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19LICHENS - ЛИШАЙНИКИ
The genus Rinodina (Physciaceae, lichenized Ascomycota) in the Magadan Region (Far East of Russia)
I. A. Galanina1, L. S. Yakovchenko1, E. V. Zheludeva2, Y. Ohmura3
'Federal Scientific Center of East Asian Terrestrial Biodiversity, Far Eastern Branch of the Russian
Academy of Sciences, Vladivostok, Russia 2Institute of Biological Problems of the North, Far Eastern Branch of the Russian Academy of Sciences, Magadan, Russia 3Department of Botany, National Museum of Nature and Science, Tsukuba, Japan Corresponding author. I. A. Galanina, [email protected]
Abstract. The lichen genus Rinodina in the Magadan Region is revised on the basis of extensive collections by the authors in 2011-2015. Fifteen species have been recorded, of which Rinodina cinereovirens, R. endospora, R. laevigata, R. metaboliza, R. olivaceobrunnea, R. parasitica, and R. sub-parieta are new for the study area. Rinodina endospora and R. sicula are rare in Russia and have only recently been found in Northeastern Asia. The presence of R. archaea and R. exigua in the Far East of Russia has not yet been confirmed.
Keywords: biodiversity, biogeography, lichens, new records, North-East Asia.
Род Rinodina (Physciaceae, lichenized Ascomycota) в Магаданской области
(Дальний Восток России)
И. А. Галанина1, Л. С. Яковченко1, Е. В. Желудева2, й. Омура3
1Федеральный научный центр биоразнообразия наземной биоты Восточной Азии ДВО РАН,
Владивосток, Россия 2Институт биологических проблем Севера ДВО РАН, Магадан, Россия 3Department of Botany, National Museum of Nature and Science, Tsukuba, Japan Автор для переписки. И. А. Галанина, [email protected]
Резюме. Изучена коллекция лишайников рода Rinodina из Магаданской обл., собранная авторами в 2011-2015 гг. В результате проведенной нами ревизии список лишайников рода Rinodina для Магаданской обл. включает 15 видов, 7 из которых приводятся впервые (Rinodina cinereovirens, R. endospora, R. laevigata, R. metaboliza, R. olivaceobrunnea, R. parasitica и R. subpa-rieta). Rinodina endospora и R. sicula являются редкими в России и только недавно были найдены в Северо-Восточной Азии. Присутствие видов R. archaea и R. exigua на Дальнем Востоке России к настоящему времени не подтверждено.
Ключевые слова: биоразнообразие, биогеография, лишайники, новые находки, Северо-Восточная Азия.
https://doii.org/1031111/Knr/2021551.97
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The genus Rinodina (Ach.) Gray belongs to the family Physciaceae and comprises approximately 300 species worldwide (Sheard, 2010; Wijayawardene et al., 2020). According to Nadyeina et al. (2011), Rinodina is polyphyletic and includes species with crustose thallus, lecanorine apothecia, one-septate brown ascospores with inner wall thickenings and Lecanora-type asci.
Rinodina is a difficult genus in terms of species identification due to the large variety of spore types. Recent studies of the genus Rinodina in Russia (including the Magadan Region), China, South Korea, and Japan have made a great progress in our understanding of the diversity of this genus in the Northeastern Asia (Sheard, 2010; Sheard et al., 2017). Sheard et al. (2017) reported many species which were previously unknown in the Russian Far East. In addition, the authors also revealed the close relationship between Asian and American species, including some that were previously considered to be endemic to North America.
The genus Rinodina in the Russian Far East has been actively studied in the last decade. However, the majority of studies are focused on the south of the area (Galanina et al., 2011, 2018, 2021; Galanina, 2013, 2016, 2019; Sheard et al., 2017; Galanina, Ezkhin, 2018, 2019; Konoreva et al, 2018; Yakovchenko et al, 2018) while data on its northern part (including the Magadan Region) remain rather poor (Korolev, Tolpy-sheva, 1980; Kotlov, 1991, 1993a, 1993b, 1995, 2004), although some new species were reported from the region as it is indicated below.
The first records of the genus Rinodina in the Magadan Region were made by Ko-rolev and Tolpysheva (1980) who reported two species, R. archaea (Ach.) Arnold and R. exigua (Ach.) Gray. Later, the species Rinodina hyperborea H. Magn., R. milvina (Wahlenb.) Th. Fr., R. mniaroea (Ach.) Körb., R. sibirica H. Magn., R. turfacea (Wah-lenb.) Körb. were reported for the Verkhnekolymskoe Highland (Kotlov, 1991, 1993a, 1993b, 1995, 2004). Finally, Sheard et al. (2017) reported R. freyi H. Magn., R. roscida (Sommerf.) Arnold, and R. septentrionalis Malme. In total, ten species of the genus Rinodina were known for the Magadan Region before the present study
The Magadan Region, with an area of 461400 km2, lies in the northeastern part of the Russian Far East. The mountainous area framing the northern shores of the Okhotsk Sea and the basin of the upper reaches of the Kolyma River is formed by the ridges of the Kolyma Highlands and the spurs of the Chersky Ridge mountain system. A narrow strip along the northern coast of the Okhotsk Sea is characterized by a temperate continental climate with relatively low winter minimum temperatures and variable weather conditions. The Okhotsk-Kolyma watershed is a climate border with very frosty winters, an average January temperature there is below -32 °C (Klyukin, 1970). The mountainous regions in the upper Kolyma River basin are characterized by the greater climate continentality. According to the geobotanical zoning by Reut (1970), different parts of the study area belong to the mountainous area of dwarf pine-larch-birch forests of the Okhotsk coast, to the bog-tussock-tundra region of the Yamsk-Tauy depression, to the area of mountain-arctic tundra, and to the lichen woodlands of the Kolyma Highlands. Yurtsev (1974) assigns this ter-
ritory to the North Okhotsk province of the East Siberian subregion of the Boreal floristic region.
