Научная статья на тему 'A REVISION OF THE LICHEN GENUS PLATISMATIA (PARMELIACEAE) IN RUSSIA, WITH A KEY TO THE SPECIES'

A REVISION OF THE LICHEN GENUS PLATISMATIA (PARMELIACEAE) IN RUSSIA, WITH A KEY TO THE SPECIES Текст научной статьи по специальности «Биологические науки»

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Ключевые слова
FOLIOSE LICHENS / HERBARIUM / COMMANDER ISLANDS NATURE AND BIOSPHERE RESERVE / RUSSIAN FAR EAST

Аннотация научной статьи по биологическим наукам, автор научной работы — Kuznetsova E.S., Stepanchikova I.S., Skirina I.F., Chesnokov S.V., Himelbrant D.E.

The paper presents the results of the study on Platismatia species in Russia. The genus Platismatia counts 11 species, distributed mainly in the Pacific region, with some endemics of western North America and northeastern Asia. Six species were known from Russia by the beginning of our studies, but a revision of the herbarium collections showed that three species (P. erosa, P. herrei, and P. lacunosa) were reported erroneously. Based on morphological and chemotaxonomical (HPTLC) examination of the herbarium collections and our own material, as well as the study of relevant literature, four species of Platismatia are here accepted for Russia: P. glauca, P. interrupta, P. lacunosa, and P. norvegica. Platismatia lacunosa is reported here as a new for Russia from the Commander Islands. The distribution of P. glauca and P. interrupta is clarified. A key to all species of the genus reported from Russia and brief descriptions of the Russian species are presented.

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Текст научной работы на тему «A REVISION OF THE LICHEN GENUS PLATISMATIA (PARMELIACEAE) IN RUSSIA, WITH A KEY TO THE SPECIES»

A revision of the lichen genus Platismatia (Parmeliaceae) in Russia,

with a key to the species

E. S. Kuznetsova1, 2, I. S. Stepanchikova1, 2, I. F. Skirina3, S. V. Chesnokov2, 4, D. E. Himelbrant1, 2

1St. Petersburg State University, St. Petersburg, Russia 2Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia 3Pacific Institute of Geography of the Far Eastern Branch of the Russian Academy of Sciences,

Vladivostok, Russia

4Botanical Garden-Institute of the Far Eastern Branch of the Russian Academy of Sciences,

Vladivostok, Russia Corresponding author. E. S. Kuznetsova, [email protected]

Abstract. The paper presents the results of the study on Platismatia species in Russia. The genus Platismatia counts 11 species, distributed mainly in the Pacific region, with some endemics of western North America and northeastern Asia. Six species were known from Russia by the beginning of our studies, but a revision of the herbarium collections showed that three species (P. erosa, P. herrei, and P. lacunosa) were reported erroneously. Based on morphological and chemotaxonomical (HPTLC) examination of the herbarium collections and our own material, as well as the study of relevant literature, four species of Platismatia are here accepted for Russia. P. glauca, P. interrupta, P. lacunosa, and P. norvegica. Platismatia lacunosa is reported here as a new for Russia from the Commander Islands. The distribution of P. glauca and P. interrupta is clarified. A key to all species of the genus reported from Russia and brief descriptions of the Russian species are presented.

Keywords: Platismatia lacunosa, foliose lichens, herbarium, Commander Islands Nature and Biosphere Reserve, Russian Far East.

Ревизия лишайников рода Platismatia (Parmeliaceae) в России с ключом для определения видов

Е. С. Кузнецова1, 2, И. С. Степанчикова1, 2, И. Ф. Скирина3, С. В. Чесноков2, 4, Д. Е. Гимельбрант1, 2

1Санкт-Петербургский государственный университет, Санкт-Петербург, Россия 2Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия 3Тихоокеанский институт географии ДВО РАН, Владивосток, Россия 4Ботанический сад-институт ДВО РАН, Владивосток, Россия Автор для переписки. Е. С. Кузнецова, [email protected]

Резюме. В статье представлены результаты исследования рода Platismatia в России. Род Platismatia насчитывает 11 видов, распространенных в основном в Тихоокеанском регионе, некоторые виды являются эндемиками западной части Северной Америки и Северо-Восточной Азии. До настоящего момента для территории России было приведено 6 видов рода Platismatia, однако ревизия гербарных коллекций показала, что имеющиеся в литературе указания трех

https://doii.org/1031111/Knr/2021551.179

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видов (P. erosa, P. herrei и P. lacunosa) являются ошибочными. По результатам морфологического и хемотаксономического анализа гербарных материалов, коллекций авторов и данных литературы для территории России приведены четыре вида рода Platismatia: P. glauca, P. in-terrupta, P. lacunosa и P. norvegica. Platismatia lacunosa найдена впервые на территории России на Командорских островах. Уточнены сведения о распространении двух видов — P. glauca и P. interrupta. В статье приведен ключ для определения видов рода Platismatia, встречающихся в России и ранее опубликованных для ее территории, а также дано краткое описание российских видов.

