The genus Rinodina (Physciaceae, lichenized Ascomycota) in the Republic of Sakha (Yakutia) with a key to the species Текст научной статьи по специальности «Биологические науки»

The genus Rinodina (Physciaceae, lichenized Ascomycota) in the Republic of Sakha (Yakutia) with a key to the species

I. A. Galanina1, S. V. Chesnokov2,3, L. A. Konoreva2, 3' 4, L. N. Poryadina5, E. A. Davydov6, A. G. Paukov7

1Federal Scientific Center of East Asian Terrestrial Biodiversity of the Far Eastern Branch

of the Russian Academy of Sciences, Vladivostok, Russia 2Botanical Garden-Institute of the Far Eastern Branch of the Russian Academy of Sciences,

Vladivostok, Russia

3Komarov Botanical Institute of the Russian Academy of Science, St. Petersburg, Russia 4The Polar-Alpine Botanical Garden-Institute of the Kola Science Centre

of the Russian Academy of Sciences, Kirovsk, Russia 5Institute for Biological Problems of Cryolithozone of the Siberian Branch of the Russian Academy of Sciences, Yakutsk, Russia 6Altai State University, Barnaul, Russia 7Ural Federal University, Ekaterinburg, Russia Corresponding author. I. A. Galanina, gairka@yandex.ru

Abstract. The lichen genus Rinodina in the Republic of Sakha (Yakutia) is revised on the basis of extensive materials collected by the authors in 1974-2022. Twenty-four species of this genus have been recorded for the lichen biota of Yakutia. Eight species are recorded for the first time. Rinodina cinereovirens, R. confragosa, R. conradii, R. intermedia, R. interpolata, R. metaboliza, R. orculata, R. trevisanii. Five previously reported species were not found. Rinodina archaea, R. exigua, R. exi-guella, R. milvina, R. sophodes. A key for identification of Rinodina known in Yakutia is given. For each species, characteristic features, differences from closely related species, and distribution in Russia and the world are discussed.

Keywords: Arctic, biodiversity, biogeography, lichens, new records, North-East Asia.

Род Rinodina (Physciaceae, лихенизированные Ascomycota) в Республике Саха (Якутия), с ключом для определения видов

И. А. Галанина1, С. В. Чесноков2, 3, Л. А. Конорева2, 3 4, Л. Н. Порядина5, Е. А. Давыдов6, А. Г. Пауков7

Федеральный научный центр биоразнообразия наземной биоты Восточной Азии ДВО РАН,

пр. 100-летия Владивостока, Россия. 2Ботанический сад-институт ДВО РАН, Владивосток, Россия 3Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия 4Полярно-альпийский ботанический сад-институт КНЦ РАН, Кировск, Россия 5Институт биологических проблем криолитозоны СО РАН, Якутск, Россия 6Алтайский государственный университет, Барнаул, Россия 7Уральский федеральный университет, Екатеринбург, Россия Автор для переписки. И. А. Галанина, gairka@yandex.ru

https://doi.org/10.31111/nsnr/2023.57.2.L49

Received. 5 April 2023 Accepted: 26 October 2023 Published: 7 November 2023

Резюме. Работа основана на изучении материала, собранного авторами в 1974-2022 гг. в Республике Саха (Якутия). В результате исследования новый список лишайников рода Rinodina для Якутии составил 24 вида, 8 из которых приводятся впервые для Якутии (Rinodina cinereovirens, R. confragosa, R. conradii, R. intermedia, R. interpolata, R. metaboliza, R. orculata, R. trevisanii). Пять видов из ранее опубликованных не найдены (Rinodina archaea, R. exigua, R. exiguella, R. milvina, R. sophodes). Представлен ключ для определения видов рода Rinodina, известных в Якутии. Для каждого вида обсуждаются характерные признаки, отличия от близких видов и распространение в России и мире.

Ключевые слова: Арктика, биоразнообразие, биогеография, лишайники, новые находки, Северо-Восточная Азия.

The genus Rinodina (Ach.) Gray belongs to the family Physciaceae Zahlbr. and comprises approximately 300 species worldwide (Sheard, 2010). Rinodina is polyphy-letic, and its members are usually characterized with crustose thalli, lecanorine apo-thecia, 2-celled brown ascospores with inner wall thickenings and Lecanora-type asci (Nadyeina et al, 2010).

Rinodina species are generally difficult to identify due to the variety of spore types. However, a study of the genus Rinodina in North America (Sheard, 2010) was helpful to our understanding of this genus in Northeastern Asia (Sheard et al., 2017). In Russia, the genus Rinodina was actively studied in the last decade, but most of these investigations concern the south of the Far East (Galanina et al., 2011, 2018, 2021a, c; Galanina, 2013, 2016a, b, 2019; Sheard et al,, 2017; Galanina, Ezkhin, 2019; Konoreva et al., 2018; Yakovchenko et al., 2018; Galanina, Yakovchenko, 2021; Galanina, Ohmura, 2022). In contrast, its northern part (Chukotka Peninsula, Magadan Region, and Yakutia) remains rather poorly studied and requires revision, although some Rinodina species were reported from this region as indicated below (Almquist, 1879, 1883; Savicz, Elenkin, 1950; Afonina et al, 1979, 1980; Korolev, Tolpysheva, 1980; Andreev, 1983, 1984; Makarova, Perfiljeva, 1984; Makarova, 1985, 1989, 1998; Makarova et al, 1988; Kotlov, 1991, 1993a, b, 1995, 2004; Samarskii et al, 1997; Poryadina, 1999a, b, 2001, 2008, 2010, 2020a, b; Zhurbenko et al, 2002, 2005; Vershinina et al, 2012, 2015; Chesnokov et al., 2016; Galanina et al., 2021b, 2022). The aim of this study is to summarise the knowledges on the genus Rinodina for the Republic of Sakha (Yakutia).

The first report of the genus Rinodina in Yakutia was found in the works of Alm-quist (Almquist, 1879, 1883, not seen, cited after Savicz, Elenkin, 1950) for the Preo-brazhen'ya Island at the mouth of the Khatanga Bay (Anabarsky District): R. exigua (Ach.) Gray, R. mniaroea (Ach.) Körb., R. turfacea (Wahlenb.) Körb. Much later, new information appeared for the Rinodina species from the forest-steppe landscapes in the middle reaches of the Indigirka River (northeast of Yakutia), including R. archaea (Ach.) Arnold, R. terrestris Tomin, and R. turfacea (Afonina et al., 1979, 1980). Also, for the northeast of Yakutia, the species R. sophodes (Ach.) A. Massal. and R. turfacea were reported at the mouth of the Sukharnaya River on the coast of the East Siberian Sea (Andreev, 1984), and R. mniaroea, R. roscida (Sommerf.) Arnold and R. turfa-cea were reported from Medvezh'i islands and mouth of the Kolyma River (Andreev,

1983; Zhurbenko et al., 2005). The species R. archaea, R. bischoffii (Hepp) A. Massal., R. turfacea were reported for the territory of the northwest of Yakutia (the vicinity of the village of Saskylakh and the coast of the Laptev Sea) (Makarova, Perfiljeva, 1984; Makarova, 1985).

For Arctic Yakutia (Novosibirskie Islands and the Lena River delta) the species R. archaea, R. bischoffii, R. olivaceobrunnea C. W. Dodge et G. E. Baker, R. roscida, R. turfacea were reported (Makarova et al., 1988; Makarova, 1989, 1998; Samarskii et al, 1997; Zhurbenko et al., 2002). For central, southern and eastern Yakutia (Aldan-sky, Olekminsky, Tomponsky districts) the species R. archaea, R. subpariata (as R. de-geliana Coppins), R. demissa (Korb.) Arnold, R. excrescens Vain., R. exigua, R. freyi H. Magn., R. laevigata (Ach.) Malme, R. milvina (Wahlenb.) Th. Fr., R. mniaroea, R. oleae Bagl., R. olivaceobrunnea, R. pyrina (Ach.) Arnold, R. septentrionalis Malme, R. sophodes, R. terrestris, and R. turfacea were reported (Poryadina, 1999a, b, 2001, 2008, 2010, 2020a, b; Vershinina et al,, 2012, 2015; Chesnokov et al,, 2016; Galanina, 2016a).