The aim of our study is to contribute to the knowledge about the diversity of Rinodina species in the Magadan Region, as well as about their ecology and distribution.
Material and Methods
The material was collected by the authors in 2011-2015 in several districts of the Magadan Region (Olsky, Omsukchansky, Yagodninsky, Severo-Evensky, Tenkinsky, and Khasynsky) (Fig. 1). Additionally, we studied vouchers from the herbaria of the Institute of Biological Problems of the North, Far Eastern Branch of the Russian Academy of Sciences (MAG) and Komarov Botanical Institute of the Russian Academy of Sciences (LE) in Russia, and the National Museum of Nature and Science (TNS) in Japan. The lichens sampled in 2012-2013 are kept at the Federal Research Center for Biodiversity of the Far Eastern Branch of the Russian Academy of Sciences (VLA). A total of 40 samples from the Magadan Region were studied.
The morphological and anatomical features of the specimens were examined by standard methods of light microscopy. Chromatography was not necessary because all the studied species can be easily distinguished by anatomy and morphology as well as by standard spot tests with KOH (K), Ca(ClO)2 (C), and C6H4(NH2)2 (P). The presence of sphaerophorin crystals was studied by the method of fluorescence in longwave UV.
Results and Discussion
In total, 15 species of Rinodina were identified, of which R. cinereovirens Vain., R. endospora Sheard, R. laevigata (Ach.) Malme, R. metaboliza Vain., R. olivaceobrun-nea C. W. Dodge et G. E. Baker, R. parasitica H. Mayrhofer et Poelt, and R. subpa-rieta (Nyl.) Zahlbr. are new for the Magadan Region. Two species, R. endospora and R. sicula H. Mayrh. et Poelt, are rare in Russia and have only recently been found in North-eastern Asia (Sheard et al, 2017; Galanina et al., 2021). Rinodina archaea, R. exigua, R. milvina, and R. mniaroea were previously reported from the Magadan Region (Korolev, Tolpysheva, 1980; Kotlov, 1995), but were not found during our field studies, as well as in the herbarium collections. The presence of R. archaea and R. exigua in the Far East of Russia has also not yet been confirmed (Sheard et al., 2017, Galanina, Ezhkin, 2019; Galanina et al., 2021).
Newly recorded species for the Magadan Region
Rinodina cinereovirens Vain. (Plate I: 1, 2)
Rinodina cinereovirens is characterized by thin light gray or brownish gray thal-lus without vegetative propagules (Plate I: 1). Apothecia quickly become narrowly attached, sometimes almost stipitate (to 0.6-1.0 mm) with disc black and plane. Thalline margin is entire and typically persistent (80-120 |im wide), with cortex expanded to 20-60 |im wide. Rinodina cinereovirens has Physcia-type spores of Type A
Fig. 1. The studied sites of the Rinodina species in the Magadan Region.
development, (21.5)23.0-25.5(27.5) x (10.0)11.5-13.5(14.0) ^m (Plate I: 2). Torus well developed. Crystals absent in cortex and present in medulla (sphaerophorin). Spot tests all negative, the secondary metabolite sphaerophorin in longwave UV turns blue-white.
Rinodina cinereovirens is closely related to R. turfacea but differs by more broadly ellipsoid spores with more bluntly rounded apices. Furthermore, R. cinereovirens inhabits bark and wood, in contrast to R. turfacea which typically grows on decaying ground vegetation, less often on wood in oroarctic environments in North America (Sheard et al., 2017).
Plate I. 1 — thallus and apothecia of Rinodina cinereovirens; 2 — Physcia-type spores of R. cinereovirens; 3 — thallus and apothecia of R. endospora; 4 — Dirinaria-type spore of R. endospora; 5 — thallus and apothecia of R. freyi; 6 — Physcia-type spores of R. freyi. Scale bars: 1, 3, 5 — 0.5 mm; 2, 4, 6 — 10 |m.
Distribution. In Russia, the species has previously been reported as Rinodina tur-facea var. cinereovirens (Vain.) H. Mayrhofer and as R. turfacea var. ecrustacea (Vain.) H. Oliv. for different regions from the Leningrad Region to the Kamchatka Territory (Urbanavichene, Urbanavichus, 1998; Kotlov, 2008; Himelbrant et al, 2009; Spi-sok..., 2010; Chesnokov, Konoreva, 2015; Sheard et al, 2017). Rinodina cinereovirens is known in Europe from Scandinavia (Norway, Sweden, and Finland) (Mayrhofer, Moberg, 2002). In North America the species was reported from Newfoundland, New Brunswick, Wapusk National Park, northern Manitoba, northern Ontario, and Alaska (Sheard, 2018).