Ключевые слова: Platismatia lacunosa, гербарий, листоватые лишайники, Командорский заповедник, Российский Дальний Восток.

The biodiversity of the parmelioid lichens is still far from completely understood in many regions of Russia. One of the problematic genera is Platismatia (L.) W. L. Culb. et C. F. Culb. Although all the Platismatia species are of significant size and easy to notice, the identification of the taxa is still a complicated task.

Platismatia is a genus including large, mainly epiphytic species. It was described as a separate genus of cetrarioid lichens by William L. Culberson and Chicita F. Culberson in 1968 based on morphological and chemical characters, with the presence of caperatic acid in the medulla being the main distinguishing feature (Culberson, Cul-berson 1968; Obermayer, Randlane, 2012). However, the chemical delimitation of the genus is quite universal, but not exhaustive: e. g., one chemotype of Platismatia erosa W. L. Culb. et C. F. Culb. lacks caperatic acid (Obermayer, Randlane, 2012). Phylo-genetically, Platismatia does not fall into any of the major clades of Parmeliaceae and remains a genus with "uncertain affinities" within the family (Thell et al, 2012; Divakar et al., 2017).

Platismatia species are characterized by usually foliose thalli with broad, loosely attached, ascending lobes. Upper surface gray, sometimes brown in exposed localities, with more or less developed (sometimes lacking) pseudocyphellae, often wrinkled. Isidia (and, more rarely, soredia) present in some species, mostly marginal, can be also laminal. Lower surface black at the center and often brown or white at the margins, speckled in some species, with a few rhizines. Cortex paraplectenchymatous. Apothecia marginal or submarginal, commonly perforate. Asci of Lecanora-type, 8-spored. Spores simple, colorless, ellipsoid or subglobose, 3.5-10.0 |im in size. Pycnidia rare, immersed, marginal (Culberson, Culberson, 1968; Thell, 2011), conidia sublageniform.

The lichen substances found in Platismatia spp. comprise atranorin, caperatic acid, in some species also fumarprotocetraric acid (Thell, 2011), additionally chloroatra-norin, jackinic, pannaric, and pseudoplacodiolic acids, and rarely yellow pigment(s?) (Culberson, Culberson, 1968; Obermayer, Randlane, 2012).

The genus Platismatia counts 11 species which have mainly Amphi-Pacific distribution with the center of diversity located in the Northern Hemisphere. Only P. glauca (L.) W. L. Culb. et C. F. Culb., the type species of the genus, occurs on all continents except Australia (Culberson, Culberson, 1968; Lumbsch et al., 2011; Allen et al., 2012; Obermayer, Randlane, 2012).

Until now, 6 taxa have been reported from Russia (Spisok..., 2010): Platismatia erosa, P. glauca, P. herrei (Imshaug) W. L. Culb. et C. F. Culb., P. interrupta W. L. Culb. et C. F. Culb., P. lacunosa (Ach.) W. L. Culb. et C. F. Culb., and P. norvegica (Lynge) W. L. Culb. et C. F. Culb. However, the revision of herbarium collections and study of relevant literature have shown that the records of three species are probably erroneous: these are P. herrei and P. erosa reported from the south of the Russian Far East (Skirina, 1995; Tchabanenko, 2002; Skirin, Skirina, 2012; Yakovchenko et al., 2013); moreover, the specimens of P. lacunosa from the Russian Arctic (Elenkin, 1907; Spisok., 2010) in LE appeared to be misidentified as well.