The Republic of Sakha (Yakutia) is the largest region of the Russian Federation and is located in the northwestern part of the Far East. Its total area is 3.103.200 square km. It measures 2000 km from north to south and 2500 km from west to east. About 40% of the territory is located inside the Arctic Circle. The climate is extremely continental, characterized by long winters and short summers. The temperature difference between the coldest month (January) and the warmest month (July) is 70-75 degrees. The absolute minimum temperature almost everywhere in the Republic is below -50 °C. In Oymyakon there is a cold pole of the Northern Hemisphere of the planet, where a temperature of -71.2 °C has been recorded (1924). The average air temperature in Yakutia is +20°C in July and -39°C in January. Annual precipitation for most of the territory is 200-250 mm, in the south and south-west — 350-500 mm. During the year, precipitation is unevenly distributed, in the cold period (from November to March) only 15-20% of the total falls, in the warm period (from April to October) — 75-80%, i. e. 4-5 times more. A particularly small amount of precipitation (from 150 to 250 mm per year) falls on the coast and islands, the Yansk and Oymyakon plateaus, the Verkhoyansk and Momo-Selennyakh basins, as well as the Central Yakut plain. In the foothills and mountainous areas, the amount of precipitation increases to 400 mm on the watersheds of the Olekma, Chara, and Aldan plateau and up to 500700 mm on the western slopes of the Aldan-Uchursky and spurs of the Verkhoyansky ranges (Klimat..., 2001). Due to the small amount of precipitation falling in winter, the snow cover is thin over most of the territory. The number of days with snow cover ranges from 200-210 in the south of Yakutia to 250 in the tundra zone. Almost the entire territory of Yakutia lies in the zone of continuous permafrost (Troeva et al., 2010; Raznoobrazie..., 2005).

More than two thirds of the territory are occupied by mountains and high plateaus (the highest point is Mount Pobeda up to 3003 m high). The North Siberian Lowland lies in the north and Central Yakutskaya Lowland in the east of the region.

Rivers drain into the Arctic Ocean. A large variety of natural conditions are found in Yakutia, from polar deserts on the islands, and tundra to forest-steppes with altitudi-nal zonation in the mountains. Most of the territory belongs to the middle taiga zone, with approximately 80% of the territory to be occupied by forests (Troeva et al, 2010; Raznoobrazie..., 2005).

Material and Methods

During the study, herbarium material stored in the herbaria of Russia (LE, SASY, VLA) and the authors' collections from different districts of the Republic of Sakha (Yakutia) were studied. The collections of A. V. Galanin from midstream of the Vi-lyuy River in the region of the tukulans (sand dunes) were also studied. All of the localities are shown in Fig. 1. Materials were identified by the first author in the laboratory of Botany of the Federal Research Center for Biodiversity of the Far East Branch of Russian Academy of Sciences. The investigation of anatomical and morphological features of lichens was made using the microscopes Zeiss Axioplan 2 and Stemi 2000-C. The study of spore structures and measurements were made using immersion oil at 1000x magnification. Anatomical examination used hand-cut sections mounted in water, and the following reagents: 10% KOH (K) and C6H4(NH2)2 (P). The protocol of Meyer and Printzen (2000) was used to characterize the epihymenium pigments. Measurements of ascospores are presented as percentiles (5)25-75(95) |im length x breadth, thus excluding outliers. Secondary products were analyzed by standard thin-layer chromatography techniques (Culberson, Kristinsson, 1970; Orange et al., 2001) in solvent systems A (toluene: 1,4-dioxane: acetic acid, 180:45:5), B (hexane: diethyl ether: formic acid, 140:72:18), C (toluene: acetic acid, 170:30), and E (cyclohexane: ethyl acetate, 75:25). Recent publications of Sheard (Sheard, 2010, 2018; Sheard et al., 2017) have been used for specimen identification. The study of secondary metabolites (TLC) was carried out by A. G. Paukov and L. A. Konoreva, and molecular genetic studies by E. A. Davydov and L. A. Konoreva.

The distribution of Rinodina species in the world is given using the following literature data: Arctic (Alstrup, 1986; Mayrhofer, Sheard, 1988; Andreev et al., 1996; Zhurbenko et al., 2005; Kristinsson et al., 2010), Europe (Mayrhofer, Poelt, 1979; Mayrhofer, 1984; Tonsberg, 1992; Mayrhofer, Moberg, 2002; Giralt et al, 1993, 1994; Giralt, Mayrhofer, 1995; Wirth, 1995; Giralt, van den Boom, 1996; Giralt, 2001, 2010; Mayrhofer et al, 2001; Aragon et al, 2004; Spribille et al, 2006; Mayrhofer, Sheard, 2007; Giavarini et al, 2009; Wirth et al, 2013), Asia (Vezda, 1965; Schubert, Kle-ment, 1971; Golubkova, 1981; Hauck, Javkhlan, 2006; Kurokawa, Kashiwadani, 2006; Hauck et al., 2013a, b; Yakovchenko et al., 2018; Galanina, Ohmura, 2022), Russia, general publications (Kotlov, 2008; Spisok..., 2010; Galanina et al., 2021c), Caucasus (Urbanavichus et al, 2010), Siberia (Almquist, 1879, 1883; Vainio, 1928; Magnusson, 1936, 1947; Makryi, 1986, 2008; Sedelnikova, 1990; Urbanavichene, Urbanavichus, 1998, 2008, 2009; Davydov, 2001; Urbanavichene, 2010; Davydov, Printzen, 2012; Kharpukhaeva, 2013; Chesnokov, Konoreva, 2015; Chesnokov, 2017; Kharpukhaeva,

Galanina, 2022), Yakutia (Andreev, 1983; Makarova, Perfiljeva, 1984, 1988; Makarova, 1985, 1989, 1998; Makarova et al, 1988; Samarskii et al, 1997; Poryadina, 1999a, b, 2001, 2003, 2005, 2006, 2008, 2010, 2020a, b; Zhurbenko et al, 2002; Vershinina et al., 2012, 2015; Chesnokov et al., 2016; Galanina, 2016a; Galanina et al., 2022), Far East (Afonina et al, 1979; Korolev, Tolpysheva, 1980; Makarova, Katenin, 1983, 1992; Andreev, 1984; Insarov, Pchelkin, 1984; Kotlov, 1991, 1993a, b, 1995, 2004, 2008; Skirina, 1996, 2012, 2015; Tchabanenko, 2002; Himelbrant et al, 2009, 2021; Himelbrant, Stepanchikova, 2011; Rodnikova, 2012; Velikanov, Skirina, 2012; Galanina, 2013, 2016b, 2019; Kondratyuk et al, 2013; Yakovchenko et al, 2013; Sheard et al, 2017; Konoreva et al, 2018; Galanina, Ezhkin, 2019; Galanina et al, 2011, 2018, 2021а, b; Galanina, Yakovchenko, 2021; Chesnokov, Konoreva, 2022); North America (Sheard, 1995, 2010, 2018; Thomson, 1997; Sheard et al, 2011); South America (Mayrhofer et al, 2001); Australia (Kaschik, 2006).

Fig. 1. Collection localities of Rinodina in the Republic of Sakha (Yakutia).

Results

In total, 19 species Rinodina were known for the Republic of Sakha (Yakutia) before the present study. As a result of our research, the list of Rinodina species of Yakutia includes 24 taxa, eight of which are new for the region [R. cinereovirens, R. confra-gosa (Ach.) Körb., R. conradii Körb., R. intermedia Bagl., R. interpolata (Stirt.) Sheard, R. metaboliza Vain., R. orculata Poelt et M. Steiner, R. trevisanii (Hepp) Körb.], and five species were not found or excluded because of misidentification (R. archaea, R. exigua, R. exiguella, R. milvina, R. sophodes].