Ecology. Rinodina cinereovirens was found on bark of Alnus sp., Betula sp., Cho-senia arbutifolia, and Larix sp. in floodplain shrub communities and coniferous forests.
Specimens examined: Olsky District, Tauy River, the Niro station, floodplain terrace, 59°47'17.2"N, 148°16'13.2"E, 36 m a. s. l., old larch forest with birch and alder, with tall grass and shrubs, on bark of Alnus sp., 6 VIII 2011, Zheludeva, MAG 0-4581; ibid., Magadansky Reserve, Ka-va-Chelomdzhinsky area, Chelomdzha River valley, 59°47'50.3"N, 148°12'52.9"E, 36 m a. s. l., flood-plain forest, on bark of Chosenia arbutifolia, 12 VII 2012, Zheludeva, MAG 0-4582; ibid., Kamenny Range, vicinity of the tourist camp Magtur, Pyany stream, 59°45'24.5"N, 149°39'17.9"E, 9 m a. s. l., floodplain shrubs, on bark of Alnus sp., 6 VIII 2013, Galanina M-13-178-3, VLA; Omsukchansky District, 500 km NE from Magadan, foothill of Kilganskie Range, vicinity of Dzhuletta mining camp, 61°11'40.9"N, 153°56'55.7"E, 969 m a. s. l., larch-dwarf pine forest with rocks along stream, on dead twigs of Larix sp., 9 VIII 2012, Yakovchenko 25-12, VLA.
Rinodina endospora Sheard (Plate I: 3, 4)
The detailed description was given by Sheard (2010), Sheard et al. (2017), and Galanina et al. (2021). The species is characterized by a gray to gray-brown verrucose or areolate thallus, narrowly attached apothecia with flakes of the epinecral layer on the thallus, large Dirinaria-type spores (20.5)22.0-24.0(27.5) x (8.5)9.0-10.0(11.0) jim, often asynchronous and of Type B development (Plate I: 3, 4). Rinodina endospora can be confused with R. metaboliza which also has Dirinaria-type spores, but they are of smaller size (13.5)17.5-19.0(23.0) x (8.5)9.0-10.0(11.0) jm (Sheard, 2010).
Distribution. This is the third location of this species in Russia and Northeastern Asia. Previously, the species was reported from Kamchatka (Ust'-Bol'sheretsky District, on bark of Chosenia arbutifolia) (Sheard et al., 2017) and from Sakhalin Island where it was found on bark of Populus sp. in riparian forest (Galanina et al., 2021). Rinodina endospora was previously considered to be a western North American endemic species distributed in California along the coastal ranges and in the Sierra Nevada (Sheard, 2010). This study suggests it is the western North American — East Asian species with disjunct distribution.
Ecology. The species was found on bark of Chosenia arbutifolia in the valley forest.
Specimen examined: Olsky District, Magadansky Reserve, Kava-Chelomdzhinsky area, Chelomdzha River valley, 59°47'50.3"N, 148°12'52.9"E, 36 m a. s. l., floodplain forest, on bark of Chosenia arbutifolia, 12 VII 2012, Zheludeva, MAG 0-4583.
Rinodina laevigata (Ach.) Malme (Plate II: 1, 2)
Rinodina laevigata is characterized by Physcia-Physconia-type spores (14.5)18.5-19.5(22.5) x (7.0)8.5-9.0(10.5) jm of Type A development, thin poorly developed thallus, and usually thick lower apothecial cortex (Plate II: 1, 2). Rinodina laevigata can be confused with R. sibirica H. Magn. but differs by a less developed thallus, grayish color, smaller spores, and more rounded and less elongated lumina. Apothecia in R. laevigata are broadly attached to the thallus, with typical plane discs and thick lower apothecial cortex.
Distribution. In Russia, the species is known from the North Caucasus to Western Siberia (Kotlov, 2008), but most probably is understudied. Rinodina laevigata is widespread and known from Norway, Sweden, Finland, and Scotland in Europe (Mayrhofer, Moberg, 2002), also from Alaska to California and in the Sierra Nevada in the western part of North America (Sheard, 2010).
Ecology. Rinodina laevigata was collected on bark of Pinuspumila (Pall.) Regel and Larix sp.
Specimens examined: Khasynsky District, ca. 25 km SW of Atka, 60°37'22.7"N, 151°33'31.6"E, 1110 m a. s. l., on twig of Larix sp., 11 VIII 2012, Ohmura 10172B, Yakovchenko, Zheludeva, TNS; Ten-kinsky District, Matrosov mine, thickets of dwarf pine, 61°38'28"N, 147°48'10"E, on bark of Pinus pumila, 8 VI 2015, Pavlov, VLA.
Rinodina metaboliza Vain. (Plate II: 3, 4)
Rinodina metaboliza is characterized by a Dirinaria-type spores (13.5)17.5-19.0(23.0) x (8.5)9.0-10.0(11.0) jm of Type B development (Plate II: 4). The species is very variable in thallus morphology, spore size and apothecial convexity (Plate II: 3). Rinodina metaboliza can be confused with R. endospora. For the differences, see the description of R. endospora above.
