichen compounds were identified with HPTLC (Arup et al. Reference Arup, Ekman, Lindblom and Mattsson1993), using the most stable solvent C (below as TLC). The colour reaction of the thallus was tested using common household bleach (C), 10% aqueous potassium hydroxide (K), 1,4-phenylendiamine in 95% ethanol (PD) and short-wave UV254 light. The material included in Joshi et al. (Reference Joshi, Nguyen, Wang, Lõkös, Koh and Hur2011) and Kondratyuk et al. (Reference Kondratyuk, Lőkös, Tschabanenko, Haji Moniri, Farkas, Wang, Oh and Hur2013) has not been seen by us. Additional material which might represent additional species will be studied further.
Taxonomy
New species from Asia
Micarea ceylanica Coppins sp. nov.
MycoBank No.: MB 836553
Similar to Micarea lignaria var. lignaria but with shorter, 1(–3)-septate ascospores; differs from Micarea rubioides in lacking soralia, having apothecia that do not form dense clusters, and in the longer ascospores, 10–18 vs (7.5–)9–11(–12) μm.
Type: Sri Lanka, Central Province, Kandy District, Dikoya, 6°52′N, 80°37′E, elev. 750 m, on the ground near the road, 13 February 1978, G. Thor 440 (S—holotype).
(Fig. 1)
Micarea ceylanica (holotype). Habit, thallus with dispersed apothecia. Scale = 1 mm. In colour online.
Thallus episubstratal, of scattered to coherent ± globose areoles, 0.12–0.2 mm diam., white to pale blue-grey. Soralia absent. In section without a well-defined cortex, but with a hyaline covering layer c. 5 μm tall; external hyphae sometimes surrounded by bluish grey pigment, K−, N+ red. Photobiont cells ‘micareoid’, 4–7 μm diam. Cephalodia absent.
Apothecia black, 0.25–0.5 mm diam., markedly convex to subglobose. Hymenium 40–55 μm tall, dilute aeruginose green, but with some vertically arranged lines of aeruginose pigment granules; dark aeruginose above (K−, N+ red), forming a distinct epithecium. Paraphyses numerous, monomorphic, sparingly branched, rather coherent in K, 1–1.5 μm wide, not or only slightly swollen at apices. Asci clavate, c. 40–50 × 12–15 μm, Micarea-type with a darker amyloid zone surrounding the apical cushion, 8-spored. Ascospores ovoid-ellipsoid to oblong and sometimes curved, 1(–3)-septate 10–18 × (3.5)–4(–5) μm. Hypothecium c. 100–120 μm tall, hyaline to dilute brownish. Exciple well developed but reflexed, dilute aeruginose, but with outwardly radiating streaks of aeruginose pigment granules; hyphae branched, outwardly radiating, c. 1 μm wide.
Conidiomata pycnidia, emergent to sessile, black, c. 50–70 μm diam.; wall green, K−, N+ red; conidia (microconidia) narrowly cylindrical, only faintly truncated at the proximal end, 6–7 × 0.8–1 μm.
Chemistry
Thallus C−, K−, PD+ red, apothecia in sections C−; argopsin detected by TLC.
Etymology
The specific epithet is an adjective derived from a Latinization of Ceylon, ‘Ceylonia’, the former name for Sri Lanka.
Notes
Similar in appearance, internal anatomy and pigmentation, and thallus chemistry to Micarea lignaria var. lignaria, but differing in the shorter (10–18 μm vs 16–36(–38) μm), mainly 1-septate ascospores which become noticeably curved with increasing length; those of M. lignaria are fusiform, not or only slightly curved, and 3–7-sepate. The new species differs from M. rubioides in lacking soralia, having apothecia that do not form dense clusters, and in the longer ascospores (10–18 μm vs (7.5–)9–11(–12) μm). Micarea pseudolignaria Brand et al. from Réunion (Brand et al. Reference Brand, van den Boom and Sérusiaux2014) differs from M. ceylanica in its shorter, ellipsoid ascospores, shorter microconidia and the presence of an unknown, PD− substance instead of argopsin, which is PD+ red.
Habitat and distribution
Known only from the type collection from mineral soil on a road cutting in a tea plantation. Coccocarpia palmicola (Spreng.) Arv. & D. J. Galloway was also collected in the vicinity (hb. S).
Micarea rubioides Coppins sp. nov.
MycoBank No.: MB 836554
Similar to Micarea lignaria var. lignaria but thallus with soralia, apothecia markedly confluent or tuberculate forming composite structures, and ascospores 1-septate and shorter (7.5–)9–11(–12) × (3–)3.5–4.5 μm.
Type: Japan, Kyushu, Kagoshima Pref. (Ohsumi Prov.), Yakushima Island, 4 km S of Miyanoura, along the road from Miyanoura to Shirotani-Unshuikyo Gorge, Kami-Yakucho, 30°23.5′N, 130°34′E, elev. 300 m, on soil at roadside, 25 October 1994, G. Thor 12310 (UPS—holotype; TNS—isotype).
(Fig. 2)
Micarea rubioides (holotype). Habit, thallus with dispersed soralia and apothecia. Scale = 5 mm. In colour online.
Thallus episubstratal, comprising crowded, convex, pale grey, matt areoles, 0.1–0.3 mm diam., which are mostly constricted below. Soralia present, efflorescent, scattered to confluent, pale bluish grey, 0.25–0.38 mm diam., but often confluent to form rounded patches up to 1.8 mm diam.; soredia 15–25 μm diam., external hyphae hyaline or with greenish, K−, N+ red pigment. In section, areoles without a well-defined cortex or amorphous covering layer; medulla absent or present in larger areoles and visible as a white core. Photobiont cells ‘micareoid’, 4–7 μm diam. Cephalodia absent.
Apothecia black, greenish black when wet, 0.15–0.38 mm diam., immarginate, mostly markedly confluent or tuberculate forming composite structures to 0.75 mm diam. Hymenium 50–65 μm tall, mostly hyaline except for a c. 10 μm tall, aeruginose-green pigmented zone in upper part (epithecium); pigment confined to the gel matrix, K−, N+ red. Paraphyses numerous, monomorphic, sparingly branched and anastomosing, rather coherent (even in K), 1.2–1.8 μm wide in mid-hymenium, not or only slightly swollen at apices to 2.5 μm. Asci clavate, c. 45–50 × 10–15 μm, Micarea-type with a darker amyloid zone surrounding the apical cushion, 8-spored. Ascospores ellipsoid to oblong-ellipsoid or ovoid-ellipsoid, 1-septate, (7.5–)9–11(–12) × (3–)3.5–4.5 μm. Hypothecium c. 150–180 μm tall, dilute olive-brownish, K−, N+ reddish, pigment confined to gel matrix; hyphae 1–1.8 μm wide, mostly vertically orientated; ascogenous hyphae with short cells swollen to 4 μm wide. Exciple indistinct, but visible in sections of young apothecia at the edge of the clusters, hyaline or tinged dilute greenish or olive-brownish; hyphae outwardly radiating, 1–1.5 μm wide, similar to paraphyses but more richly branched.