Rinodina milvina (Poryadina, 2001), was identified as Buellia sp. (brown hypo-thecia, no thallus margin, Buellia-type spores) and therefore excluded. Rinodina de-missa (Poryadina, 2001) was also excluded, the specimen belongs to Dimelaena oreina (Ach.) Norman. The material of Rinodinapyrina by Vershinina et al. (2012, 2015) requires verification, but these specimens are not available for revision. Other samples of R. pyrina from this region (Poryadina, 1999a, 2001, 2005) were reidentified by us as R. laevigata. However, R. pyrina is known for Yakutia based on another specimen (see below).

Two samples of Rinodina exigua (Makarova, 1998; Poryadina, 2001) we identified as R. turfacea and R. sibirica H. Magn., respectively. We were unable to verify another record of R. exigua in the literature (Vershinina et al., 2012, 2015) as the samples were not available for study. Rinodina exiguella (Vainio) H. Magn. (Poryadina, 2005) was reidentified as R. laevigata.

Samples of R. archaea (Poryadina, 2001, 2008, 2020a, b) were identified as R. ci-nereovirens Vain., R. laevigata, R. sibirica, and R. turfacea, respectively. The sample of R. sophodes (25 VII 1993 Poryadina 1993-07-25/11-3-4) was identified as R. sibirica.

The complete labels of examined specimens are in the Supplementary file.

New species for the Republic of Sakha (Yakutia) Rinodina cinereovirens (Vain.) Vain.

Rinodina cinereovirens is characterized by a thin light gray or brownish gray thallus without vegetative propagules. The apothecia quickly become narrowly attached, sometimes almost stipitate (to 0.6-1.0 mm in diam.) with a plane, black disc. The thalline margin is entire and typically persistent (80-120 |im wide), with a cortex expanded to 20-60 |im wide below. Rinodina cinereovirens has Physcia-type spores of Type A development, (21.5)23.0-25.5(27.5) x (10.0)11.5-13.5(14.0) ^m and a well developed torus. Crystals absent in cortex and present in medulla (sphaerophorin). Spot tests all negative, the secondary metabolite sphaerophorin turns blue-white in longwave UV.

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).

Distribution. Eurasia [Scandinavia (Norway, Sweden, and Finland), Russia (from Leningrad Region to Kamchatka Territory and Magadan Region)], North America (USA: Alaska; Canada: northern Manitoba, northern Ontario, Newfoundland, New Brunswick).

Ecology. On bark of Alnus sp., Larix gmelinii (Rupr.) Kuzen. and Pinus sp. in mixed forests and on wood in mountain epilithic-lichen tundra.

Specimens examined: Nizhnekolymsky District, Poryadina, SASY L-2008-09-19/24-3; Tom-ponsky District, Poryadina, SASY L-1996-07-14/12-7-8 (det. as Rinodina archaea), Poryadina, SASY L-1996-07-04/6-1; Olekminsky District, 16 VII 2008, Poryadina, SASY; Aldansky District, 16 VII 2000, Poryadina, SASY; Neryungrinsky District, 30 VI 2015, Konoreva SC-238, LE L-25116; Sofronova, SASY L-2007-07-04/15-3.

Rinodina confragosa (Ach.) Körb.

Rinodina confragosa is characterized by its thin, light-gray, areolate thallus and dark fimbriate prothallus. Its apothecia are narrowly attached, to 0.7-1.0 mm in diam., with a black and plane disc (Fig. 2 C). The thalline margin is entire and typically persistent, up to 100 |im wide, with a columnar cortex expanded to 30-50 |im wide below. Rinodina confragosa has Physcia-type spores of Type A development, (16.5)19.0-20.5(22.5) x (8.0)9.0-10.0(11.0) ^m and a pigmented torus (Fig. 2 D). Spot tests are K+ yellow, P+ faint yellow.

Rinodina confragosa is similar to R. jacutica Galanina et Konoreva in thallus morphology and spot tests, but the latter possesses Dirinaria-type spores (16.5)18.0-20.0(23.0) x (7.5)8.5-10.0(11.5) (Galanina et al., 2022). Rinodina confragosa is closely related to R. brandii Giralt et van den Boom (a rare European species) which differs in the presence of pannarin located only in the epihymenium (Giralt, van den Boom, 1996).

Distribution. Eurasia [from northern Europe to Turkey, Caucasus, Mongolia, Russian Far East (Primorye Territory and Southern Kuril Islands), and Japan], North America (widely distributed throughout the western United States and into Alberta, and rare in coastal Newfoundland in the east), South Africa, and Australia.

Ecology. On quartz pebbles in sparse pine forest on a river terrace.

Specimen examined: Zhigansky District, Galanina, VLA L-3148.

Rinodina conradii Körb.

Rinodina conradii is characterized by its 3-septate spores with Type B development. This species has a light to dark gray, brownish-gray continuous thallus with spot tests all negative.

Rinodina conradii can be mistaken for R. intermedia but the latter species differs in submuriform spores with Type A development. Rinodina intermedia can have rose-violet medullary reaction with KOH (Sheard, 2010).

Distribution. Widespread in the world in both hemispheres, with a cold temperate to oroarctic distribution, in Russia it is widespread from the Murmansk and Leningrad regions to the Primorye Territory in the Far East.

Ecology. On bark in floodplain forests, mossy slope with birch trees at 807 m a. s. l. Specimens examined: Tomponsky District, 17 VII 2016, Konoreva SC-302, LE L-25121.

Rinodina intermedia Bagl.

Rinodina intermedia is characterized by its submuriform spores with Type A development. This species has a thin light gray-green to light brown continuous thallus and grows on soil or terricolous mosses.

Rinodina intermedia can be confused with R. conradii as discussed above. Distribution. Widespread in the world: Europe (from the Iberian Peninsula in the south to the Channel Islands in the north), Africa (Kenya), Asia (Himalayas, Japan), North America (southwestern US states, Mexico), South America (Ecuador). In Russia the species is known from Kaliningrad Region, southern Siberia, Olkhon Island in Lake Baikal, Buryatia (Volcano valley), Trans-Baikal Territory (Sokhondins-kiy Nature Reserve), Khabarovsk Territory (Kukanskii Ridge).

Ecology. On bark of Larix gmelinii and on a dead tree in larch forests. More information about this species is in Galanina (2019).

Specimens examined: Oymyakonsky District, 8 VII 2016, Konoreva SC-291, LE L-25117; 13 VII 2016, Yatsyna SC-286, LE L-25118.

Rinodina interpolata (Stirt.) Sheard

Rinodina interpolata is characterized by a Physcia-type to Physconia-type spores (14)15-17(20) x (6)7-8(9) |m of Type A development (Mayrhofer, Moberg, 2002). This species has a gray to dark brown rimose-areolate thallus and grows on siliceous rocks.

Rinodina interpolata might be confused with R. parasitica H. Mayrhofer et Poelt which also has Physcia-Physconia-type spores, but they have more rounded lumina and this species grows on the Aspicilia. Rinodina interpolata can also be confused with R. tephraspis (Tuck.) Herre, but it is distinguished by its larger Teichophila-type spores, (17.0)20.0-21.0(24.0) x (8.0)10.5-11.5(14.0) |m (Sheard, 2010).

Distribution. Europe (central part, British Isles, Scandinavia, and Iceland), Russia (Republic of Karelia, the Caucasus, the Urals, and southern Siberia), North America.

Ecology. On rocky outcrops in the river valley.

Specimen examined: Tomponsky District, Poryadina, SASY L-1996-07-08/9-5-6-7.

Fig. 2. A — apothecia of Rinodina calcigena; B — Bischoffii-type spores of R. calcigena with unusually narrow lumina canals during early development, Physconia-like lumina, lack of a torus, and a very poorly developed septal pigment band at maturity; C — apothecia of R. confragosa; D — Physcia-type spores of R. confragosa with retaining septal and apical wall thickening and a pigmented torus; E — apothecia of R. pyrina; F — mature spores of R. pyrina with Physconia-like lumina, torus lacking, and septal disck present. Scale bars: A, C, E — 0.5 mm; B, D, F — 10 |m.