Distribution. The species was described by Vainio (1928) from several places along the Yenisei River in Siberia. In Russia, it is known from the republics of Karelia and Komi, and South Siberia (Eastern Sayan Mountains) (Kotlov, 2008; Urbanavi-chene, Urbanavichus, 2009). In Europe, the species was found in Scandinavia (central Sweden and Norway) (Mayrhofer, Moberg, 2002). In North America, it is a western species extending from coastal Alaska to southern California and inland to the Rocky Mountains (Sheard, 2010).
Ecology. Rinodina metaboliza was collected on bark of Populus sp. in floodplain forest.
Specimen examined: Olsky District, Magadansky Reserve, Chelomdzha River, 59°47'50.3"N, 148°12'52.9"E, 36 m a. s. l., floodplain forest with Chosenia arbutifolia and Alnus sp., on bark of Populus sp., 12 VII 2012, Zheludeva, MAG 0-4584.
Rinodina olivaceobrunnea C. W. Dodge et G. E. Baker (Plate II: 5, 6)
Rinodina olivaceobrunnea is characterized by Physcia-type spores (16.5)20.5-21.5(26.0) x (8.0)9.5-10.0(12.0) jm of Type A development (Plate II: 6). The species
has abundant and small, narrowly attached apothecia (Plate II: 5). Rinodina olivaceo-brunnea can be confused with R. turfacea, but differs in smaller spores and apothecia, absence of a massive columnar lower cortex, and lack of sphaerophorin. R. olivaceo-brunnea can also be confused with R. archaea, but it is distinguished by its Physcia-type spores, thick lower cortex, and typically muscicolous rather than lignicolous habitat (Sheard, 2010).
Distribution. In Russia, R. olivaceobrunnea has wide distribution and was previously found in the Arctic (Novaya Zemlya, Taimyr, and Chukotka), Murmansk Region, Komi Republic, North Caucasus (Karachayevo-Circassian Republic), and West Siberia (Kotlov, 2008). Rinodina olivaceobrunnea is distributed in both hemispheres being known in Europe, central Africa, Australasia, Antarctica, and North America (Mayrhofer, Moberg, 2002; Sheard, 2010).
Ecology. Rinodina olivaceobrunnea was collected on old wood near the airport.
Specimen examined: Severo-Evensky District, Evensk village, 61°55'02.8"N, 159°14'20.8"E, 3 m a. s. l., old vegetable gardens behind the airport, on old logs (wood), 11 VII 2015, Zheludeva, MAG CЭ-4585.
Rinodina parasitica H. Mayrhofer et Poelt (Plate III: 1, 2)
Rinodina parasitica is characterized by a lichenicolous habit and brown thallus, Physcia-Physconia-type spores (13.5)16.0-16.5(19.0) x (7.0)8.5-9.0(10.5) jm of Type A development (Plate III: 1, 2). It can be confused with R. obnascens (Nyl.) Oliv., another lichenicolous species, but the latter has Milvina-type spores and gray thallus with minute dark brown to black consoredia (Sheard, 2010). R. obnascens has not been found in Russia yet.
Distribution. In Russia, R. parasitica was found in Siberia (Krasnoyarsk Territory, Baikal Region) (Kotlov, 2008). The species occurs in scattered localities in Europe (Norway, Sweden, and Finland) (Mayrhofer, Moberg, 2002), Asia (Mongolia) (Byazrov, 2013), and North America, where it is known in Alaska and western Canada, southern Rocky Mountains, coastal ranges, and Sierra Nevada (Sheard, 2010).
Ecology. Rinodina parasitica was found on the thallus of an unidentified Aspi-cilia. It is arctic-alpine lichen growing on species of Aspicilia and Rhizocarpon (Kotlov, 2008).
Specimen examined: Olsky District, Magadansky Reserve, Yamsky area, Studenaya station, a small mountain, 59°45'38.0"N, 153°34'24.9"E, 69 m a. s. l., on Aspicilia sp. growing on rock, 15 VII 2015, Zheludeva, MAG 0-4586.
Plate II. 1 — thallus and apothecia of Rinodina laevigata; 2 — Physcia-Physconia-type spores of R. laevigata; 3 — thallus and apothecia of R. metaboliza; 4 — Dirinaria-type spore of R. metaboliza; 5 — thallus and apothecia of R. olivaceobrunnea; 6 — Physcia-type spores of R. olivaceobrunnea. Scale bars: 1, 3, 5 — 0.5 mm; 2, 4, 6 — 10 |m.
Rinodina subparieta (Nyl.) Zahlbr. (Fig. 2)
Rinodina subparieta is characterized by a Physconia-type spores (15.0)18.5-19.5(23.0) x (8.5)10.0-11.0(13.0) ^m of Type A development, light gray scattered areoles, producing atranorin (K+ yellow) and whitish soredia forming along the areole margins in a labriform soralia (Fig. 2). Rinodina subparieta are similar to R. willeyii Sheard et Giralt, but the latter has Pachysporaria-type spores, whitish areolate thallus (than subsquamulose), and producing pannarin (P+ cinnabar).