Morphologically closest to the genus Platismatia are Cetrelia W. L. Culb. et C. F. Culb., Asahinea W. L. Culb. et C. F. Culb., Esslingeriana Hale et M. J. Lai, in part also Tuckermannopsis Gyeln. and Punctelia Krog. The species of Cetrelia are chemically variable, but they always lack caperatic acid. Other distinguishing characters are not exclusive, however, they can help in identification: the majority of Cetrelia occurring in Russia have marginal soredia and distinct whitish pseudocyphellae on the lower surface; the lobes of Cetrelia are not reticulate; some species have C+ red medullar reaction or medulla UV+ white (Randlane, Saag, 1991). Some Cetrelia are superficially very similar to Platismatia, and chromatography is needed for certain identification of the species. For example, the isidiate Cetrelia braunsiana (Müll. Arg.) W. L. Culb. et C. F. Culb., occurring in Asia, can be confused with Platismatia interrupta, but differs from the latter by KC+ red medullary reaction; C. braunsiana produces atra-norin in the cortex, and alectoronic, a-collatolic, 4-O-methylphysodic and physodic acids in the medulla (Luo et al., 2007). Most of Tuckermannopsis species are smaller in size, have narrower lobes with no isidia, and are greenish-brown, not gray. Other distinguishing features of Tuckermannopsis spp. are the absence of pseudocyphellae or whitish spots on the upper surface; also Tuckermannopsis never contains caperatic acid in medulla and rarely has lichen compounds in the cortex (Ahti, Thell, 2011). Esslingeriana idahoensis (Essl.) Hale et M. J. Lai (North American monotypic genus, at the moment not known in Eurasia) is very close to Platismatia in morphology, but its medulla lacks caperatic acid and contains endocrocin, and also, in contrast to Platis-matia, E. idahoensis usually has abundant pycnidia (see Goward et al., 1994). Asahinea can be distinguished by the absence of rhizines and the presence of alectoronic and a-collatolic acids in the medulla instead of caperatic acid. Punctelia spp. are usually more attached to the substrate, lack isidia, and contain lecanoric or gyrophoric acid (C+ red) in the medulla. Some lichens containing usnic acid, e. g. Nephromopsis laureri (Kremp.) Kurok., can also resemble Platismatia, but they can be easily distinguished by the chemistry and yellowish color of the thallus.

Material and Methods

The treatment is based on published data, studies of own material deposited in LE and LECB, and herbarium collections in H, INEP, LE, MSK, VBGI, VGEO. Specimens in TU were checked by Professor Tiina Randlane. About 40 samples were

subjected to chemical studies based on standard high-performance thin-layer chromatography (HPTLC) using solvent systems A and C (Culberson, 1972; Orange et al., 2001). We used standard color reactions detected with 10% potassium hydroxide (K), sodium hypochlorite (C), K followed by C on the same fragment (KC), para-phe-nylenediamine (P), and operated long-wave (365 nm) UV lamp (CAMAG UV Cabinet 4) (Smith et al., 2009). More than 150 herbarium specimens from the study area were examined, but only selected ones are listed in detail.

Results The species

Platismatia glauca (L.) W. L. Culb. et C. F. Culb., 1968, Contr. U. S. Natl. Herb. 34(7): 530.

= Lichen glaucus L., 1753, Sp. Pl. 2: 1148. = Lobaria glauca (L.) Hoffm., 1796, Deutschlands Flora oder botanisches Taschenbuch. Zweiter Theil für das Jahr 1795. Cryptogamie: 149. = Certaria glauca (L.) Ach., 1803, Method. Lich.: 296. = Physcia glauca (L.) DC., 1805, Flore française 2: 401. = Platysma glaucum (L.) Frege, 1812, Deutsches Botanisches Taschenbuch für Liebhaber der deutschen Pflanzenkunde 2: 167. = Parmelia glauca (L.) Hepp, 1824, Lichenen-Flora von Würzburg: 23. — Lecto-type: Europe, LINN 1273.139.

Platismatia glauca is a highly variable lichen recognized by weakly attached thallus with broad ascending lobes with simple or coralloid isidia, and sometimes with granular soredia on the lobe margins or rarely on the upper surface. Some lobes can become nearly fruticulose and coralloid. Upper surface smooth or slightly reticulate-wrinkled, gray, brown or blackish, without distinct pseudocyphellae but often somewhat maculate, lower surface brown to black, paler to even white close to the margins. Apo-thecia rare, to 10 mm diam., submarginal, disc brown, sometimes perforate; asci IKI+ blue/blue-green, subhymenium IKI+ purple; spores ellipsoid to ovoid, 3-4 x 5-8 |im. Pycnidia marginal, immersed, very rare.

Chemistry. Upper cortex contains atranorin, K+ yellow, KC, C, and P-, IKI+ purple. Medulla contains caperatic acid, K, KC, C, and P-, UV-. Some authors report the presence of pseudoplacodiolic acid and chloroatranorin (Obermayer, Randlane, 2012), but in our material only atranorin and caperatic acid were revealed by HPTLC.

Ecology. Corticolous on conifers, birch and other deciduous trees, sometimes on wood, soil, and rocks, inhabits various biotopes, especially common in taiga zone, also can be found in deciduous forests and tundras (including mountain tundras).