Rinodina metaboliza Vain.

Rinodina metaboliza is characterized by its Dirinaria-type spores (13.5)17.5-19.0(23.0) x (8.5)9.0-10.0(11.0) |m of Type B development that typically inflate at the septum in KOH. The species is very variable in thallus morphology, spore size, and apothecial convexity.

Rinodina metaboliza can be confused with R. endospora Sheard which also has Dirinaria-type spores, but they are larger size (20.5)22.0-24.0(27.5) x (8.5)9.0-10.0(11.0) |m (Sheard, 2010).

Distribution. Eurasia [Scandinavia (central Sweden and Norway), Russia (the republics of Karelia and Komi, South Siberia — Eastern Sayan Mountains, and the Far East — Magadan Region)], North America (from coastal Alaska to eastern Canada and the United States).

Ecology. On bark of Alnus sp., Betula sp., Picea sp., Populus sp., P. tremula L., Salix sp., in floodplain and mixed forests at 130-295 m a. s. l.

Specimens examined: Bulunsky District, Poryadina, SASY L-2009-07-19/10-1; Zhigansky District, Galanina, VLA L-3152, L-3153; Vilyuysky District, Galanina,VLA L-2937, Galanina, VLA L-2963, Galanina, VLA L-2938, L-2956, L-2957, L-2958, L-2959, L-2970, L-2971, L-2977, L-2978, L-2980, L-3123, L-3127, L-3133, Galanin, VLA L-2950, L-2951, L-2966, L-2967, L-2969, L-2972, L-3121, L-3122, L-3124; Kobyaysky District, Poryadina, SASY L-2002-07-07/0-2; Tom-ponsky District, 21 VII 2016, Konoreva SC-307, LE L-25119; Poryadina, SASY L-2016-07-21/24-1, L-2011-07-07/3-4; Khangalassky District, Ahti, SASY L-2002-06-30/0-1; Olekminsky District, 16 VII 2008, Poryadina, SASY.

Rinodina orculata Poelt et M. Steiner

Rinodina orculata is characterized by a Physcia-Physconia-type spores (12.5)16.0-16.5(20.0) x (6.0)8.0-8.5(9.5) |m of Type A development. The species is very variable in thallus and apothecia morphology (Sheard, 2010).

Rinodina orculata can be confused with R. trevisanii, which has larger [(14.5)18.0-18.5(23.0) x (7.5)8.5-9.0(10.5) |m] Physconia-type spores. Unlike R. trevisanii, R. or-culata has smaller spores on average and often shows persistent Physcia-type apical thickening (see Sheard, 2010).

Distribution. Eurasia [rarely and scattered in Germany, Scandinavia and often in the south (Iberian Peninsula, Portugal), Russia (Caucasus, Krasnoyarsk Territory, the Urals)], North America (widespread from the Gaspe Peninsula in eastern Canada to the west coast of North America, and often in the Rocky Mountains).

Ecology. On bark of Pinuspumila (Pall.) Regel in mixed forests at 836 m a. s. l.

Specimen examined: Neryungrinsky District, 30 VI 2015, Konoreva SC-254, LE L-25120.

Rinodina trevisanii (Hepp) Körb.

Rinodina trevisanii is characterized by a Physconia-type spores (14.5)18.0-18.5(23.0) x (7.5)8.5-9.0(10.5) |m of Type A development (Fig. 3 B, C). The species

possesses a thin continuous to rimose thallus and scattered apothecia with the discs frequently becoming convex, and the narrow thalline margins often becoming biato-rine (Sheard, 2010). Young apothecia of Rinodina trevisanii are with plane discs and prominent apothecial margins (Fig. 3 A).

Rinodina trevisanii can be confused with R. archaea and R. orculata. See above for differences from R. orculata. Rinodina archaea is distinguished by a thick and areolate thallus, and apothecia quickly become contiguous, and angular by compression, with persistently plane discs and a well-developed thalline margin. Also, the spores of R. archaea have more prolonged Physcia-type stage of development. Rinodina convexula is synonymous with R. trevisanii (Mayrhofer, Sheard, 2007; Sheard, 2010).

Distribution. Europe [the Alps, rarely in Scandinavia (Finland and Sweden)], Asia (Mongolian part of Altai, Saur Range in Kazakhstan). Rinodina trevisanii is poorly studied in Russia and is rarely recorded [eastern Siberia (Krasnoyarsk Territory), Far East (Khabarovsk Territory, Bastak Nature Reserve)]. In North America, it is rarely found in Arizona, in the Cascade Mountains from Oregon to British Columbia.

Ecology. On bark in floodplain forests and on wood in thickets of dwarf birch at 807-1046 m a. s. l.

Specimens examined: Tomponsky District, 17 VII 2016, Konoreva SC-302, LE L-25121; Oymyakonsky District, Poryadina, SASY L-2016-07-13/13-2.

Species previously noted for the Republic of Sakha (Yakutia) Rinodina bischoffii (Hepp) A. Massal.

Rinodina bischoffii is variable in morphology. It is characterized by an inconspicuous, often endolithic thallus, but sometimes it can be well developed. Apothecia range from submerged in the thallus to narrowly attached, the disc quickly becomes convex to hemisphaerical, the thallus margin is concolorous with the thallus. Rinodina bischoffii has spores Bischoffii-type of Type A development, the torus is absent, the walls are not ornamented. The hymenium contains oil drops.

Rinodina bischoffii is highly polymorphic species and may or may not lack oil droplets in hymenium, J. Sheard (2010) considers R. immersa (Körb.) J. Steiner to be its synonym. Rinodina bischoffii is similar to R. guzzinii, but the latter has a more developed thallus and larger spores (21-22 |im length), and the spores of R. bischoffii (16-18 |im) have a more ellipsoidal shape, in contrast to R. guzzinii, in which the spores are broadly round at the ends. Rinodina castanomelodes H. Mayrhofer et Poelt is also a similar species, but it has a well-developed thallus which is almost lo-bate at the edges, larger apothecia (up to 0.9 mm in diam.) and the spores are also wide-rounded at the ends. Giralt (2010) considers R. castanomelodes to be a variety of R. bischoffii. In the Arctic, R. calcigena can be confused with R. bischoffii, but the former has larger, broadly ellipsoid spores (19-20 |im length), and a rapidly disappearing thin canal. A pigmented belt around the septum is visible only in mature spores (Sheard, 2010).

Fig. 3. A — young apothecia of Rinodina trevisanii with the plane discs and prominent apothecial margins; B — mature spores of R. trevisanii have Physcia- to Physconia-type lumina with relatively

narrow canals when immature; C — overmature spore with inflated lumina, thin apical walls

and narrow torus.

Scale bars: A — 0.5 mm; B, C — 10 |im.

Distribution. Widespread in both hemispheres: Eurasia [Europe, Russia (from the Arctic to Southern Siberia and from the European part to the Far East], North and South Africa, Asia, and Australasia, North America (widespread in the temperate zone, excluding the northwestern Pacific and southeastern states). In Yakutia, Rinodina bischoffii was found on the New Siberian Islands (Zemlya Bunge Island) and in the Saskylakh village vicinity on lime-containing rocks (Makarova, 1985; Makarova et al., 1988).

Ecology. On lime-containing rocks at 507 m a. s. l.

Specimen examined: Anabarsky District, Perfiljeva, SASY L-1981-08-11/42-5.