Distribution. In Russia, it is a widespread species known from the Caucasus to the Far East (Spisok..., 2010; Himelbrant, Stepanchikova, 2011; Galanina, 2013; Sheard et al, 2017). In Europe it is known from Scotland (Giavarini et al, 2009), Scandinavia and Austria (Tonsberg, 1992; Mayrhofer, Moberg, 2002). Rinodina subparieta is also known from Mongolia (Hauck, Javkhlan, 2006) and eastern Asia with a distribution spanning Russia, Japan, and Korea (Sheard et al., 2017). In North America, R. subparieta has a Pacific and North Atlantic disjunct distribution (Sheard, 2010; Resl et al, 2016).
Ecology. Rinodina subparieta was collected on bark of Alnus sp. in old larch forest. In Russia, the species was found on coniferous and deciduous trees (Abies sp., Alnus sp., Betula sp., Castanea sp., Chosenia arbutifolia, Picea sp., Prunus sp., Quercus sp., Salix sp., and Taxus sp.) in deciduous and mixed forests, often along river valleys. It occurs from the sea level to 2550 m (Sheard et al., 2017).
Specimen examined: Olsky District, Tauy River, the Niro station, floodplain terrace, 59°47'17.2"N, 148°16'13.2"E, 36 m a. s. l., old larch forest with Betula sp., Alnus sp., tall grasses and shrubs, on bark of Alnus sp., 6 VIII 2011, Zheludeva, MAG 0-4587.
Fig. 2. Thallus of Rinodina subparieta consisting of light gray scattered areoles with whitish soredia forming along the areole margins. Scale bar: 0.5 mm.
Species previously known from the Magadan Region Rinodina freyi H. Magn. (Plate I: 5, 6)
Rinodina freyi is characterized by gray-green continuous thallus, apothecia frequently becoming contiguous on small thalli (Plate I: 5), and relatively small, darkly pigmented Physcia-type spores (12.0)15.0-16.0(18.5) x (6.0)7.5-8.0(9.0) pm of Type A development with a heavy torus (Plate I: 6). Rinodina freyi can be confused with R. septentrionalis, but the latter species has copper-brown thallus consisting of small discrete verrucae, narrowly attached and scattered apothecia.
Distribution. In Russia, R. freyi was previously recorded from the Magadan Region, Kamchatka Territory, Sakhalin Island, Khabarovsk and Primorye territories (Sheard et al., 2017; Galanina et al, 2021). It has also been reported from Japan (Sheard et al, 2017) and western Mongolia (Hauck et al, 2013). European distribution of this species is poorly known although originally it was described from Europe (Switzerland) (Magnusson, 1947); the species is also published from Germany (Wirth et al, 2013). Sheard (2010) indicated that R. freyi is closely related to the European R. septentrionalis (Giralt, Mayrhofer, 1995). Rinodina freyi is the most common species of the genus in North America, being frequent in both the east and west of the continent (Sheard, 2010).
Ecology. In the Magadan Region, R. freyi was found on bark of Alnus, Chosenia, and Salix in floodplain Chosenia forest and alder forest. In North America it inhabits various tree species, including Abies spp., Acer spp., Alnus crispa (Aiton) Pursh, Betula spp., Fraxinus sp., Picea spp., Pinus spp., Quercus macrocarpa Michx., Sorbus sp., Ulmus americana L. mainly in the southern boreal zone (Sheard, 2010).
Specimens examined: Olsky District, Ola River, floodplain forests, on bark of Salix sp., 8 VII 2013, Galanina M-13-203-1, VLA; ibid, Galanina M-13-186-1, M-13-210-1, VLA (Sheard et al, 2017); ibid, Spafareva Island, N part of the Iisland, 59°11'41.7"N, 149°05'09.8"E, 110 m a. s. l., alder forest along the snowfield, on Alnus sp., 20 VII 2013, Zheludeva, MAG 0-4588; ibid, the upper reaches of the Ola River, 138 km of the federal highway, near the bridge over the river, 60°24'49.5"N, 151°30'49.0"E, 13 m a. s. l., floodplain forest, on bark of Chosenia arbutifolia, 13 VII 2013, Galanina M-13-193-1, VLA; Severo-Evensky District, Evensk village, 61°55'04.2"N, 159°14'05.8"E, 3 m a. s. l., larch in the planting, on bark of Larixsp., 11 VII 2015, Zheludeva, MAG 0-4589; Yagodninsky District, Myakit River valley, 295 km of the federal highway, 61°24'25.8"N, 152°05'25.9"E, 664 m a. s. l., single old larch trees at the bottom of a slope, on bark of Larix sp., 13 VII 2013, Galanina M-13-195-1, VLA.
Rinodina milvina (Wahlenb.) Th. Fr.
Rinodina milvina is characterized by thick dark brown thallus and Milvina-type spores (15.5)18.0-19.0(22.0) x (7.5)9.5-10.5(12.0) pm of Type A development. Young thalli of R. milvina can be confused with R. parasitica, which is well distinguished by its smaller Physcia- Physconia-type spores.
Distribution. Rinodina milvina was previously noted in the Magadan Region by Kotlov (1995). In Russia, the species is widespread from the European part to the Far
East, and from the Southern Siberia to the Arctic (Novaya Zemlya) (Kotlov, 2008). Rinodina milvina has scattered distribution in Europe (Mayrhofer, Moberg, 2002) and also occurs in Asia (Turkey, Iraq, Georgia, Armenia, Azerbaijan, Kazakhstan, Mongolia, and Japan) (Wagner, Spribille, 2005; Kotlov, 2008; Byazrov, 2010; Ohmura, Kashi-wadani, 2018), North Africa (Kotlov, 2008), and North America (Greenland, Rocky Mountains, Sierra Nevada, Cascades and costal range) (Sheard, 2010).