Conidiomata pycnidia, inconspicuous, immersed to emergent and sessile, black, 40–100 μm diam.; wall olive-brown or more strongly green in upper part, K−, N+ red; conidia (microconidia) narrowly cylindrical, only faintly truncated at the proximal end, 5–7 × 0.8–1 μm.
Chemistry
Thallus C−, K−, PD+ red, apothecia in sections C−; argopsin detected by TLC.
Etymology
The specific epithet refers to the resemblance of the clustered apothecia to the fruits of Rubus caesius L., as used in Acharius (Reference Acharius1803: 324) for the basionym of Psora rubiformis (Ach.) Hook.
Notes
Similar to Micarea lignaria var. lignaria, especially with regard to the internal anatomy and pigmentation of the apothecia, but with much smaller ((7.5–)9–11(–12) μm vs 16–36(–38) μm), 1-septate ascospores. Although similar in appearance, the thallus areoles lack the external, amorphous, hyaline covering-layer of M. lignaria, and sometimes break down to form efflorescent, bluish grey soralia. Sterile material of M. rubioides could be confused with the sorediate M. leprosula (Th. Fr.) Coppins & A. Fletcher, but the latter contains gyrophoric acid in addition to argopsin. When fertile, M. leprosula is distinguished by its 3-septate ascospores. Micarea ceylanica differs from M. rubioides in having an external, amorphous, hyaline covering-layer to the thallus areoles; longer, often curved ascospores; and lacking clustered apothecia. Micarea pseudolignaria Brand et al. from Réunion (Brand et al. Reference Brand, van den Boom and Sérusiaux2014) also resembles M. rubioides and has 1-septate, ascospores which are shorter (8.0–9.2 μm). In addition, M. pseudolignaria lacks soralia, has shorter microconidia and contains an unknown, PD− substance instead of argopsin, which is PD+ red. The recently described M. rubiformis P. M. McCarthy & Elix (McCarthy & Elix Reference McCarthy and Elix2020a), growing on soil in New Zealand, has blackberry-like apothecia but differs from M. rubioides in having a dark red-brown hypothecium, larger ascospores, an absence of soralia and in containing 2’-O-methylsuperperlatolic acid rather than argopsin.
Habitat and distribution
The type collection is from mineral soil on the side of a dirt road. Yakushima is the southernmost island where the conifer Cryptomeria japonica still occurs. It differs from the surrounding islands by the presence of high mountains, the highest peak reaching 1935 m. The annual precipitation is extremely high, with 3000–5000 mm in the lowlands and up to 10 000 mm in the montane area. The lichen flora is distinctive with, for example, the endemic Calicium muriformis Tibell (Tibell & Thor Reference Tibell and Thor2003) and Gymnoderma (e.g. Kashiwadani & Gradstein Reference Kashiwadani and Gradstein1982; Yoshimura Reference Yoshimura1982). The lowland vegetation is subtropical. A section of the montane area belongs to a national park, partly covered with forests of very large and old trees of Cryptomeria japonica. Micarea rubioides has also been found amongst collections from Malaysia (Borneo, Sarawak) and the Philippines (Luzon). The Sarawak collection was from a montane ridge at 1525 m, on mineral soil and bryophytes in crevices between stones on a landslip, along with Baeomyces heteromorphus Nyl. ex C. Bab. & Mitt., and the collection from Luzon was found in an open area in the cloud forest zone.
Additional specimens examined
Japan: Kyushu: Kagoshima Pref. (Ohsumi Prov.), Yakushima Island, c. 14 km W of Anbo, along the path to the peak Miyanoura-dake, Hananoego marsh 2 km NW of Yodokawa lodge, 30°18.5′N, 130°31′E, 1650 m, on mossy stump, 1994, G. Thor 12500 (TNS, UPS; TLC, argopsin; no apothecia).—Malaysia: Borneo, Sarawak: Gunong Mulu National Park, ridge between top of Pantu Ridge and Gunong Tamacu, 1525 m, on soil and bryophytes in crevices between stones on landslip, 1978, Coppins 5712 (E).—Philippines: Luzon: Benguet Prov., Baguio, Mt Santa Tomas, 120°35′E, 16°18′N, cloud forest, 2200 m, on soil, 1987, Aptroot 20305 (hb. Aptroot).
Additional reports of Micarea species from Japan and South Korea
The species of Micarea so far known from Japan are enumerated, with a small selection of cited specimens. The total number of collections for each species made during G. Thor's visits to Japan from 1994, 1995, 1996, 1997, 2006, 2010, and 2012 and South Korea in 2001 and 2006, are indicated after the name of the country. The examination of collections from Japan has increased the number of Brianaria and Micarea species known from that country from eight to 19. Eleven species are reported as new to Japan (M. botryoides (Nyl.) Coppins, M. denigrata, M. erratica, M. hedlundii Coppins, M. lithinella (Nyl.) Hedl., M. micrococca and M. misella) or new to Asia: M. byssacea (Th. Fr.) Czarnota et al., M. deminuta Coppins and M. xanthonica Coppins & Tønsberg (new to Asia, Japan); M. nitschkeana (J. Lahm ex Rabenh.) Harm. (new to Asia, South Korea).
Brianaria sylvicola (Flot. ex Körb.) S. Ekman & M. Svensson
Syn.: Micarea sylvicola (Flot. ex Körb.) Vězda & Wirth.
Notes
This species has recently been transferred from Micarea to the new genus Brianaria S. Ekman & M. Svensson in the Psoraceae (Ekman & Svensson Reference Ekman and Svensson2014).
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
Shaded siliceous rocks; 750–1220 m. A widely distributed species in Japan but few findings. First reported from Japan by Inoue & Moon (Reference Inoue and Moon1998, as Micarea sylvicola). Also found in Asia from Turkey (Yildiz et al. Reference Yildiz, John and Yurdakulol2002), eastern Russia (Andreev et al. Reference Andreev, Kotlov and Makarova1996), Mongolia (Schubert & Klement Reference Schubert and Klement1977) and Taiwan (Aptroot & Sparrius Reference Aptroot and Sparrius2003); elsewhere from Europe, North and South America, and Australia (Tasmania (Kantvilas & Coppins Reference Kantvilas and Coppins2019)).
Specimens examined
Japan (3): Izu Islands: Hachijo-Island, Mt Hachijo-Fuji, 33°08′N, 139°46′E, 840 m, under overhang of rock, 1994, Thor (12036) & Kashiwadani (TNS, UPS). Kochi: Ino-cho town, Teregawa, S slope of Mt Kamegamori, path along the stream Shirai-dani, humid old-growth, mixed deciduous/coniferous forest with e.g. Abies firma, Cryptomeria japonica, Quercus spp. and Tsuga sieboldii, 33°45.09–16′N, 133°11.48–54′E, 750–770 m, vertical shaded rock, 2006, Thor 21228 (UPS). Honshu: Toyama Pref. (Etchu Prov.), 30 km ESE of Toyama, Tateyama-cho, Nakashinkawa-gun, Syomyo-zaka, 600 m W of Shomyo Waterfall, just S of the river, path on the steep slope, mixed deciduous/Cryptomeria forest, 36°34′N, 137°31′E, 1160–1220 m, 1994, on shaded rock, Thor 12625 (TNS, UPS).