Rinodina calcigena (Th. Fr.) Lynge

Rinodina calcigena is characterized by a thin gray or ochraceous-brownish gray thallus present around the base of the apothecia or as scattered areoles on the substrate. Its apothecia are erumpent at first, broadly attached, to 0.5-0.9 mm in diam., the disc is black and plane (Fig. 2 A). The thalline margin is entire and narrow, typically persistent or rarely disappearing. Rinodina calcigena has Bischoffii-type spores of Type A development (Fig. 2 B), (15.0)19.0-20.0(23.5) x (9.0)11.5-11.5(13.0) |m and torus absent. The spores of R. calcigena have unusually narrow lumina canals during

development for the Bischoffii-type, never show apical thickening during development, a characteristic of the Physconia-type. A pigmented band is rarely evident around the septum at maturity but immature spores may possess very broad pigmented bands (Sheard, 2010). Spot tests are all negative.

Rinodina calcigena is closely related to R. bischoffii but differs in its longer spores with unusually narrow lumina canals during development. Rinodina calcigena is closely related to R. guzzinii but differs in its shorter spores with unusually narrow lumina canals during development, a poorly developed thallus and very symmetrical apothecia with a narrow or disappearing thallus margin (Mayrhofer, Sheard, 1988; Sheard, 2010).

Distribution. Eurasia [Europe (Scandinavia), Russia (Polar Urals, Novaya Zemlya, Southern Siberia, and Kamchatka)], North America (Arctic, Rocky Mountains, Greenland). In Yakutia (Mayrhofer, 1984; Kotlov, 2008).

E c o l o g y . On rock in larch forests.

Specimen examined: Tomponsky District, Poryadina, SASY L-1996-07-02/0-1-2-3 (was previously determined as R. gennarii Bagl.).

Rinodina excrescens Vain.

Rinodina excrescens is characterized by its discrete, coarse, bullate areoles that often bear erumpent soredia or blastidia. It contains pannarin, resulting in a Pd+ orange spot test, and its often lightly pruinose apothecia (pannarin) include Physcia-type spores.

Rinodina granulans Vain. is similar but its blastidia are finer and meld to form a continuous leprose crust (Giralt et al, 1994; Sheard, 2010; Galanina et al, 2011).

D i s t r i b u t i o n . Eurasia [Europe (scattered and rare in Austrian province of Styria, Croatian island of Mljet, Spain, Greek island of Crete), Russia (West Siberia, Altay and Trans-Baikal territories, and Far East (Primorye and Khabarovsk territories, Sakhalin and Shikotan islands), Japan — Hokkaido Island], North America (the Great Lakes region). Known from Yakutia (Chesnokov et al, 2016).

Ecology. On bark of Betula sp. and Larix gmelinii also on wood and mosses in larch and mixed forest at 127-1108 m a. s. l.

Specimens examined: Tomponsky District, 19 VII 2016, Konoreva SC-303, 305, LE L-25122, L-25123; 17 VII 2016, Konoreva SC-295, 297, 298, 299, LE L-25124, L-25125, L-25137, L-25142; Poryadina, SASY L-2018-09-22/8-6, L-2018-09-22/8-9, L-2018-09-22/8-10, L-2018-09-22/8-11, L-2018-09-22/8-13, L-2018-09-22/8-14; Oymyakonsky District, 13 VII 2016, Konoreva SC-292, LE L-25126; Namsky District, Poryadina, SASY L-2019-06-01/1-9.

Rinodina freyi H. Magn.

Rinodina freyi is characterized by a gray-green continuous thallus, apothecia frequently becoming contiguous on small thalli, 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) ^m of Type A development, and a heavy torus.

Rinodina freyi can be confused with R. septentrionalis, but the latter species has copper-brown thallus consisting of small discrete verrucae (convex when moist), narrowly attached and scattered apothecia.

Distribution. 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). In Russia, R. freyi was previously recorded from the Magadan Region, Kamchatka Territory, Sakhalin Region (Sakhalin Island), Khabarovsk and Primorye territories. It has also been reported from Japan and western Mongolia. Rinodina freyi is the most common species of the genus in North America, being frequent in both the East and West of the continent. In Yakutia, Rinodina freyi was found on the Tukulan Mahatta in the Vilyuysky District (Galanina, 2016a).

Ecology. On bark of Alnus sp., Picea sp., Salix sp. in different types of forest at 352 m a. s. l.

Specimens examined: Vilyuysky District, Galanina, VLA L-2952, L-2973, L-2974; Olekminsky District, 18 VII 2008, Poryadina, SASY; Lensky District, Egorova, SASY L-2002-06-24/0-2; Neryungrinsky District, 12 VII 2015, Poryadina, SASY.

Rinodina jacutica Galanina et Konoreva

The species was recently described from Yakutia (Galanina et al, 2022). It was found in the Oymyakonsky District, on stones in coniferous moss-lichen forest in a river valley.

Rinodina jacutica is well distinguished by its Dirinaria-type spores, (16.5)18.0-20.0(23.0) x (7.5)8.5-10.0(11.5) |m (Fig. 4 B, C), type A development, light-gray thallus with a slight yellowish tinge and spot tests of thallus K+ yellow and P+ yellow (atranorin), as well as by growing on stony substrate (Fig. 4 A).

Rinodina jacutica can be confused externally with the saxicolous R. santorinensis J. Steiner from Southern Europe. Rinodina santorinensis also contains atranorin and has a similar habit but differs in its Pachysporaria-type spores (14.0)15.0-21.0(22.0) x 8.0-12.0(14.0) |m, type B development, well developed torus and the presence of pannarin (P+ orange) in the cortex and sometimes in the epihymenium (Giralt, 2001). It is a maritime Macaronesian-Mediterranean species unlikely to be found in Yakutia. Rinodina gennarii is similar to R. jacutica in its Dirinaria-type spores, but its spores are smaller (12.0)15.0-16.5(19.0) |m in length, and brown thallus has no reactions with K and P. Rinodina confragosa similar to R. jacutica by its thallus morphology and spot tests, but it has Physcia-type spores (16.5)19.0-20.5(22.5) x (8.0)9.0-10.0(11.0) |m (Sheard, 2010). Rinodina brouardii B. de Lesd. is a North American species that grows on calcium-containing rocks and has the Dirinaria-type spores (15.0)18.0-18.5(22.0) x (7.5)8.5-9.0(10.0) |m. It can be distinguished by dark gray to ochraceous, areolate thallus, lacking prothallus, and negative spot test reactions (Sheard, 2010).

Distribution. Russia (Yakutia, Oymyakonsky District) (Galanina et al., 2022).

Ecology. On stone in larch forest with mosses-lichen cover at 1179 m a. s. l.

Fig. 4. Rinodina jacutica. A — habit; B, C — Dirinaria-type spores (B — note the Physcia-like lumina but lack of a torus, C — spore structure, note the septal swelling, the lack of a torus, and presence of a septal disc). Scale bars: A — 1 mm; B, C — 10 |im. Photo from Galanina et al. (2022).

Specimens examined: Oymyakonsky District, 7 VII 2016, Konoreva 65, 75, LE L-15578, L-15579; doublet, VLA L-2928.

Rinodina laevigata (Ach.) Malme

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) |m of Type A development, a thin poorly developed thallus, broadly attached apothecia with plane discs, and a usually thick lower apo-thecial cortex.

Rinodina laevigata can be confused with R. sibirica but differs by a less developed thallus, often its darker colour, smaller spores, and thick lower apothecial cortex. Often morphological and anatomical features of these two species have overlapping sizes, therefore genetic analysis is necessary for better understanding.

Distribution. Eurasia [Europe (Norway, Sweden, Finland, and Scotland), Asian Russia (from the North Caucasus to the Magadan Region)], North America (the western part, from Alaska to California and the Sierra Nevada). In Russia probably understudied. In Yakutia, Rinodina laevigata was found on the Tukulan Mahatta in the Vilyuysky District (Galanina, 2016a).

Ecology. On bark of Alnus sp., Betula fruticosa Pall., Betula sp., Crataegus sp., Duschekia sp., Juniperus sp., Larixgmelinii, Larix sp., Picea sp., Pinus sylvestris, Populus tremula in different types of forest. Rinodina laevigata is found very often in Yakutia at 70-861 m a. s. l.