Ecology. Rinodina milvina inhabits siliceous rocks, sometimes parasitizing crus-tose lichens (Mayrhofer, Moberg, 2002; Kotlov, 2008). In North America, the species was found on granites, quartzites and other acidic rocks, on sandstone, volcanic rocks, from 900 to 3660 m a. s. l. (Sheard, 2010).
Rinodina mniaroea (Ach.) Korb.
Rinodina mniaroea is characterized by its apothecia becoming convex and thalline margin becoming excluded, large Physcia-type spores (20.5)24.5-25.5(30.0) x (9.5)11.5-12.5(14.5) jm of Type A development. Rinodina mniaroea is similar to R. turfacea, which differs by producing sphaerophorin (UV+ blue-white) and persistently plane apothecial discs with prominent thalline margins. Another species to be distinguished from R. turfacea is R. olivaceobrunnea, which differs by its smaller apo-thecia and spores.
Distribution. Rinodina mniaroea was previously reported from the Magadan Region by Kotlov (1995). In Russia, the species is widespread in the northern and mountain regions (Arctic, northern European part, North Caucasus, Siberia, Urals, Altai, Far East) (Makarova, Katenin, 1983, 1992; Sedelnikova, 1990; Kotlov, 2008). In Europe it occurs in central part (Wirth, 1995; Giavarini et al., 2009), Iberian Peninsula (Giralt, 2010), and Scandinavia (Mayrhofer, Moberg, 2002). In Asia R. mniaroea was reported from the Caucasus, Himalayas, Mongolia, and China (Vezda, 1965; Kotlov, 2008). In North America, R. mniaroea is common in the western Arctic, the Rocky Mountains and scattered in the eastern Arctic and Greenland (Sheard, 2010).
Ecology. Rinodina mniaroea grows on mosses and plant debris in the arctic-alpine, less often in the subalpine and boreal zones (Mayrhofer, Moberg, 2002; Kotlov, 2008). In North America it was recorded up to 4070 m a. s. l. in Arizona (Sheard, 2010).
Rinodina roscida (Sommerf.) Arnold
Rinodina roscida is characterized by large Physcia-type spores (22.5)30.0-32.0(39.5) x (10.5)12.5-13.5(16.0) jm and Type A or rarely B development. R. roscida can be confused with R. turfacea and differs in its light gray thallus lacking sphaero-phorin, pruinose apothecial discs, J+ blue lower cortex of apothecial margins, significantly larger spores with often submucronate apices (Sheard, 2010).
Distribution. Rinodina roscida is widespread in the northern and mountainous regions of Northern Hemisphere (Mayrhofer, Moberg, 2002; Wirth et al., 2013): in the Arctic from Scandinavia to Chukotka and southbound to Southern Siberia (Trans-Bai-
kal Territory, Sayany) and Asia (Caucasus, Himalayas, Mongolia, China) (Mayrhofer, Moberg, 2002; Kotlov, 2008; Spisok..., 2010). In North America it is common in the Arctic, especially in the west, extending southwards in the Rocky Mountains (Sheard, 2010).
Ecology. Rinodina roscida was collected on mosses and plant debris on rock in mountains.
Specimens examined: Omsukchansky District, 500 km NE from Magadan, foothill of Kilganskie Range, vicinity of Dzhuletta mining camp, 61°11'39.8"N, 153°58'49.8"E, 1480 m a. s. l., upper part of the slope with calcareous rocks, on mosses, 11 VIII 2012, Yakovchenko M-12-Ca-3, VLA; ibid., Yakovchenko M-12-Ca-1, M-12-Ca-2, VLA (Sheard et al., 2017); Khasynsky District, small mountain ca. 120 km NE of Atka, 61°11'43.4"N, 153°58'38.8"E, 1240 m a. s. l., on plant debris on rock, 11 VIII 2012, Ohmura 10145, Yakovchenko, Zheludeva, TNS.
Rinodina septentrionalis Malme = Rinodina hyperborea H. Magn.
Rinodina septentrionalis is characterized by Physcia-type spores (13.5)16.0-17.0(19.5) x (6.5)7.5-8.5(9.5) pm with well-developed torus and Type A development. Rinodina septentrionalis has copper-brown thallus consisting of small discrete verrucae, narrowly attached and scattered apothecia. For the differences from similar R. freyi see its description.
Distribution. Rinodina septentrionalis was previously reported for the Magadan Region by Kotlov (1995) as R. hyperborea. In Eurasia, R. septentrionalis is widely distributed in the Arctic and boreal zones and in mountainous regions from northern Scandinavia to Kamchatka Peninsula and southbound to Georgia, Kazakhstan, Altai Mts, Sikhote-Alin Mts, Japan (Insarov, Pchelkin, 1984; Davydov, 2001; Giralt, 2001, 2010; Mayrhofer, Moberg, 2002; Tschabanenko, 2002; Kotlov, 2008; Himelbrant et al, 2009; Galanina, 2013; Hauck et al, 2013; Wirth et al, 2013; Yakovchenko et al, 2013; Chesnokov, Konoreva, 2015; Sheard et al., 2017; Galanina et al., 2021). In North America it is common in the Arctic, extending southwards in the Rocky Mountains to Colorado (Sheard, 2010).