Micarea adnata Coppins
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
A rare species on lignum of stumps and logs, and bases of old trees in old-growth forest; 220–1480 m. Previously reported for Asia from Japan (Kashiwadani et al. Reference Kashiwadani, Kobayashi and Moon2011) and Taiwan (Aptroot & Sparrius Reference Aptroot and Sparrius2003); elsewhere from Europe (Czarnota Reference Czarnota2007), Macaronesia and North America (Fryday & Coppins Reference Fryday, Coppins, Nash, Gries and Bungartz2007).
Specimens examined
Japan (4): Kochi: Ino-cho town, Teregawa, NE slope of Mt Iwaguro-yama, along path from the road, tall, humid, old-growth, mixed deciduous/coniferous forest with e.g. Abies homolepis, Fagus crenata and Quercus spp., 33°45′10ʺN, 133°09′54ʺE, 1300 m, 2006, Thor 21326 (TNS, UPS). Honshu: Wakayama Pref. (Kii Prov.), Mt Koya, the walk from Ichinohashi bridge to the temple Okuno-in, old Cryptomeria japonica forest, 34°13′N, 135°35′E, 800 m, on stump in dense forest, 1994, Thor 12125 (TNS, UPS); Tochigi Pref. (Shimotsuke Prov.), Nikko National Park, 18.5 km WNW of Nikko, 1 km SSE of the village Yumoto, above Yutaki Falls, dense old-growth mixed coniferous/deciduous forest, 36°48′N, 139°26′E, on a log, 1997, Thor 15553b (UPS). Hokkaido: Kitami Prov., Shari-gun, Shari-cho, Shiretoko National Park, NW slope of Shiretoko Peninsular, c. 7 km NE of Utoro town, N of the small road to Iwaobetsu hot-spring hotel (Onsen), old-growth lowland forest dominated by Abies sachalinensis and Quercus crispula, 44.10947°N, 145.08242°E, 220 m, on a stump, 2010, Thor 24878 (UPS).
Micarea botryoides (Nyl.) Coppins
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
A rare species on bark of old trees (Abies, Betula, Picea) in old-growth humid forest; 600–1640 m. New to Japan, previously reported for Asia from Taiwan (Aptroot & Sparrius Reference Aptroot and Sparrius2003); elsewhere from Europe, Macaronesia and North America (Czarnota Reference Czarnota2007).
Specimens examined
Japan (3): Honshu: Tochigi Pref. (Shimotsuke Prov.), Nikko National Park, 19.5 km WNW of Nikko, 2.2 km NNE of the village Yumoto, along trail from Yumoto, N slope just S of Lake Karigome-ko, 36°49′N, 139°26′E, 1640 m, dense old-growth forest dominated by coniferous trees, on Betula bark, 1997, Thor 15608 (TNS, UPS). Hokkaido: Ishikari Prov., Higashikawa-cho, 140 km NE of Sapporo city, W slope of Mt Asahidake, village of Asahidake Onsen, 200–600 m SW of the ropeway station, along the trail ‘Kumagera course’ from Lodge Nutapukaushipe to the ropeway station, old-growth montane forest, 43°39.0395′N, 142°47.4070′E, 1100 m, on Picea sp., 2010, Thor 25852 (UPS); Kushiro Prov., Kushiro-shi, 53 km NNW of Kushiro city, Mt O-Akan dake, along the trail from the road at the easternmost end of Lake Akan (elev. 420 m) to the top of the mountain (elev. c. 1350 m), old-growth montane forest with e.g. Abies sachalinensis, Acer spp., Larix leptolepis, Magnolia sp., Picea sp. and Quercus crispula, 43°27′N, 144°09′E, 600–1100 m, on Abies sachalinensis, 2010, Thor 26046 (UPS).
Micarea byssacea (Th. Fr.) Czarnota et al.
Description and illustrations
Czarnota & Guzow-Krzemińska (Reference Czarnota and Guzow-Krzemińska2009).
Habitat and distribution
Common and widely distributed on bark and lignum of trees, logs and stumps in old-growth or humid forest; 20–2000 m. New to Asia, but most records from Japan of Micarea prasina probably belong here (e.g. Thor et al. Reference Thor, Kashiwadani and Moon2005); it might be the most common Micarea species in Japan, at least in forests. It was reported as Micarea sp. from the Imperial Palace grounds, Tokyo, by Ohmura et al. (Reference Ohmura, Thor, Frisch, Kashiwadani and Moon2014). It is common in Europe and was recently reported as new to North America from coastal Maine (Launis & Myllys Reference Launis and Myllys2014) and is considered widespread and common in Tasmania (Kantvilas & Coppins Reference Kantvilas and Coppins2019). All specimens examined have Sedifolia-grey pigment (greenish, K+ violet) in the thallus and apothecia, and methoxymicaraeic acid determined by TLC; thus, they do not represent the newly described Micarea microareolata Launis et al. or M. laeta Launis & Myllys in the M. byssacea complex (Launis et al. Reference Launis, Pykälä, van den Boom, Sérusiaux and Myllys2019a, Reference Launis, Malíček, Svensson, Tsurykau, Sérusiaux and Myllysb). Recent advances using molecular data for unravelling the complexities of Micarea prasina s. lat. (sensu Coppins Reference Coppins1983) have been based almost solely on European material. We strongly recommend that such studies should be expanded to other temperate regions of the world, where this group is well represented.
Selected specimens examined
Japan (c. 25): Honshu: Tokyo-to = Tokyo Metropolis (Musashi Prov.), Tokyo, Shiroganedai, Minato-ku, garden of the Institute for Nature Study, National Science Museum, old-growth evergreen forest, 35°38′N, 139°43′E, 30 m, on log of deciduous tree, 1999, Thor 16609 (TNS, UPS); Aomori Pref. (Mutsu Prov.), Mt Hakkoda, S of Suiren-numa pond, open mixed coniferous/deciduous forest, 40°37′N, 140°55′E, 900 m, on 2 m high stump, 1994, Thor 12000 (TNS, UPS). Hokkaido: Kitami Prov., Shari-gun, Shari-cho, Shiretoko National Park, NW slope of Shiretoko Peninsular, c. 7 km NE of Utoro town, N of the small road to Iwaobetsu hot-spring hotel (Onsen), old-growth lowland forest dominated by Abies sachalinensis and Quercus crispula, 44.1112°N, 145.0850°E, 249 m, on a stump, 2010, Thor 25021 (TNS, UPS).