Distribution. Platismatia glauca is a cosmopolitan species, recorded on all continents except Australia (Culberson, Culberson, 1968; Lumbsch et al., 2011; Allen et al., 2012; Obermayer, Randlane, 2012). It has been reported throughout almost all regions of the forest belt of Russia (e. g., Tchabanenko, 2002; Blinkova, 2004; Poryadina, 2005; Fadeeva et al., 2007; Skirina, 2007, 2016; Korchikov, 2011; Davydov, 2014; Selivanov et al., 2015). The species has repeatedly been cited for the south of the Russian Far

East (Tchabanenko, 2002; Tchabanenko et al, 2002; Skirina, 2007, 2016). However, in course of active studies in recent years many specimens of the genus Platismatia were collected in Primorye Territory, Sakhalin, and the Kuril Islands (LE, H), and they all belong to P. interrupta. In addition, all the revised herbarium specimens from these regions previously assigned to P. glauca (LE, VGEO, VBGI) we also re-identified as P. interrupta. There are no certain records of P. glauca in Japan and Southeast Asia (Culberson, Culberson, 1968; Buaruang et al., 2017; Ohmura, Kashiwadani, 2018), but it has been recorded in China (Wei, 1991). In Siberia the species has been recorded from Khanty-Mansi Autonomous Area (LE), Sverdlovsk Region (LE), Tomsk Region (Kovaleva, 2006), Chelyabinsk Region (Urbanavichene, 2011), Novosibirsk Region (Romanova, Sedelnikova, 2010), Altai Territory (Davydov, 2014; LE), Kemerovo Region (Sedelnikova, 1990), republics of Altai (LE), Khakassia (Sedelnikova, 2001), Tyva (Sedelnikova, 1985), Buryatia (Sedelnikova, 2001; TU), and Krasnoyarsk Territory (Sedelnikova, 2001). All records of P. glauca from Yakutia (Fesko, 1990; Po-ryadina, 2005, 2007) are erroneous and refer to Cetrelia cetrarioides (Delise et Duby) W. L. Culb. et C. F. Culb. (L. N. Poryadina, pers. comm.).

Notes. Platismatia wheeleri Goward, Altermann et Bjork segregated from P. glauca in 2011 (Lumbsch et al., 2011) is distinguished by strongly undulate marginal soralia, erumpent from the medulla, and absence of isidia. Currently known only from North America and Slovakia (Lumbsch et al., 2011; Allen et al., 2012), probably can be found in Russia as well.

Selected specimens examined. Murmansk Region: Khibiny Mountains, NE slope of the Yukspor Mountain, forest belt, dry branches of Picea sp., 1930, Rassadina, LE L-15404. Nenets Autonomous Area: NE part of Malozemelskaya Tundra, N end of Nenetskaya Gryada, yernik (Betula shrubs) on the slope, branches of Betula nana L., 15 VIII 1998, Lavrinenko, LE L-15397. Kostroma Region: vicinity of Kologriv, spruce forest, dry Picea sp., 26 VIII 1929, Ladyzhenskaya, LE L-15406. Krasnodar Territory: Lagonaki Upland, left bank of the Molochka River, bark of Abies sp., 23 VII 1980, Krivoro-tov, LE L-15412. Crimea: Crimean Nature Park, Chyornaya Mountain, 18 km N of Alushta, bark of Fagus sp., 28 VII 1953, Vasil'kov, LE L-15399. Sverdlovsk Region: Severouralsk District, vicinity of Vsevolodo-Blagodatskoe, branches of Picea sp., 20 VIII 1962, Savicz, LE L-15396.

Platismatia interrupta W. L. Culb. et C. F. Culb., 1968, Contr. U. S. Natl. Herb. 34(7): 539. — Holotype: Japan, Prov. Shinano (Pref. Nagano), 1952, Asahina, TNS.

The species can be easily distinguished from other Platismatia species by the presence of abundant large conspicuous pseudocyphellae on the smooth or weakly reticulate-wrinkled gray upper surface. Lower surface of P. interrupta lacks pseudocyphellae, in contrast to P. erosa which has plenty of punctiform pseudocyplellae on the lower side of the thallus. Isidia simple, cylindrical or globose, sometimes disintegrating into soredia. Apothecia and pycnidia unknown.

Chemistry. Upper cortex contains atranorin, K+ yellow, KC, C, and P-, IKI- or pale purple. Medulla contains caperatic acid and unidentified yellow pigment (the latter not seen in the revised specimens), K, KC, C, and P-, UV-.

Ecology. Corticolous on bark of coniferous and deciduous trees, lignicolous, rarely terricolous, inhabits various forest biotopes.

Distribution. Outside of Russia P. interrupta is known only from Japan (Ohmu-ra, Kashiwadani, 2018). It is a widespread and common species in the southern Russian Far East. It has been confirmed from Khabarovsk Territory (H, LECB), Amur Region (LE), Jewish Autonomous Region (VGEO), Sakhalin Region (LE), and Pri-morye Territory (Tchabanenko et al, 2002; Skirina, 2016; LE). There are also several records of the species for Irkutsk Region and Republic of Buryatia (Parn, Trass, 1990; Krasnaya..., 2010; TU).