Ecology. Rinodina septentrionalis was collected from the bark of Alnus sp., Larix sp., and Salix sp. In the Altai Mts, it was found on the bark of deciduous and coniferous trees in the wet habitats of the forest belt (Davydov, 2001). In North America it was recorded up to 3050 m a. s. l. in the Rocky Mountains (Sheard, 2010).
Specimens examined: Olsky District, Magadansky Reserve, Kava River, 59°47'37.6"N, 148°00'30.2"E, 30 m a. s. l., aspen forest with scarce larch, birch and dwarf pine, on wood, 9 VII 2012, Zheludeva, MAG 0-4590; ibid, Zheludeva 735, MAG (Sheard et al, 2017); Khasynsky District, ca. 25 km SW of Atka, 60°37'22.7"N, 151°33'31.6"E, 1110 m a. s. l., on twig of Larix sp., 11 VIII 2012, Ohmura 10172B, Yakovchenko, Zheludeva, TNS; Magadansky District, 3 km W of Magadan, 59°33'48.4"N, 150°42°'39.0"E, 60 m a. s. l., on twig of Alnus sp., 6 VIII 2012, Ohmura 9942, Yakovchenko, Zheludeva, TNS; ibid., vicinity Snezhnaya Dolina village, floodplain forest, on bark of Salix sp., 4 VII 2001, Lysenko, MAG M-4593; ibid., 59°44'21.1"N, 150°51'23.3"E, 169 m a. s. l., floodplain forest, on bark of Salix sp., 26 VII 2001, Artemova, MAG M-4591.
Rinodina sibirica H. Magn. (Plate III: 3, 4)
Rinodina sibirica is characterized by Physcia-Physconia-type spores (17.0)20.0-21.5(25.5) x (8.5)10.0-11.5(13.0) jm with the well-developed torus, the dark-colored cell walls, and Type A development (Plate III: 4). Rinodina sibirica has erumpent apothecia broadly attached at first and then becoming narrow at the base, scattered, numerous with the dark brown to black disk becoming convex, sometimes hemispherical (Plate III: 3). Rinodina sibirica can be often confused with R. archaea, but the latter species has Physconia-type spores and mainly lignicolous (Sheard, 2010). The young apothecia of R. sibirica with plane discs can be confused with such of R. cinere-ovirens, but the latter has large Physcia-type spores and crystals in medulla (sphaero-phorin, UV+ blue-white). Specimens of R. sibirica can be often referred to R. sophodes, which differs by small Milvina-type spores and European distribution (Sheard, 2010). Rinodina sophodes was probably erroneously reported for the Far East of Russia.
Distribution. Rinodina sibirica was previously reported for the Magadan Region by Kotlov (1995). Rinodina sibirica is an American-Asian species with a wide amphi-beringian range (Galanina et al, 2021). The species was described from the environs of Yeniseisk, Tomsk, and Tobolsk in Russia (Magnusson, 1936). Later, it was additionally reported from numerous places in Siberia along the rivers Yenisei, Ob, Irkut, Lena, and Aldan (Magnusson, 1947), as well as from Altai and Sayan (Kotlov, 2008), and from Mongolia (Golubkova, 1981). Recently R. sibirica was reported from the Trans-Baikal and Khabarovsk territories, the Magadan Region, Sakhalin Island, and the Kamchatka Peninsula (Sheard et al., 2017; Galanina et al., 2021). The species was also reported from North America (Thomson, 1997, as R. granulans Vain.; Sheard, 2010).
Ecology. Rinodina sibirica was collected on bark of Betula sp., Larix sp., and Salix sp. The species grows mainly on bark of deciduous trees (e. g. Alnus spp., Betula spp., Padus avium Mill., Populus spp., Salix spp., Sorbus sp.), less often on conifers (Abies sp., Larix sp., Picea sp., Pinus sp.) and on wood (Magnusson, 1936, 1947; Kotlov, 2008; Sheard, 2010; Sheard et al., 2017).
Specimens examined: Olsky District, floodplain of Kava River, Bogot Island, 59°46'16.6"N, 147°59'26.7"E, 39 m a. s. l., larch-birch forest, on bark of Betula sp., 7 VII 2012, Zheludeva, MAG 0-4592; ibid, Iamborko M-02-1, LE (Sheard et al., 2017); Magadansky District, vicinity of Sne-zhnaya Dolina village, floodplain forest, on bark of Salix sp., 4 VII 2001, Lysenko, MAG M-4593; ibid, on bark of deciduous tree, 28 VI 2007, students, MAG.
Rinodina sicula H. Mayrh. et Poelt (Plate III: 5, 6)
Rinodina sicula is characterized by thin gray to black-brown rimose-areolate thallus (Plate III: 5), Physconia-type spores (16.0)18.0-19.5(23.0) x (7.5)9.0-10.0(11.5) jm
Plate III. 1 — thallus and apothecia of Rinodina parasitica; 2 — Physcia-Physconia-type spores of R. parasitica; 3 — thallus and apothecia of R. sibirica; 4 — Physcia-Physconia-type spore of R. sibirica; 5 — thallus and apothecia of R. sicula; 6 — Physconia-type spores of R. sicula. Scale bars: 1, 3, 5 — 0.5 mm; 2, 4, 6 — 10 |m.