Micarea cf. czarnotae
A collection from South Korea resembles Micarea byssacea in apothecial anatomy and pigmentation, and in thallus chemistry (methoxymicareic acid). However, it differs in the thallus being composed of small convex to subglobose areoles, 70–200 μm diam., rather than small, granule-like goniocysts, 12–40 μm diam. It is also similar to M. meridionalis van den Boom et al. (van den Boom et al. Reference van den Boom, Brand, Coppins and Sérusiaux2017) but that species contains micareic acid and has slightly smaller areoles, 40–70 μm diam. The recently described M. pseudotsugae van den Boom et al. (van den Boom et al. Reference van den Boom, Guzow-Krzemińska and Kukwa2020) has an areolate thallus containing methoxymicareic acid, but Superba-brown replaces Sedifolia-grey as its apothecial pigment. This South Korean collection should be compared with another recently described species, M. czarnotae Launis et al. (Launis et al. Reference Launis, Pykälä, van den Boom, Sérusiaux and Myllys2019a), which has goniocysts that can coalesce to form a warted-areolate thallus and also an identical thallus chemistry. This species is now known to occur in North America (Spribille et al. Reference Spribille, Fryday, Pérez-Ortega, Svensson, Tønsberg, Ekman, Holien, Resl, Schneider and Stabentheiner2020).
Specimen examined
South Korea (1): Pusan: Songdo-gu, Songdo-Annam Park, path along the coast, open Quercus dentata/Pinus densiflora forest, 35°02′N, 129°03′E, 0–120 m, on Pinus densiflora, 2001, Thor 16905 (UPS).
Micarea deminuta Coppins
Descriptions and illustrations
Coppins (Reference Coppins1995), Czarnota (Reference Czarnota2007).
Habitat and distribution
In Japan, mainly on bark, especially of Cryptomeria japonica, in old-growth but also managed forests; 750–c. 1500 m, widely distributed but scattered. Interestingly, most collections from Europe are on lignum. New to Asia; elsewhere from Europe, North America (California (Fryday & Coppins Reference Fryday, Coppins, Nash, Gries and Bungartz2007)) and Australia (Tasmania (Czarnota Reference Czarnota2007); New South Wales, on soil (McCarthy & Elix Reference McCarthy and Elix2020b)). A widely distributed species with scattered findings.
Selected specimens examined
Japan (10): Kyushu: Kagoshima Pref. (Ohsumi Prov.), Yakushima Island, c. 14 km W of Anbo, along the path to the peak Miyanoura-dake, from c. 500 m E of Yodokawa lodge to Hananoego marsh 2 km NW of Yodokawa lodge, dense forest, 30°18′N, 130°31′E, 1400–1650 m, on bark of Cryptomeria japonica, 1994, Thor 12486 (TNS, UPS). Kochi: Ino-cho town, Teregawa, S slope of Mt Kamegamori, path along the stream Shirai-dani, humid old-growth, mixed deciduous/coniferous forest with e.g. Abies firma, Cryptomeria japonica, Quercus spp. and Tsuga sieboldii, 33°45.09–16′N, 133°11.48–54′E, 750–770 m, on coniferous tree, 2006, Thor 21250, 21274, 21275 (UPS). Honshu: Aomori Pref. (Mutsu Prov.), Mt Hakkoda, around Yachi Hot Spring, dense forest near small stream, 40°38′N, 140°56′E, 750 m, on Cryptomeria japonica [bark], 1994, Thor 11873a (TNS).
Micarea denigrata (Fr.) Hedl.
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
A rare but widespread species on lignum of tree stumps in old-growth forests; 600–c. 1640 m. New to Japan. Previously reported for Asia from South Korea (Kondratyuk et al. Reference Kondratyuk, Lőkös, Tschabanenko, Haji Moniri, Farkas, Wang, Oh and Hur2013), eastern Russia (Siberia (Konoreva et al. Reference Konoreva, Chesnokov and Poryadina2018)) and Taiwan (Aptroot & Sparrius Reference Aptroot and Sparrius2003); elsewhere from Europe, North America and Australia (Czarnota Reference Czarnota2007).
Specimens examined
Japan (2): Honshu: Tochigi Pref. (Shimotsuke Prov.), 19.5 km WNW of Nikko, 2.2 km NNE of the village Yumoto, along trail from Yumoto, c. 100 m S of the small lake Karigomeko, north slope with old-growth, dense forest dominated by coniferous trees, 36°49′N, 139°26′E, 1640 m, on lignum of old, large Tsuga diversifolia stump, 1996, Thor 15062 (UPS; TLC, nil). Hokkaido: Kushiro Prov., Kushiro-shi, 53 km NNW of Kushiro city, Mt O-Akan dake, along the trail from the road at the easternmost end of Lake Akan to the top of the mountain, old-growth montane forest, 43°27′N, 144°09′E, 600–1100 m, on stump, 2010, Thor 26033 (TNS, UPS; TLC, nil).
Micarea erratica (Körb.) Hertel et al.
Syn.: Leimonis erratica (Körb.) R. C. Harris & Lendemer.
Syn. nov.: Micarea coreana L. Lőkös et al. Described and reported from South Korea (Kondratyuk et al. Reference Kondratyuk, Lőkös, Tschabanenko, Haji Moniri, Farkas, Wang, Oh and Hur2013). In the protologue it was compared only with M. farinosa Coppins, with which it bears no resemblance. In our opinion it is undoubtedly M. erratica.
Descriptions and illustrations
Czarnota (Reference Czarnota2007), Kondratyuk et al. (Reference Kondratyuk, Lőkös, Tschabanenko, Haji Moniri, Farkas, Wang, Oh and Hur2013).
Habitat and distribution
A rare but widespread species on siliceous stones; 1080–2205 m. New to Japan. Previously reported for Asia from Indonesia (Java (Hertel Reference Hertel1977)) and South Korea (Kondratyuk et al. Reference Kondratyuk, Lőkös, Tschabanenko, Haji Moniri, Farkas, Wang, Oh and Hur2013, as M. coreana); elsewhere from Europe, North America, Australasia (Czarnota Reference Czarnota2007; Kantvilas & Coppins Reference Kantvilas and Coppins2019) and Réunion (Brand et al. Reference Brand, van den Boom and Sérusiaux2014).
Specimens examined
Japan (2): Honshu: Yamanashi Pref. (Kai Prov.), Yamanashi-city, Makioka-cho, at the road to Odarumi Pass, roadside with grass and scattered stones, 35°51.726′N, 138°39.624′E, 2205 m, on stone, 2012, Thor 28153 (TNS, UPS). Hokkaido: Ishikari Prov., Higashikawa-cho, 140 km NE of Sapporo, W slope of Mt Asahidake, village of Asahidake Onsen, 600 m SW of the ropeway station, at Lodge Nutapukaushipe, 43°38.5582′N, 142°47.3782′E, 1080 m, on stone in wall at the lodge, 2010, Thor 25831 (UPS).