Specimens examined: Bulunsky District, 20 VII 2009, Poryadina, SASY; Ust-Yansky District, Perfiljeva, SASY L-1995-07-29/46-1-2; Nizhnekolymsky District, 1977, Andreev, LE; Zhigansky District, Galanina, VLA L-3157, L-3158, L-3159, L-3160, L-3161, L-3162, L-3163, L-3164, L-3165, L-3166; Mirninsky District, Poryadina, SASY L-2006-08-12/8-2, L-2006-08-12/8-3, L-2006-08-12/8-1, L-2006-08-15/0-2, L-2006-08-17/18-1; Vilyuysky District, Galanin, VLA L-2098, L-2687, L-2688, L-2965, L-2979, L-2981, L-2982, L-3129, L-3135, L-3136, Galanina, VLA L-2973, L-2987, L-2988, L-2992, L-2998, L-3001, L-3002, L-3003, L-3004, L-3005, L-3006, L-3007, L-3008, L-3009, L-3010, L-3133; Namsky District, Poryadina, SASY L-2019-06-01/1-8; Kobyaysky District, Poryadina, SASY L-2002-07-04/2-3; Tomponsky District, 19 VII 2016, Konoreva SC-304, LE; Poryadina, SASY L-1993-09-03/1-1-2-3-4 (the two specimens together were determined as Rinodina exiguella and R. exigua); Poryadina, SASY L-1996-07-04/5-1-2, L-1996-07-05/7-1-2-3, L-1996-07-14/11-7-8-9 (det. as R. archaea); Oymyakonsky District, 8 VII 2016, Konoreva SC-291, LE L-25117; 5 VII 2016, Konoreva SC-287, LE L-25128; Khangalassky District, Poryadina, SASY L-2017-07-09/7-13; Lensky District, Egorova, SASY L-2002-06-29/0-5; Poryadina, SASY L-2014-06-01/0-1; Olekminsky District, Chikidov, SASY L-2006-07-13/0-1, L-2006-07-19/02-26; Aldansky District, 4 VII 2015, Chesnokov SC-208, LE L-25129; 10 VII 2015, Chesnokov SC-214, LE L-25130; 6 VII 2015, Chesnokov SC-210, LE L-25131; 16 VII 2000, Poryadina, SASY (two specimens).

Rinodina mniaroea (Ach.) Korb.

Rinodina mniaroea is characterized by its apothecia becoming convex with thalline margin becoming excluded, and large Physcia-type spores (20.5)24.5-25.5(30.0) x (9.5)11.5-12.5(14.5) |m of Type A development.

Rinodina mniaroea is similar to R. turfacea, which differs in producing sphaero-phorin (UV+ blue-white) and its persistently plane apothecial discs with prominent thalline margins. Another species to be distinguished from R. turfacea is R. olivaceo-brunnea, which differs in its smaller apothecia and spores.

Distribution. Eurasia [central European part, Iberian Peninsula, and Scandinavia, the Caucasus, Himalayas, Mongolia, China, Russia (widespread in the northern and mountain regions (Arctic, northern European part, North Caucasus, Siberia, Urals, Altai, Far East)], North America (common in the western Arctic, the Rocky Mountains and scattered in the eastern Arctic and Greenland). In Yakutia, Rinodina mniaroea was found in the mouth of the Kolyma River on the northeast of the region and on the Tuku-lan Mahatta in the Vilyuysky District (Zhurbenko et al, 2005; Galanina, 2016a).

Ecology. On mosses and plant debris in the birch and pine forests at 357727 m a. s. l.

Specimens examined: Vilyuysky District, Galanina, VLA L-2939, L-2940, L-2941, L-2961, L-2953; Aldansky District, 4 VII 2015, Chesnokov SC-206, LE L-25132; 13 VII 2015, Konoreva SC-244, LE L-25133; 12 VII 2015, Konoreva SC-233, LE L-25134.

Rinodina oleae Bagl.

Rinodina oleae is characterized by its relatively small Dirinaria-type spores (12.0)15.0-16.5(19.0) x (6.5)7.5-8.0(9.5) ^m of Type A or B development and innate to broadly attached apothecia (Sheard, 2010).

Rinodina oleae can be confused with R. endospora which also has Dirinaria-type spores, but they are a larger size (20.5)22.0-24.0(27.5) x (8.5)9.0-10.0(11.0) ^m. Rinodina oleae is often regarded as being synonymous with R. gennarii but this species grows on rocks in seaside habitats, while R. oleae occurs on bark (Sheard, 2010). The close relationship of these species was discussed in several papers (Giralt, Mayrhofer, 1995; Trinkaus et al., 1999; Giralt, 2001), also using molecular methods (Helms et al., 2003; Kaschik, 2006). Currently, R. gennarii is treated as being separate by Sheard (2010), and therefore the distribution of the R. oleae should be studied in Russia.

Distribution. Eurasia [Southern Europe, Russia (Sakhalin and Kuril Islands, Khabarovsk and Primorye territories, reported from many regions as a synonym of R. gennarii), China, Korea, Japan], North America, where it has a scattered distribution. In Yakutia, Rinodina oleae was found on the Tukulan Mahatta in the Vilyuysky District (Galanina, 2016a).

Ecology. On bark of Salix sp., twig of Picea sp. in the spruce forests at 62-81 m a. s. l.

Specimens examined: Zhigansky District, Galanina, VLA L-3165; Vilyuysky District, Galanina, VLA L-3145, L-3146.

Rinodina olivaceobrunnea Dodge et Baker

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) ^m of Type A development. The species has abundant and small, narrowly attached apothecia.

Rinodina olivaceobrunnea might be confused with R. turfacea, but differs in its smaller spores and apothecia, absence of a massive columnar lower cortex, and the lack of sphaerophorin. Rinodina olivaceobrunnea can also be confused with R. archaea, but it is distinguished by its Physcia-type spores, and its typically muscicolous rather than lignicolous habitat (Sheard, 2010).

Distribution. Rinodina olivaceobrunnea is distributed in both hemispheres being known in Europe, central Africa, Australasia, Antarctica, and North America. In Russia the species is widely distributed [Arctic (Novaya Zemlya, Taimyr, and Chu-kotka), Murmansk Region, Komi Republic, North Caucasus (Karachayevo-Circassian Republic), West and South-Eastern Siberia and Far East (Khabarovsk Territory)]. In Yakutia, R. olivaceobrunnea was found on the New Siberian Islands and in the Vilyuysky District (Tukulan Mahatta) (Samarskii et al., 1997; Galanina, 2016a).

Ecology. On bark of Salix sp., wood, plant debris in the spruce forests and arctic lichen-moss rubble desert at 81-1280 m a. s. l.

Specimens examined: Bulunsky District, Nikolin, SASY L-1982-08-13/16-4, 12 VIII 1980, Per-filjeva, SASY, Poryadina, SASY L-1996-07-14/11-10-12 (det. as Rinodina exigua); Vilyuysky District, Galanina, VLA L-3147; Namsky District, Poryadina, SASY L-2019-06-01/1-7; Tomponsky District, 14 VII 2016, Konoreva SC-293, SC-294, LE L-25135, L-25136; 17 VII 2016, Konoreva SC-297, LE L-25137.

Rinodina pyrina (Ach.) Arnold

Rinodina pyrina is characterized by its crowded apothecia, large algal cells and its small, Dirinaria-type spores with Physconia-like lumina that quickly becoming expanded to exclude the septal and apical wall thickenings (Fig. 2 E, F). The dark pigmented apices of the paraphyses form a dark brown rather than red-brown epihymenium (Mayrhofer, Moberg, 2002; Sheard, 2010, 2018; Sheard et al., 2011).

D i s t r i b u t i o n . The species is widespread worldwide: Europe, Asia, northern Africa, North America, and Australasia. In Russia, the species is widely distributed according to the literature data, but it seems to be very rare based on studying samples from the Far East and Yakutia. Rinodina pyrina was reported from Yakutia (Poryadina, 1999a, 2005; Vershinina et al., 2012, 2015). The specimens (Vershinina et al., 2012, 2015) were not available for verification. One specimen (4 VII 1996, Poryadina, SASY L-1996-07-04/5-1-2) (Poryadina, 1999a) was reidentified as R. laevigata during this study. Rinodina pyrina was confirmed to occur in Yakutia during specimen revision from the Central Yakutia in the Namsky District.