Selected specimens examined. Amur Region: Zeysky Strict Nature Reserve, the main watershed of the ridge in the upper Bolshaya Erakinga River, 54°07'12.7"N, 126°56'02.4"E, spruce forest, bark of Betula ermanii Cham. subsp. lanata (Regel) A. K. Skvortsov, 6 IX 2016, Dudov, LE L-15380. Khabarovsk Territory: Ulchsky District, vicinity of Mariinsky Reid, 51°42'N, 140°12'E, relict dunes, on soil, VIII 2012, Tugi, LE L-15409; ibid, Chikhaeva Bay, Davydova Cape, spruce forest, bark of coniferous tree, 9 IX 2001, Skirina, VGEO 13970 (det. Skirina sub P. glauca); Komsomolsk District, 20 km SSW from Boktor village, Gorin River basin, interfluve of the Muolgu and Koldka rivers, 50°58'00.4"N, 137°18'04.0"E, 202 m a. s. l., coniferous forest with birch, bark of Pinus sp., 18 VIII 2011, Yakovchenko, VBGI Fs17; Bikin District, Kukansky Ridge, 950 m a. s. l., coniferous forest, bark of Abies sp., 23 VIII 2012, Galanina, INEP (det. Galanina sub P. herrei). Jewish Autonomous Region: Bastak Strict Nature Reserve, Bogdyr Mountain, 800-1150 m a. s. l., coniferous forest, bark of Abies sp., 18 VII 2003, Skirina, Skirin, VGEO 15847 (det. Skirina sub P. glauca); ibid,, 800-1100 m a. s. l., coniferous forest, on bark of Abies sp., Picea sp., and wood of fallen trunks, 18 VII 2003, Skirina, Skirin, VGEO 15887 (det. Skirina sub P, norvegica); ibid,, 800-1000 m a. s. l., coniferous forest, on wood of fallen trunks, 18 VII 2003, Skirina, Skirin, VGEO 15889 (det. Skirina sub P, erosa). Primo-rye Territory: Shkotovsky District, Ussuriisky Strict Nature Reserve, watershed of Koryavaya Pad' and the Suvorovka River, fir-spruce forest, deadwood, 1975, Efimova, VGEO 9414 (det. Guseva sub P. erosa); Terney District, Sikhote-Alinsky Strict Nature Reserve, Lysaya Mountain, 1000 m a. s. l., shrubs of Siberian dwarf pine, bark of Pinuspumila (Pallas) Regel, 23 VI 1977, Skirina, VGEO 5924 [det. Guseva sub P. erosa and published in Skirina (1995)], 9415 [det. Guseva, Skirina sub P, norvegica and published in Skirina (1995)]; ibid,, Sikhote-Alin, 45°42'16.5"N, 136°40'00.6"E, pine forest on the slope, dead Pinus koraiensis Siebold et Zucc., 13 VIII 2010, Kuznetsova, LE L-15374; Dalnegorsk District, valley of the Rudnaya River, oak forest, bark of Betula sp., 1981, Skirina, VGEO 5924 (det. Guseva, Skirina sub P. erosa); Partizansky District, pass between Golets and Lasovaya mountains, 1130 m a. s. l., coniferous forest, bark of Abies sp., 18 VIII 2009, Skirina, VGEO 26098 (det. Skirina sub P, norvegica), 26125 (det. Skirina sub P, glauca). Sakhalin Region: Sakhalin Island, Dolinsky District, 106 km of the Federal Highway, larch forest with alder, 13 VII 2008, Bogacheva, Tsarenko, VBGI (det. Galanina sub P. glauca); Korsakovsky District, Prigorodnoe settlement, larch forest, bark, 22 VI 2008, Tsarenko, Chervonnaya, VBGI; ibid,, Protected Area "Laguna Busse", vicinity of Lake Vyselkovoye, 46°33'57.1"N, 143°16'54.7"E, 26 m a. s. l., fir-spruce forest, on rotten wood of stump, 24 V 2017, Konoreva 229, VBGI 79339; ibid,, between Lake Vyselkovoye and Laguna Busse, 46°33'31.1"N, 143°16'55.5"E, 12 m a. s. l., swamp fir-larch-spruce forest, 26 V 2017, Chesnokov 125, LE, MSK; vicinity of Yuzhno-Sakhalinsk, 47°11'N, 142°35'E, larch forest with bamboo, bark of Abies sp., 15 IX 2005, Galanin, VBGI; Makarovsky District, Poreche settlement, coniferous forest, bark, 19 VII 2008, Tsarenko, Bogachova, VBGI; Tomarinsky District, Krasnogorsky Protected Area, road between Lake Ajnskoe and Okhotsk Sea, 48°27'29.1"N, 142°03'06.2"E, 4 m a. s. l., grassy seashore, on bark of Betula sp., 13 V 2017, Chesnokov 25, LE; ibid,, 48°27'34.2"N, 142°03'12.5"E, 7 m a. s. l., fir forest, on bark of Abies sachalinensis (F. Schmidt) Mast., 13 V 2017, Konoreva 78, VBGI 79614; ibid,, vicinity of Lake Uglovoe, 48°33'39.9"N, 141°58'15.0"E, 37 m a. s. l., fir forest with mosses, on