Micarea globulosella (Nyl.) Coppins
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
Not found during the recent field visits and known from Japan only from the type collection (Yokohama, on bark, 1879, E. Almquist (S, H-NYL)). It occurs elsewhere in Europe and North America but is rarely collected, except in old-growth coniferous forests in northern Sweden.
Micarea hedlundii Coppins
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
A rare species found once on the base of an old coniferous tree in old-growth forest; 750–770 m. New to Japan; elsewhere from shaded habitats in old-growth woodlands in wet forests or valley woodlands of Europe (including Russia, Komi Republic (Hermansson et al. Reference Hermansson, Pystina and Kudrycheva1998)), Macaronesia, Asia (eastern Siberia (Konoreva et al. Reference Konoreva, Chesnokov and Poryadina2018)), North America (New Brunswick (Gowan & Brodo Reference Gowan and Brodo1988); California (Fryday & Coppins Reference Fryday, Coppins, Nash, Gries and Bungartz2007); Maine, Michigan, Québec (Launis & Myllys Reference Launis and Myllys2014)), South America (Chile, specimens cited below), Central Africa (Rwanda (Czarnota Reference Czarnota2007)) and Réunion (Brand et al. Reference Brand, van den Boom and Sérusiaux2014). We also report a new locality in the near-coastal region of the Russian Far East.
Specimens examined
Japan (1): Kochi: Ino-cho town, Teregawa, S slope of Mt Kamegamori, path along the stream Shirai-dani, 33°45.09–16′N, 133°11.48–54′E, 750–770 m, humid old-growth, mixed deciduous/coniferous forest with e.g. Abies firma, Cryptomeria japonica, Quercus spp. and Tsuga sieboldii, in hollow at base of coniferous tree, 2006, Thor 21503 (UPS).—Russia: Khabarovskiy Krai: Komsomolsk-De Kastri route, Khomi Mountains, c. 20 air km E of Chernyy Mys, hills on N bank of the Salasu River, mixed coniferous/broad-leaved mature forest, 51°05.896′N, 138°46.303′E, 316 m, on rotten wood, 2009, Spribille (30498), Yakovchenko, Printzen, Kanz & Malashkina (GZU).—Chile: Region X: Rio Enco, between Choshuenco and Enco, 39°54′S, 72°9′W, 400 m, on underside of a fallen charred trunk, 1986, B. J. Coppins, D. J. Galloway, G. Guzman & P. W. James 6042 (E); ibid., in hollowed trunk of Nothofagus dombeyi, B. J. Coppins et al. 6043 (E).
Micarea inopinula (Nyl.) Coppins & T. Sprib. comb. nov.
MycoBank No.: MB 836556
Lecidea inopinula Nyl., Lich. Jap., 71 (1890).—Catillaria inopinula (Nyl.) Zahlbr., Cat. Lich. Univ. 4, 18 [as ‘inopulina’] (1926); type: Japan, Fuji, Itjigome [Itchigômé], ‘super truncos putrescentes inter Jungermannias’, 1879, E. Almquist (H-NYL 13003 = H9507484—lectotype, selected here, MBT 394864, material also in S).
Biatorina prasinella Jatta, Bull. Soc. Bot. Ital. 1911, 257 (1911).—Micarea prasinella (Jatta) I. M. Lamb, Lilloa 26, 413 (1953); type: Tasmania, [West Coast Range, near Macquarie Harbour], Dubbilbarril, Mt Lyell railway, on Jungermanniaceous hepatic on bark, 1899, W. A. Weymouth 977 (FI—holotype; BM—isotype!).
Lecidea inopinula, described by Nylander (Reference Nylander1890) from Mt Fuji, is an earlier name for the cephalodiate taxon widely known as Micarea prasinella, but it was overlooked in previous work on Micarea because the name has been maintained in Catillaria until now (Ohmura & Kashiwadani Reference Ohmura and Kashiwadani2018). A locality for this species in Japan extends the range of this charismatic, temperate rainforest species from Australasia (Tasmania (Lamb Reference Lamb1953); New Zealand (Kantvilas & Coppins Reference Kantvilas and Coppins2019)), North America (Alaska (Spribille et al. Reference Spribille, Pérez-Ortega, Tønsberg and Schirokauer2010, Reference Spribille, Fryday, Pérez-Ortega, Svensson, Tønsberg, Ekman, Holien, Resl, Schneider and Stabentheiner2020); Oregon (Andersen & Ekman Reference Andersen and Ekman2005), as ‘U.S.A.’, GenBank voucher AY567745)), South America (Chile (Kantvilas & Coppins Reference Kantvilas and Coppins2019)) and Europe (Scotland (Coppins Reference Coppins2008)). It was not discovered during the recent visits to Japan by G. Thor, so a diligent search of the forested slopes of Mt Fuji is required to see if it is extant in Japan; likewise for M. globulosella, also described from there.
Micarea inquinans (Tul.) Coppins
This lichenicolous species was reported as new to Japan by Zhurbenko et al. (Reference Zhurbenko, Frisch, Ohmura and Thor2015) from one single locality in Yamanashi Prefecture. One additional nearby locality from Nagano Prefecture is included here. Both findings are on the terricolous lichen Dibaeis arcuata (Stirt.) Kalb & Gierl in alpine regions.
Specimens examined
Japan (2): Honshu: Nagano Pref. (Shinano Prov.), Ueda-city, along the trail from Sugadaira Pastureland to Mt Neko-dake, stone field below the top, 36°32.721′N, 138°23.758′E, 2150 m, on exposed soil, 2012, Thor 28250 (TNS, UPS); Yamanashi Pref. (Kai Prov.), Kofu-city, along the trail between Odarumi Pass and Mt Kinpu, 35°52.194′N, 138°37.885′E, 2570 m, on soil, 2012, Thor 27977 (UPS).
Micarea lignaria (Ach.) Hedl. var. lignaria
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
A rare species found only at one locality in South Korea. First reported from South Korea by Aptroot & Moon (Reference Aptroot and Moon2015). Widely distributed, in Asia, Europe, Macaronesia, North and South America (specimen from Chile cited below), Australasia (Papua New Guinea (Aptroot et al. Reference Aptroot, Diederich, Sérusiaux and Sipman1997)), Réunion (Brand et al. Reference Brand, van den Boom and Sérusiaux2014) and the Subantarctic (South Shetland Islands (Czarnota Reference Czarnota2007)).
Specimens examined
South Korea (2): Cheju Island: the rim surrounding the crater of Mt Halla, 33°22′N, 126°33′E, 1850–1950 m, on mossy rock, 2001, Thor 17070, 17129 (UPS); ibid., on soil/plant debris on dead Abies koreana, Thor 17038 (UPS).—Chile: Region X: Parque Nacional Vicente Pérez Rosales, Petrohue, Rio Petrohue near falls, 41°8′S, 72°27′W, 500 m, amongst mosses on wet rocks with Toninia thiopsora, 1986, B. J. Coppins et al. 6074 (E).