E c o l o g y . On wood on the steppe slope surrounded by larch forest.

Specimen examined: Namsky District, Poryadina, SASY L-2019-06-01/3-5-6 with Rinodina sibirica.

Rinodina roscida (Sommerf.) Arnold

Rinodina roscida is characterized by its very large Physcia-like spores (sometimes similar to Dirinaria-type), (22.5)30.0-32.0(39.5) x (10.5)12.5-13.5(16.0) ^m of Type A or rarely B development, often with submucronate apices of spores and the lack of a torus.

Rinodina roscida can only be confused with R. turfacea and differings in its light gray thallus lacking sphaerophorin, pruinose apothecial discs. Rinodina roscida also has similar spore type and type of development to R. terrestris but R. roscida has much larger apothecia and spores (Sheard, 2010).

Distribution. Rinodina roscida is widespread in the northern and mountainous regions of Northern Hemisphere: Arctic (from Scandinavia to Chukotka and from Alaska to Greenland), Asia (Caucasus, Himalayas, Mongolia, China, Southern Siberia (Trans-Baikal Territory, Sayany), North America (Rocky Mountains and with an outlier population in Newfoundland). In Yakutia, R. roscida was found in the mouth of the Kolyma River, on the New Siberian Islands (Kotelnyi Island) (Makarova, Perfil-jeva, 1988; Samarskii et al., 1997; Zhurbenko et al., 2005).

Ecology. On mosses and plant debris in pine forest, shrub-lichen-moss tundra, and moss-ledum-blueberry flowing swamp at 81-410 m a. s. l.

Specimens examined: Anabarsky District, 4 VIII 1974, Perfiljieva, SASY, Perfiljieva, SASY L-1981-08-09/32-1-2; Bulunsky District, Karpov, SASY L-1980-08-12/12-2; Vilyuysky District, Galanina, VLA L-2997; Aldansky District, 16 VII 2015, Konoreva, LE L-13353; 13 VII 2015, Konoreva SC-237, LE L-25138; 13 VII 2015, Chesnokov SC-215, LE L-13333, L-25139.

Rinodina septentrionalis Malme

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) |m with a well-developed torus and Type A development. Rinodina septentrionalis has copper-brown thallus consisting of small discrete verrucae (convex when moist), and narrowly attached and scattered apothecia.

The differences from the similar Rinodina freyi see under that species.

Distribution. Eurasia (Arctic, boreal zones and mountainous regions from northern Scandinavia to Chukotka and Kamchatka, and southbound to Georgia, Kazakhstan, Altai Mts, Sikhote-Alin Mts, Japan); North America (Arctic, extending southwards in the Rocky Mountains to Colorado). In Yakutia, Rinodina septentrionalis was found in the Suntar-Khayata Ridge and Tukulan Mahatta (Poryadina, 2001; Galanina, 2016a).

Ecology. On the bark of Alnus sp., Larix sp., Populus tremula, Salix sp., twig of Crataegus sp. and Picea sp. in different forest and moss-ledum-blueberry flowing swamp at 31-579 m a. s. l.

Specimens examined: Zhigansky District, Galanina, VLA L-3153; Vilyuysky District, Galanina, VLA L-2963, L-2971, L-2989, L-2990, L-2991, L-3137, L-3138, Galanin, VLA L-1246, L-2974, L-2981, L-2983, L-2985; Kobyaysky District, Poryadina, SASY L-2002-07-07/0-1, L-2002-07-15/0-17; Aldansky District, 16 VII 2000, Poryadina, SASY. Neryungrinsky District, Chikidov, SASY L-2007-07-01/9-2-8.

Rinodina sibirica H. Magn.

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) |m with a well-developed torus, the darkly

pigmented walls, and Type A development. Rinodina sibirica has erumpent apothecia broadly attached at first, becoming narrow at the base, scattered, with the dark brown to black discs that may become convex or even hemisphaerical.

Rinodina sibirica might be confused with R. archaea, but the latter has Physco-nia-type spores and is mainly lignicolous (Sheard, 2010). The young apothecia of R. sibirica with plane discs can be confused with R. cinereovirens, which has large Phy-scia-type spores and crystals in medulla (sphaerophorin, UV+ blue-white). Specimens of R. sibirica have often been referred to R. sophodes, which differs in its small Milvi-na-type spores and European distribution (Sheard, 2010).

Distribution. This species has a wide Amphiberingian range, only slightly extending to Europe in the Ural Mountains (Galanina et al., 2021c). 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., 2021a, b, c). The species is known from North America (Thomson, 1997, as R. granulans; Sheard, 2010). Rinodina sibirica was also reported from Yakutia (Kotlov, 2008).

Ecology. On the bark of coniferous and deciduous trees in different forest types and flowing swamp at 62-1640 m a. s. l. More information about this species is in Galanina et al. (2021).

Specimens examined: Bulunsky District, 2 VII 1998, Makarova, LE; Poryadina, SASY L-2009-07-22/15-1; Zhigansky District, 8 VI 1901, Cajander, H; 6 VIII 1901, Cajander (11 specimens s. n.), H; 7 VIII 1901, Cajander (3 specimens s. n.), H; 16 VIII 1901, Cajander (3 specimens s. n.), H; 17 VIII 1901, Cajander, H; Cajander (2 specimens s. n.), H; Verkhoyansky District, Ahti, Efimo-va, H 65400, 65400b; Momsky District, Poryadina, SASY L-1993-07-25/11-3-4 (det. as Rinodina sophodes); Mirninsky District, 15 VIII 2006, Poryadina, SASY (det. as R. exigua and as R. archaea); Poryadina, SASY L-2006-08-12/8-4; Vilyuysky District, 23 VII 2015, Galanina, VLA; 25 VII 2015, Galanina, VLA; 14 VII 2015, Galanina, VLA; 12 VII 2015, Galanina Ya-15-18, VLA; 11 VII 2015, Galanina, VLA; 4 VI 2012, Galanina, VLA L-2166, L-2167, L-2168, L-2169, L-2973, L-3128, L-3131, L-3132, L-3134; Galanin, VLA; 2 VI 2012, Galanin, VLA; Galanin, VLA L-2098, L-2158, L-2159, L-2160, L-2161, L-2162, L-2163, L-2164, L-2165, L-3126, L-3130; Namsky District, Poryadina, SASY L-2018-09-22/8-6 (det. as R. archaea), L-2019-06-01/3-5-6, L-2019-06-01/1-9; Ust-Al-dansky District, Zakharova, SASY L-2011-08-23/0-3; Kobyaysky District, 9 VII 2002, Poryadina, SASY L-2002-07-09/0-6; 9 VII 2002, Poryadina, SASY; 13 VII 2002, Poryadina, SASY (det. as R. exigua); 25 VIII 1995, Poryadina, SASY L-1995-08-25/15-1, L-1995-08-25/15-2; 10 VII 2002, Poryadina, SASY; Poryadina, SASY L-2002-07-09/0-3, L-2002-07-09/0-5); 14 VII 2002, Poryadina, SASY; 14 VII 2002, Poryadina, SASY; 8 VII 2002, Poryadina, SASY; 18 VII 2002, Poryadina, SASY; Poryadina, SASY L-2002-07-15/0-16, L-2002-07-04/2-2; Tomponsky District, 17 VII 2016, Konoreva SC-301, LE L-25140, 17 VII 2016, Konoreva SC-296, 299, LE L-25141, L-25142; Poryadina, SASY 1996-07-05/7-1-2-3 (det. as R. archaea); Poryadina, SASY 1996-07-01/3-1-2-3 (det. as R. archaea); 14 VII 1996, Poryadina, SASY (det. as R. archaea); Khangalassky District, Ahti, H 61412, 61412d, duplet in SASY L-2002-06-30/0-1, Ahti, Efimova, H 42348, duplet in SASY; Ahti, Timofeev, H 64406d, Ahti, H 61800, Poryadina, SASY L-2007-09-15/0-2; 20 IX 2007, Poryadina,

SASY; Olekminsky District, 18 VII 2008, Poryadina, SASY; Poryadina, SASY L-2008-07-26/27-6; Aldansky District, Poryadina, SASY L-2015-07-16/41-1; 21 VII 2006, Poryadina, SASY (det. as R. archaea); 16 VII 2000, Poryadina, SASY; 4 VII 2015, Chesnokov SC-209, LE L-25143; 10 VII 2015; Chesnokov SC-213, LE L-25144; 6 VII 2015; Chesnokov SC-211, LE L-25145; 4 VII 2015, Chesnokov SC-205, LE L-25146; 6 VII 2015, Chesnokov SC-212, LE L-25147; Neryungrinsky District, 1 VII 2015, Konoreva SC-253, LE L-25148; Fes'ko, SASY L-1987-08-12/525; 11 VII 2015, Poryadina, SASY; 12 VII 2015, Poryadina, SASY.