(Plate III: 6) of Type A development, and the presence of gyrophoric acid in the apoth-ecial margin (C+ red).
Distribution. Rinodinasicula is described from Italy (Mayrhofer, Poelt, 1979). It is known in Europe from Italy, Iberian Peninsula, Denmark, Sweden (Mayrhofer, Poelt, 1979; Giralt, Llimona, 1997; Mayrhofer, Moberg, 2002; Mayrhofer, Sheard, 2007) and in Northeastern Asia from Kamchatka Peninsula, Magadan Region, Trans-Baikal Territory (Sheard et al., 2017).
Ecology. Rinodina sicula is a saxicolous species. In the Magadan Region it was found on rocks in the middle part of the slope with cliffs.
Specimen examined: Omsukchansky District, 500 km NE to Magadan, foothill of Kilganskie Range, vicinity of Dzhuletta mining camp, 61°11'43.3"N, 153°58'34.9"E, 1321 m a. s. l., middle part of the slope with cliffs, on rocks, 9 VIII 2012, Yakovchenko M-12-27-4, VLA; ibid., Yakovchenko M-12-27-1, VLA (Sheard et al, 2017).
Rinodina turfacea (Wahlenb.) Körb.
Rinodina turfacea is characterized by brownish-gray thallus, large apothecia with concave or plane disc, persistent thalline margin containing crystals of sphaeropho-rin, Physcia-type spores (22.0)27.5-29.5(35.0) x (10.5)12.5-13.5(15.5) pm of Type A development (Shaerd, 2010). Rinodina turfacea can be confused with R. cinereovirens; for the differences, see the description of R. cinereovirens.
Distribution. Rinodina turfacea was previously reported from the Magadan Region by Kotlov (1995). It is mainly northern circumpolar species restricted to the Arctic and Subarctic territories in Eurasia from Scandinavia to the Kamchatka Peninsula with southernmost locations in the Mongolian and Chinese parts of the Altai (Schubert, Klement, 1971; Afonina et al., 1980; Makarova, Katenin, 1983; Makryi, 1986; Sedelnikova, 1990; Abbas et al, 2001; Davydov, 2001; Mayrhofer, Moberg, 2002; Tschabanenko, 2002; Kotlov, 2008; Himelbrant et al, 2009; Skirina, 2012; Wirth et al, 2013; Sheard et al., 2017; Galanina et al., 2021). In North America, it is common in the Arctic, southward to the Rocky Mountains in Montana and Wyoming, also in the state of Colorado (Sheard, 2010).
Ecology. Rinodina turfacea was found on plant debris.
Specimens examined: Olsky District, Kamenny Range, vicinity of the Magtur tourist base, flood-plain of Pyany stream, 59°45'24.5"N, 149°39'17.9"E, 9 m a. s. l., floodplain shrubs with Alnus sp., on plant debris, 6 VIII 2013, Galanina M-13-178-3, VLA; ibid, Atargan Peninsula, 59°32'35.6"N, 151°29'27.2"E, 3 m a. s. l., seashore, on dry pebble on plant debris, 3 VIII 2016, Zheludeva, MAG 0-4595; Omsukchansky District, 500 km NW from Magadan, foothill of Kilganskie Range, vicinity of Dzhuletta mining camp, 61°11'43.3"N, 153°58'34.9''E, 1321 m a. s. l., middle part of the slope with cliffs, on plant debris, 9 VIII 2012, Zheludeva, MAG; ibid, on plant debris, Yakovchenko 27-12-2,2712-3, VLA (Sheard et al, 2017); ibid, on plant debris, 11 VIII 2012, Ohmura 10145B, TNS.
As a result of the study, the list of Rinodina known from the Magadan Region consists of 15 species, 7 of which are reported for the first time for the region (R. cinereovirens, R. endospora, R. laevigata, R. metaboliza, R. olivaceobrunnea, R parasitica, and
R. subparieta). The genus is mainly represented by species typically occurring in the northern regions (R. cinereovirens, R. metaboliza, R. mniaroea, R. olivaceobrunnea, R. roscida, R. septentrionalis, and R. turfacea).
Despite the new data, the Rinodina genus in the Magadan Region requires further study. Noteworthy are the species R. endospora and R. sicula which are rare in Russia and have only recently been found in Northeastern Asia (Sheard et al, 2017; Galanina et al, 2021). These species might be rarely reported because saxicolous Rinodina spp. are poorly investigated in general. It can be assumed that they occur more frequently in the northern regions of Eurasia within epilithic lichen communities. Other species requiring attention are R. laevigata and R. sibirica which are probably widespread in Russia, although rarely found in some regions. The distribution of both taxa is worthy of further research.
Acknowledgments
We are very grateful to Dr. J. W. Sheard, a monographer of the Rinodina genus in North America, for the consultations and assistance in identifying the specimens. The authors also thank anonymous reviewers for valuable corrections and comments. The research was carried out within the state assignment of Ministry of Science and Higher Education of the Russian Federation (FWFS-2021-0002, 0290-2015-0018).
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