Micarea lithinella (Nyl.) Hedl.
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
A rare species on siliceous stones, boulders and semi-exposed rocks found at three nearby localities on Hokkaido; 395–650 m. New to Japan; previously reported for Asia from China (Yunnan (Czarnota Reference Czarnota2007)) and Taiwan (Aptroot & Sparrius Reference Aptroot and Sparrius2003); elsewhere from Europe and North America (Czarnota Reference Czarnota2007).
Selected specimens examined
Japan (3): Hokkaido: Kitami Prov., Shari-gun, Shari-cho, Shiretoko National Park, NW slope of Shiretoko Peninsular, c. 10 km NE of Utoro town, along the trail from Iwaobetsu hot-spring hotel (Onsen) to Mt Rausu-dake, old-growth montane forest, 44°10′N, 145°09′E, 395 m, on a boulder, 2010, Thor 23690 (TNS, UPS); ibid., 44°06.243′N, 145°05.716′E, 400 m, on stones, 2010, Thor 24536 (TNS, UPS); ibid., 44°10′N, 145°10′E, 650 m, on rock, 2010, Thor 24804 (UPS).
Micarea melaena (Nyl.) Hedl.
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
A common species on bark and lignum of trees and stumps, especially conifers, in old-growth and managed forests; 620–1280 m. Widely distributed, in Asia, Europe, Macaronesia, North America, South America (Falkland Islands (Calvelo & Fryday Reference Calvelo and Fryday2006)), Australasia (Tasmania (Kantvilas & Coppins Reference Kantvilas and Coppins2019)) and the Subantarctic (South Shetland Islands (Czarnota Reference Czarnota2007)).
Selected specimens examined
Japan (14): Kyushu: Kagoshima Pref. (Ohsumi Prov.), Yakushima Island, 5 km S of Miyanoura, Shirotani-Unsuikyo Gorge, Kami-Yaku-cho, 30°22.5′N, 130°34′E, 640–700 m, on old Cryptomeria japonica [bark], 1994, Thor 12335, 12464 (TNS, UPS). Kochi: Ino-cho town, Teregawa, S slope of Mt Kamegamori, path along the stream Shirai-dani, humid old-growth, mixed deciduous/coniferous forest, 33°45.09–16′N, 133°11.48–54′E, 750–770 m, on moss on deciduous tree, 2006, Thor 21240 (UPS). Honshu: Yamanashi Pref. (Kai Prov.), Minamikoma-gun, Hayakawa-cho, 25 km W of Kofu town, along the small dirt road from the starting point 1 km S of the village Narada (and 100 m S of the dam) and c. 1.5 km towards SW until the road finish, deciduous forest with scattered, young, planted Cryptomeria japonica stands, on lignum of stump, 35°34′N, 138°18′E, 800–900 m, 1999, Thor 16493 (UPS). Hokkaido: Ishikari Prov., Kamikawa-gun, Kamikawa-cho, 200–400 m E of Obako Gorge tourist centre, at S shore of Niseicharomappu-gawa stream, mixed Abies sachalinensis/deciduous forest, 43°42′N, 143°01′E, on Picea jezoensis stump [lignum], 1994, Thor 14647 (TNS, UPS).
Micarea micrococca (Körb.) Gams ex Coppins
Description and illustrations
Czarnota & Guzow-Krzemińska (Reference Czarnota and Guzow-Krzemińska2009).
Habitat and distribution
A rare species on bark and lignum of trees, logs and stumps in old-growth or humid forest on Hokkaido; 220–1200 m. Reported as new to South Korea by Aptroot & Moon (Reference Aptroot and Moon2015). New to Japan. Its distribution outside Europe is uncertain following the recent revision of the Micarea prasina group (Czarnota & Guzow-Krzemińska Reference Czarnota and Guzow-Krzemińska2009; Guzow-Krzemińska et al. Reference Guzow-Krzemińska, Sérusiaux, van den Boom, Brand, Launis, Łubek and Kukwa2019; Launis et al. Reference Launis, Pykälä, van den Boom, Sérusiaux and Myllys2019a, Reference Launis, Malíček, Svensson, Tsurykau, Sérusiaux and Myllysb), although it has since been reported from Alaska (Spribille et al. Reference Spribille, Fryday, Pérez-Ortega, Svensson, Tønsberg, Ekman, Holien, Resl, Schneider and Stabentheiner2020). The M. micrococca complex is recognized by having apothecia up to 0.4 mm wide, rarely adnate, cream-white or greyish and by containing methoxymicareic acid. All Japanese collections have whitish, K− (lacking the Sedifolia-grey pigment) apothecia and spores 3–4.5 μm wide, and thus do not belong to the recently recognized M. czarnotae Launis et al. or M. pseudomicrococca Launis & Myllys (Launis et al. Reference Launis, Pykälä, van den Boom, Sérusiaux and Myllys2019a, Reference Launis, Malíček, Svensson, Tsurykau, Sérusiaux and Myllysb).
Specimens examined
Japan (3): Hokkaido: Kushiro Prov., Kushiro-shi, 53 km NNW of Kushiro city, Mt O-Akan dake, along the trail from the road at the easternmost end of Lake Akan to the top of the mountain, old-growth montane forest with e.g. Abies sachalinensis, Acer sp., Larix leptolepis, Magnolia sp., Picea sp. and Quercus crispula, 43°27′N, 144°09′E, 600–1100 m, on log, 2010, Thor 26038 (UPS); Kitami Prov., Shari-gun, Shari-cho, Shiretoko National Park, NW slope of Shiretoko Peninsular, c. 10 km NE of Utoro town, along the trail from Iwaobetsu hot-spring hotel (Onsen) to Mt Rausu-dake, old-growth subalpine forest dominated by Betula ermannii, 44.08942°N, 145.11675°E, 1021 m, on bark of Betula ermannii, 2010, Thor 24505 (UPS); ibid., c. 7 km NE of Utoro town, along the trail from Iwaobetsu hot-spring hotel (Onsen) to Mt Rausu-dake, old-growth lowland forest dominated by Abies sachalinensis and Quercus crispula, 44.10947°N, 145.08242°E, 220 m, on stump, 2010, Thor 24879 (UPS).
Micarea misella (Nyl.) Hedl.
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
A rare species on lignum of logs and stumps, and also on bark of horizontal trunk of Betula ermannii; c. 900–1030 m. New to Japan. Previously reported in Asia from eastern Russia (Konoreva et al. Reference Konoreva, Chesnokov and Poryadina2018), Bhutan (Aptroot & Feijen Reference Aptroot and Feijen2002), Taiwan (Aptroot & Sparrius Reference Aptroot and Sparrius2003) and South Korea (Aptroot & Moon Reference Aptroot and Moon2015); elsewhere from Europe and North and South America (Czarnota Reference Czarnota2007).