rotten wood, 14 V 2017, Chesnokov 33, Konoreva 85, LE; ibid., 48°34'30.6"N, 141°58'57.3"E, 21 m a. s. l., fir forest, on bark of Abies sachalinensis, 14 V 2017, Chesnokov 40, LE; ibid., 48°34'55.1"N, 141°58'38.0"E, 48 m a. s. l., fir forest with birch and yew, on bark of Taxus cuspidata Siebold et Zucc., 14 V 2017, Chesnokov 43, 45, LE; ibid, 48°34'51.5"N, 141°57'48.7"E, 16 m a. s. l., fir forest with mosses forest along road, on bark of Abies sachalinensis, 14 V 2017, Chesnokov 47, Konoreva 120, LE; ibid., 48°34'45.5"N, 141°56'34.7"E, 12 m a. s. l., larch forest with lichens and mosses, 16 V 2017, Chesnokov 67, LE; ibid, Lake Baklanie, 48°32'08.0"N, 141°59'04.2"E, 10 m a. s. l., larch forest, on bark of Larix sp., Chesnokov 51, LE; Shikotan Island, neighborhood of Tserkovnaya Bay, 43°44'17"N, 146°41'02"E, 38 m a. s. l., yew grove with ferns, on deadwood Taxus cuspidata, 15 VI 2017, Konoreva 379, LE; Iturup Island, Ostrovnoy Protected Area, vicinity of Lesozavodskoy, 44°46'02.1"N, 147°11'41.5"E, 21 m a. s. l., fir forest with bamboo, bark of Sorbus sp., 14 VIII 2017, Chesnokov 275, LE L-15395; ibid, 44°46'07.7"N, 147°11'47.9"E, 18 m a. s. l., fir forest with bamboo and maple, 14 VIII 2017, Chesnokov 276, LE, Konoreva 597, VBGI 79363; ibid., neighborhood of Iodny Cape, 44°43'40.3"N, 147°19'41.2"E, 67 m a. s. l., willow-birch forest with bamboo along the old road, on bark of Salix sp., 16 VIII 2017, Chesnokov 298, LE; Kunashir Island, Kurilsky Strict Nature Reserve, neighborhood of the Saratovsky cordon, left bank of the Saratovskaya River, fir-spruce forest, on bark of Abies sachalinensis, 4 VI 2018, Chesnokov 24, LE.

Platismatia lacunosa (Ach.) W. L. Culb. et C. F. Culb., 1968, Contr. U. S. Natl. Herb. 34(7): 541. (Fig. 1)

= Cetraria lacunosa Ach., 1803, Method Lich.: 295. = Platysma lacunosum (Ach.) Nyl., 1855, Bot. Not.: 137. — Neotype: North America, Herb. Swartz, S. = Cetraria lacunosa f. cavernosa (Menzies) G. Merr., 1910, Bryologist 13: 20, tab. II, fig. 4.

Platismatia lacunosa is characterized by gray or whitish-gray upper surface, strongly reticulate and ridged up to the margins. Pseudocyphellae sometimes present on the ridges, but poorly visible on a whitish thallus. Soredia and isidia absent. Lower surface black at the center and brown or white at the margins with scarce or abundant rhizines. Apothecia relatively common, submarginal. Spores ellipsoid 7.0-10.0 x 3.0-4.5 ^m. Pycnidia marginal to submarginal or even superficial (on the crests of the reticulations).

Chemistry. Upper cortex contains atranorin, K+ yellow, KC, C, and P-. Medulla contains caperatic and fumarprotocetraric acids, K, KC, C, and IKI-, P+ orange-red, UV-.

E co logy. Saxicolous in lichen and dwarf shrub tundra. In western North America the species inhabits different substrates (bark, wood, stones, mosses) in coastal rainforests.

Distribution. Pacific coast of North America (Brodo et al, 2001) and the Aleutian Islands, including the westernmost Attu Island (Talbot et al, 1991). In Russia, it is known only from Medny Island (Commander Islands, Kamchatka Territory).