Specimens examined
Japan (3): Honshu: Ibaraki Pref. (Hitachi Prov.), Hirosawa, 11.5 km N of Tsukuba-city, S slope of Mt Tsukuba, 36°11′N, 140°06′E, on log at roadside, 1994, Thor 11735 (UPS, apothecia and pycnidia); Aomori Pref. (Mutsu Prov.), S of Suiren-numa pond, Mt Hakkoda, 40°37′N, 140°55′E, 900 m, on a c. 2 m high stump [lignum] in open mixed coniferous/deciduous forest, 1994, Thor 11999 (TNS, UPS, apothecia and pycnidia). Hokkaido: Kitami Prov., Shari-gun, Shari-cho, Shiretoko National Park, NW slope of Shiretoko Peninsular, c. 10 km NE of Utoro town, along the trail from Iwaobetsu hot-spring hotel (Onsen) to Mt Rausu-dake, old-growth subalpine forest dominated by Betula ermannii, 44.08905°N, 145.11694°E, 1032 m, on Betula ermannii, 2010, Thor 24678 (UPS, pycnidia only).—South Korea (1): Cheju Island: Namceju-gun, Namwon-up, along the Songpanak trail on the E slope of Mt Halla, above the Azalea field shelter, Abies koreana forest with scattered deciduous trees, 33°31′N, 126°32′E, 1500–1700 m, on snag, 2001, Thor 17500 (UPS, pycnidia only).
Micarea nitschkeana (J. Lahm ex Rabenh.) Harm.
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
Rare, only observed once on a snag. New to Asia; elsewhere from Europe and North America (California (Fryday & Coppins Reference Fryday, Coppins, Nash, Gries and Bungartz2007)). The report from Australia (Tasmania (Czarnota Reference Czarnota2007)) is based on a misidentification of M. denigrata (G. Kantvilas, personal communication). A recent report of the species as new to Asia was corrected in a subsequent publication (Konoreva et al. Reference Konoreva, Chesnokov and Poryadina2018).
Specimen examined
South Korea (1): Gangwon Prov.: Yangyang-gun, Ser-myun, Osaeck-ri, the southern part of the massif Sorak Mts, Sorak-san National Park, S slope of Mt Dachong, along the trail from the shelter c. 500 m WNW of the top of Mt Dachong to the village at Hangyeryong Pass, c. 1–3 km SW of the shelter, open mixed coniferous/deciduous forest and siliceous rocky outcrops, 38°07.06–10′N, 128°27.25–26′E, 1400–1600 m, on snag of Abies koreana, 2006, Thor 20430 (UPS).
Micarea peliocarpa (Anzi) Coppins & R. Sant.
Descriptions and illustrations
Coppins (Reference Coppins1983), Czarnota (Reference Czarnota2007).
Habitat and distribution
Widely occurring and common on bark and lignum of old trees, especially conifers and Betula, stumps and logs, and shaded siliceous rocks, also on plant remains in more open situations; 10–1650 m. Widely distributed, in South Korea and Japan (Kondratyuk et al. Reference Kondratyuk, Lőkös, Tschabanenko, Haji Moniri, Farkas, Wang, Oh and Hur2013; Ohmura & Kashiwadani Reference Ohmura and Kashiwadani2018), though not yet found in eastern Siberia (Konoreva et al. Reference Konoreva, Chesnokov and Poryadina2018), Europe, Macaronesia, North, Central and South America, Africa (Madagascar (Aptroot Reference Aptroot1990)), Réunion (Brand et al. Reference Brand, van den Boom and Sérusiaux2014) and Australasia (Czarnota Reference Czarnota2007; Kantvilas & Coppins Reference Kantvilas and Coppins2019).
Selected specimens examined
Japan (27): Kyushu: Kagoshima Pref. (Ohsumi Prov.), Yakushima Island, 7.5 km W of Anbo, Yakusugi-land, Yaku-cho, dense forest, 30°18′N, 130°34′E, 950–1100 m, on shaded mossy rock, 1994, Thor 12387 (TNS, UPS). Kochi: Ino-cho town, Teregawa, NE slope of Mt Iwaguro-yama, along path from the road, tall, humid, old-growth, mixed deciduous/coniferous forest with e.g. Abies homolepis, Fagus crenata and Quercus spp., 33°45.10′N, 133°09.54′E, 1300 m, on log, 1996, Thor 21308 (TNS, UPS); ibid., on base of coniferous tree, Thor 21314 (TNS, UPS). Honshu: Okayama Pref. (Kii Prov.), Maniwa-gun, Kawakamimura, shrine 2 km SW of Gohara, 35°15′N, 133°38′E, 590 m, on Pinus bark, 1994, Thor 12230, 12232 (TNS, UPS). Hokkaido: Kitami Prov., Rishiri-to Island, Rishiri-gun, Rishirifuji-cho, along the trail from Rishiri-hokuroku campsite 2.5 km S of Sakae town to Himenuma Pond, old-growth lowland forest dominated by Abies sachalinensis and Quercus crispula, 45°13′N, 141°14′E, 140–360 m, on Betula bark, 1995, Thor 13819 (TNS, UPS).—South Korea (2): Cheju Island: Namcheju-gun, Namwon-up, along the Songpanak Trail on the E slope of Mt Halla, above the Azalea field shelter, Abies koreana forest with scattered deciduous trees, 33°21′N, 126°32′E, 1500–1700 m, on lignum of living Abies koreana, 2001, Thor 17563 (UPS).
Micarea prasina Fr.
The presence of Micarea prasina s. str. from Japan needs to be confirmed. No collection was found in this study.
Micarea synotheoides (Nyl.) Coppins
Type citations, descriptions and illustrations
Coppins (Reference Coppins1983: fig. 31D, ascospores from lectotype), Czarnota (Reference Czarnota2007).
Notes
This species was originally collected by Almquist in 1879 from Itjgome [on slopes of Mt Fuji] and described twice by Nylander, as Lecidea synotheoides Nyl. (Nylander Reference Nylander1890) and as L. longella Nyl. (Nylander Reference Nylander1900). None of Almquist's collections under either name have mesoconidia-containing pycnidia, but fortunately most of the 26 recent collections from Japan have such pycnidia in abundance, and indeed the species was often encountered as ‘sterile’ with numerous sessile to short-stalked pycnidia (Czarnota Reference Czarnota2007: fig. 56A) and no apothecia. This answers the problem discussed by Czarnota (Reference Czarnota2007) regarding the possibility that two distinct species may be included in the concept of M. synotheoides as adopted by Coppins (Reference Coppins1983). The result is that the material from westernmost Europe (and probably some from north-western America) requires a new name; it is introduced below as M. longispora. Micarea synotheoides has recently been recorded from Australia (New South Wales (McCarthy & Elix Reference McCarthy and Elix2020b)). The specimen concerned lacked both mesoconidia and microconidia, although it did have macroconidia-containing pycnidia which have not been previously documented for either M. longispora or M. synotheoides.
Habitat and distribution
On bark of a variety of trees and shrubs, including Abies, Alnus, Betula, Picea and Sorbus, in old-growth and subalpine forest; c. 220–2000 m. A northern species mainly occurring on Hokkaido, rarely on Honshu and only at high altitudes.
Selected specimens examined
Japan (26): [Cited specimens all with both apothecia and mesopycnidia] Honshu: Toyama Pref. (Etchu Prov.), 33 km ESE of Toyama, Tateyama-cho, Nakasinkawa-gun, Midagahara, along the path to Tateyama crater 500 m SSE of Midagahara bus stop, subalpine, open mixed deciduous/Abies mariesii forest, 36°34′N, 137°34′E, 1960–2000 m, on shaded Abies mariesii, 1994, Thor 12674 (TNS, UPS). Hokkaido: Ishikari Prov., Higashikawa-cho, 140 km NE of Sapporo, W slope of Mt Asahidake, village of Asahidake Onsen, 200–600 m SW of the ropeway station, along the trail ‘Kumagera course’ from Lodge Nutapukaushipe to the ropeway station, old-growth montane forest with e.g. Abies sachalinensis, Acer spp., Alnus sp. and Picea sp., 43°39.0395′N, 142°47.4070′E, 1100 m, on Picea, 2010, Thor 25863 (TNS, UPS); Kitami Prov., Shari-gun, Shari-cho, Shiretoko National Park, NW slope of Shiretoko Peninsula, c. 10 km NE of Utoro town, along the trail from Iwaobetsu hot-spring hotel (Onsen) to Mt Rausu-dake, old-growth subalpine forest dominated by Betula ermannii, 44.09406°N, 145.10672°E, 811 m, on Sorbus commixta, 2010, Thor 24324 (TNS, UPS); ibid., 44.08376°N, 145.12332°E, 1220 m, on Alnus maximowiczii, 2010, Thor 25447 (TNS, UPS).
Micarea xanthonica Coppins & Tønsberg
Description and illustrations
Coppins & Tønsberg (Reference Coppins and Tønsberg2001).
Habitat and distribution
A rare species found once on bark in humid forest. New to Asia; elsewhere from oceanic Western Europe and western North America. When described, Micarea xanthonica was shown to belong to a list of species that apparently have a Western European (including Macaronesia for some species) and north-western North American disjunct distribution (Coppins & Tønsberg Reference Coppins and Tønsberg2001). The lichen flora in many parts of NW Asia is still insufficiently studied and more species known currently to have a European and North American distribution are certain to be found in Asia (and maybe elsewhere as well).
Specimen examined
Japan (1): Kochi: Tsuno-mura village, Mt Tongu-kogen, path along ridge, humid, open, mixed deciduous/coniferous forest with e.g. Abies homolepis, Quercus spp. and Weigela decora, 33°28.38–50′N, 133°00.20–55′E, 1360–1485 m, on tree, 2006, Thor 21467 (UPS, no apothecia).
Additional new species
Micarea longispora Coppins sp. nov.
MycoBank No.: MB 836555
Similar to M. synotheoides but with longer ascospores (14–35(–43) vs 14–26 μm), 1–7(–11)-septate, immersed pycnidia and longer mesoconidia (4.5–6 vs 3.5–5 μm).
Type: Scotland, Argyll (V.C. 98), Loch Fyne, 8 km E of Inverary, Ardkinglass, 27(NN)/17.10 [56°14′50.24ʺN, 4°57′13.95ʺW], elev. 30 m, on Quercus bole, 18 November 1977, B. J. Coppins 4035 (E—holotype; UPS—isotype).
Notes
Micarea longispora differs from M. synotheoides in having somewhat longer ascospores, often with more septa, immersed mesoconidia-containing pycnidia, and longer mesoconidia, 4.5–6 × 1.2–1.5 μm. For a full English description see Coppins (Reference Coppins1983: 188–189, figs 31E, 43D & E) and for a habit photograph see Czarnota (Reference Czarnota2007: fig. 56B).
As previously discussed by Czarnota (Reference Czarnota2007), the material of Micarea synotheoides s. lat. from strongly oceanic areas (e.g. western Scotland and Norway) differs from the type and other material from Japan in having longer ascospores that are often more than 3-septate. Material from Central Europe (Czech Republic, Poland and Ukraine) has shorter ascospores, as in the Japanese collections, and additionally has sessile to short-stalked, mesoconidia-containing pycnidia, with shorter conidia. As the 19th century type material of M. synotheoides does not have any mesoconidia-containing pycnidia, it was not possible to assign either the Western or Eastern European material to M. synotheoides s. str. However, such pycnidia are amply present in the recently collected material from Japan, so it is now possible to identify the Central European material as M. synotheoides s. str.
Habitat and distribution
The species is usually corticolous though often overgrowing bryophytes, in mainly old-growth woodlands, on ‘acid-barked’ coniferous and deciduous trees and shrubs. It is very occasionally found on the lignum of rotting tree trunks. The species occurs in oceanic areas of Europe from Norway, the western British Isles, north Spain and Macaronesia, and probably equivalent areas along the Pacific coast from Washington to British Columbia, but material named as ‘M. synotheoides’ from North America requires critical examination to confirm this.
Selected additional specimens examined
Great Britain: England: V.C. 3, South Devon, Dartmoor National Park, Cornwood, Dendles Wood NNR, 20(SX)/61.61, 1991, Coppins [13869] & O'Dare (E).—Ireland: V.C. H35, West Donegal: Glenveagh National Park, woodland on SE shore of Lough Veagh, C/00.19, c. 60 m, on Betula, 1991, Coppins [14694] & H. Fox (E); ibid., on Pinus, Coppins 14709 (E).—Spain: Tenerife: Aquamansa, near Los Organos, 1100 m, on Erica arborea, 1978, Topham s. n. (E).
Acknowledgements
We thank Paweł Czarnota for earlier discussions relating to Micarea synotheoides, the curators at S and UPS, and André Aptroot for making available his collection of Micarea rubioides from the Philippines. GT's stay in Japan in 1994–1996 was funded by a postdoctoral fellowship from the Japan Society for the Promotion of Science (JSPS) and a fellowship for Priority area research in Japan from the JSPS. The field trip in South Korea by GT in 2006 was financially supported by a grant to KHM (no. 052-052-040) from the Core Environmental Technology Development Project for Next Generation funded by the Ministry of Environment of the Korean Government. TS acknowledges funding from the National Sciences and Engineering Research Council of Canada (NSERC). GT wishes to thank Dr Akira Mori, Yokohama National University, for inviting him to participate in the inventory in Shiretoko, Hokkaido in 2010 and for arranging collecting permits.
We are delighted to be able to offer this paper in celebration of Peter Crittenden's 70th birthday and in recognition of his many contributions to lichenology, in particular his dedicated stewardship of The Lichenologist.
Author ORCIDs
Brian J. Coppins, 0000-0001-9464-0495; Toby Spribille, 0000-0002-9855-4591.