Notes. Platismatia lacunosa is distinguished from all other Platismatia species by its positive (red) medullary reaction with P (fumarprotocetraric acid). It differs from the morphologically closest P. norvegica by the absence of isidia, and by chemistry

Fig. 1. Platismatia lacunosa (thallus with apothecium), Medny Island, Commander Islands.

Scale bar: 1 cm. Photo by I. Stepanchikova.

(presence of fumarprotocetraric acid). Before 1926 (Savicz, 1926) the species was erroneously recorded for Eurasia, being confused with Asahinea chrysantha (Tuck.) W. L. Culb. et C. F. Culb. (= Cetraria chrysantha Tuck.) (e. g., Mattirolo, Belli, 1903; Elenkin, 1907). All further reports of Platismatia lacunosa from the Russian Arctic, namely Novaya Zemlya, Franz Josef Land, and Taimyr Peninsula (Kristinsson et al., 2010; Spisok..., 2010) are based on incorrect identifications or citing of previous erroneous publications and refer to Asahinea chrysantha (LE). Hence, Platismatia lacuno-sa is reported here as new for Russia.

Selected specimens examined. Kamchatka Territory: Aleut District, Komandorsky Strict Nature Reserve, Commander Islands, Medny Island, top of the SE spur of the Mt. Sen'kina, 54°43'40.4"N, 167°38'53.2"E, 210 m a. s. l., rocky outcrops in lichen-dwarf shrub tundra, on rock, 22 VIII 2019, Himelbrant, Stepanchikova, LE L-15387; ibid,, slope of the hill W of Lake Gladkovskoe, 54°44'08.7"N, 167°42'17.9"E, 146 m a. s. l., rocky outcrops in lichen-dwarf shrub tundra, on rock, 20 VIII 2019, Himelbrant, Stepanchikova, LECB.

Platismatia norvegica (Lynge) W. L. Culb. et C. F. Culb., 1968, Contr. U. S. Natl. Herb. 34(7): 543.

= Cetraria lacunosa f. norvegica Lynge, 1921, Vidensk. Selsk. Skr., Math. Naturvid. Kl. [Kristiania] 1921(7): 196. = Cetraria norvegica (Lynge) Du Rietz, 1924, Bot. Not.: 59. — Lectotype: Norway, Hordaland, Store Kalso, 1920, Lynge, O.

= Cetraria norvegica f. sorediata Degel., 1935, Acta Phytogeogr. Suec. 7: 52. = Cetra-ria tuckermanii var. sorediata (Degel.) Rasanen, 1952, Kuopion Luonnon Ystavain Yhdistyksen Julkaisuia, ser. B, II, no. 6: 31, 46.

The diagnostic characters of P. norvegica are gray, strongly ridged reticulate upper surface, small pseudocyphellae of irregular shape confined to the crests of the ridges, and simple or coralloid isidia, usually associated with the pseudocyphellae and abundant along the margins of the ridges. Lower surface is also reticulate, brown or black at the center and brown or white at the margins with scarce black rhizines. Apothecia uncommon, submarginal. Spores ellipsoid 6.0-8.5 x 3.0-4.0 |im. Pycnidia rare, immersed, marginal.

Chemistry. Upper cortex contains atranorin, K+ yellow, KC, C, and P-. Medulla contains caperatic acid, K, KC, C, and P-, IKI- or + purple, UV-.

Ecology. Oceanic species, inhabiting rocks and bark of spruces in coniferous and mixed forests in humid conditions.

Distribution. The species is distributed in Europe and North America (Culberson, Culberson, 1968; Obermayer, Randlane, 2012). In Russia, it is known with certainty only from Olovgora Mountain in Arkhangelsk Region (Tarasova, 2014). The species was also reported from most regions of southern Far East (Khabarovsk Territory, Jewish Autonomous Region, Sakhalin Region, and Primorye Territory) (Tolpy-sheva, 1990; Tchabanenko, 2002; Tchabanenko et al., 2002; Skirina, 2007, 2016), but all these records are doubtful. All available samples (VGEO) collected in the Far East and previously identified as P. norvegica belong to P. interrupta.

Notes. Platismatia norvegica is similar to P. glauca, but differs by the distinct reticulation and more distinct pseudocyphellae usually associated with the ridges. P. in-terrupta also can be slightly reticulate, but far not as explicitly as P. norvegica, and also the pseudocyphellae of P. interrupta are much more abundant and conspicuous.

Key to Platismatia species published for Russia

(the species erroneously reported from Russia are in square brackets) 1. Lower surface of the thallus covered with distinct punctiform pseudocyphellae. Lobes usually

wide, rounded, reticulate, and isidiate...................................................................................[P. erosa]

— Pseudocyphellae present only on the upper surface or absent..................................................................2

2. Thallus consists of two types of lobes simultaneously: first, broad and rounded, and second,

lacerate, elongated, thin and richly branched, often heavily isidiate.............................P. glauca

— All lobes of thallus are morphologically more or less equal, rounded or elongated.............................3

3. Lobes broad and rounded...................................................................................................................................4

— Lobes narrow (5 mm wide or less), elongate to ribbon-shaped, branched,

margins isidiate..........................................................................................................................[P. herrei]

4. Soredia and/or isidia present.............................................................................................................................5

— Soredia and isidia totally absent.......................................................................................................................9

5. Soredia present (often together with isidia)..................................................................................................6

— Soredia absent, isidia present...........................................................................

6. Pseudocyphellae on the upper surface are distinct, round to irregular,

7

elongate or confluent.....................................................

— Pseudocyphellae absent or indistinct..................................

7. Thallus highly reticulate (check lower surface as well!)

— Thallus surface smooth or weakly reticulate.....................

8. Pseudocyphellae abundant on the upper surface..............

— Pseudocyphellae absent or indistinct..................................

P. interrupta

.......P. glauca

. P. norvegica

P. interrupta .......P. glauca

8

9. Upper surface strikingly lacunose, prominently ridged by strong reticulations rising at right

angles to the surface, spaces between ridges very deep, medulla P+ orange-red.....P. lacunosa

Discussion

In the course of our study, we encountered that many Platismatia records in eastern Siberia and the south of the Russian Far East are not confirmed by herbarium specimens. Thus, we draw conclusions about the distribution of the species based on the revision of available herbarium materials, consultations with colleagues, and analysis of the global ranges of species.

All available specimens previously identified as Platismatia erosa and P. herrei (LE, VGEO) we re-identified as P. interrupta. Platismatia herrei inhabits the western coast of North America (Brodo et al, 2001). The range of P. erosa is limited by the southern part of Asia: China, India, Indonesia, Japan, Nepal, Taiwan, Philippines, and Vietnam (Culberson, Culberson, 1968; Poelt, 1990; Wei, 1991; Awasthi, 2007; Obermayer, Randlane, 2012).

Finally, four species of Platismatia are known to occur in Russia, namely P. glauca, P. interrupta, P. lacunosa, and P. norvegica. Platismatia glauca is distributed throughout the European part of Russia, Ural, and Siberia. Platismatia interrupta is a widespread and common species in the southern Russian Far East and quite rare in Eastern Siberia. Unfortunately, the revision of Russian specimens in the lichen Herbarium of the University of Tartu (TU, Estonia) is currently impossible due to the Covid-19 pandemic. However, thanks to the kind assistance of Professor Tiina Randlane, we made sure that both species (P. glauca and P. interrupta) are present in the Khamar-Daban Range (T. Randlane, pers. comm.). There are numerous (some pretty rich) samples of P. glauca and three samples of P. interrupta from this area in TU. All three P. interrupta specimens are from the right shore of the Solzan River. They are not very healthy, but definitely have pseudocyphellae. Thus, the contact zone of the two most common Platismatia species in Russia is Eastern Siberia.

Platismatia norvegica shows a disjunct distribution pattern, being present in Russia only in the Archangelsk Region; the records from the Russian Far East are not supported by specimens. P. lacunosa is reported here for the first time from a single island at the easternmost limit of the Russian Far East.

Upper surface without strong reticulations or interspaces remain shallow, medulla P

7

In some cases, the revised herbarium specimens were previously misidentified as members of the genus Platismatia, for example, Asahinea chrysantha, Cetrelia braun-siana, C. cetrarioides, C. olivetorum, Nephromopsis laureri, etc. However, these species have a different set of secondary metabolites from Platismatia and can be identified using spot-tests and chromatography methods even in old herbarium material.

Acknowledgments

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We are grateful to L. Konoreva, L. Poryadina, T. Randlane, I. Galanina, and L. Yakovchenko for valuable consultations about distributions of the species. Special thanks are due to L. Konoreva, who kindly provided samples from the South of the Russian Far East. V. Bakalin, K. Klimova, and E. Borovichev are acknowledged for assistance in the loan of herbarium specimens. We thank E. Mamaev and the team of the Komandorsky Strict Nature Reserve for critical help during field studies. The authors are grateful to the reviewers for their valuable corrections and comments. The study of E. Kuznetsova, I. Stepanchikova, and D. Himelbrant was carried out within the framework of the institutional research project "Flora and systematics of algae, lichens, and bryophytes of Russia and phytogeographically important regions of the world" (no. 121021600184-6) of the Komarov Botanical Institute RAS and supported by Russian Foundation for Basic Research (grant 18-05-60093).

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