Rinodina subpariata (Nyl.) Zahlbr.

Rinodina subpariata is characterized by its light gray thallus, whitish labriform soralia in early development, abundant presence of atranorin, and Physcia-type spores (Sheard, 2010) in early development and Physconia-type spores in well developed esorediate morphs (Resl et al., 2016).

Rinodina subpariata might be confused with light gray forms of R. efflorescens Malme, but the latter species has pannarin and its soralia do not contrast with the thallus in colour (Sheard, 2010). Rinodinagriseosoralifera Coppins is also characterized by the presence of atranorin and zeorin but is distinguished by its bluish- or greenish-gray soralia over large areas of the thallus and by its Pachysporaria-type spores (Sheard, 2010). Rinodina subpariata is similar to R. willeyii Sheard et Giralt, but the latter has Pachysporaria-type spores, and an areolate thallus producing pannarin (P+ cinnabar).

Distribution. Eurasia [Scotland, Scandinavia, and Austria (as R. degeliana Coppins)], Mongolia (Khentey Mts.), Russia (from the Caucasus to the Far East), Japan, and Korea], North America (a Pacific and North Atlantic disjunct distribution). Rinodina subpariata was reported in Yakutia as R. degeliana (Vershinina et al., 2012, 2015; Chesnokov et al., 2016).

Ecology. On bark of Picea obovata Ledeb., Salix sp. in different forests at 110582 m a. s. l.

Specimens examined: Zhigansky District, Galanina, VLA L-3154; Aldansky District, 19 VII 2015, Chesnokov SC-216, LE L-25148; 11 VII 2015, Chesnokov SC-232, LE L-25150; Neryungrinsky District, 11 VII 2015, Poryadina, SASY; Chikidov, SASY L-2007-07-01/9-1.

Rinodina terrestris Tomin

Rinodina terrestris is characterized by a thin brownish-gray thallus, small apothe-cia with a persistently plane or rarely convex black disc, thalline margin concolou-rous with thallus, Physcia-Physconia-type spores (17.5)22.5-24.0(29.0) x (8.0)10.0-10.5(12.5) |im of Type A or B development, without torus and often mucronate.

Rinodina terrestris can be confused with R. roscida; for the differences see the description of R. roscida.

Distribution. Rinodina terrestris is scattered throughout the temperate and Arctic Northern Hemisphere, North America (from western intermontane dry areas of British Columbia and USA, occurring southwards into Colorado and New Mexico). Despite R. terrestris has been several times reported for Yakutia (Afonina, 1980, 1979; Poryadina, 2020a, b) it was only once found in the studied collections (27 VI 1976,

Makarova, LE). One specimen (Poryadina, 2020b) was reidentified as R. sibirica. It had atypical "bloated" thick thallus growing on wood along with R. pyrina (1 VI 2019, Poryadina, SASY L-2019-06-01/3-5-6). We have already studied samples of R. sibi-rica with a similar non-typical thallus from Yakutia and they were also seen by John Sheard (Galanina et al, 2021c). Three specimens (not published) (16 VII 2015, Ko-noreva, LE L-13353; 13 VII 2015, Chesnokov, LE L-13333; 13 VII 2015, Konoreva SC-237, LE) were reidentified as R. roscida.

Ecology. On soil near the mouth of river in meadow-steppe. The species is characteristic for dry steppes and desert-like sites.

Specimen examined: Oymyakonsky District, 27 VI 1976, Makarova, LE.

Rinodina turfacea (Wahlenb.) Korb.

Rinodina turfacea is characterized by a brownish-gray thallus, large apothecia with concave or plane discs, persistent thalline margin containing crystals of sphaerophorin, Physcia-type spores (22.0)27.5-29.5(35.0) x (10.5)12.5-13.5(15.5) ^m of Type A development.

Rinodina turfacea can be confused with R. cinereovirens; for the differences see the description of the latter species.

Distribution. It is mainly northern circumpolar species restricted to the Arctic and Subarctic territories. Eurasia (from Scandinavia to the Kamchatka Peninsula with southernmost locations in the Mongolian and Chinese parts of the Altai), North America (Arctic, southwards to the Rocky Mountains in Montana and Wyoming, and Colorado). Rinodina turfacea was previously reported from the Yakutia (Afonina et al., 1979, 1980; Andreev, 1983, 1984; Makarova, Perfiljeva, 1984; Makarova et al, 1988; Makarova, 1989; Samarskii et al, 1997; Poryadina, 2001, 2020a; Zhurbenko et al, 2002, 2005).

Ecology. On mosses and plant debris in different forests and tundras, flowing swamp at 81-1092 m a. s. l.

Specimens examined: Anabarsky District, 1981, Anonymous, SASY; 8 VII 1981, Stepanova, SASY; Bulunsky District, 12 VIII 1982, Nikolin, SASY; Nikolin, SASY L-1982-08-14/20-1; 12 VIII 1982, Nikolin, SASY; 12 VIII 1982, Perfiljieva, SASY; 18 VIII 1982, Perfiljieva, SASY; Poryadina, SASY L-2009-07-24/19-1; 2 VII 1988, Makarova, LE (2 speciments, det. as Rinodinaexigua);Nikolin, SASY L-1982-08-13/16-4; Momsky District, 25 VII 1993, Poryadina, SASY; Vilyuysky District, Galanina, VLA L-2997; Tomponsky District, 14 VII 1996, Poryadina, SASY (det. as R. archaea), Poryadina, SASY L-2016-07-17/19-5; Oymyakonsky District, Yatsyna SC-286, LE L-25151.

Doubtful and excluded species Rinodina archaea (Ach.) Arnold

Rinodina archaea is characterized by its broadly attached and contiguous apothe-cia with disc rather persistently plane, and by its relatively large Physconia-type spores (Sheard, 2010, 2018).

Distribution. Europe and western North America with some outliers in the eastern part, Russia (European part, the Urals, Siberia, and the Far East). Before our study, the species was several times reported for Yakutia (Poryadina, 2001, 2003, 2006, 2020a; Vershinina et al., 2015). We assume that Rinodina archaea can be found in Yakutia, but the available specimens identified as R. archaea (Poryadina, 2001, 2003, 2006, 2020a) were reidentified as R. sibirica (21 VII 2006, Poryadina, SASY; 14 VII 1996, Poryadina, SASY; 5 VII 1996, Poryadina, SASY L-1996-07-05/7-1-2-3; 1 VII 1996, Poryadina, SASY L-1996-07-01/3-1-2-3; 22 IX 2018, Poryadina, SASY L-2018-09-22/8-6), R. turfacea (14 VII 1996, Poryadina, SASY), R. laevigata (14 VII 1996, Poryadina, SASY L-1996-07-14/11-7-8-9), and R. cinereovirens (14 VII 1996, Poryadina, SASY L-1996-07-14/12-7-8). The samples published by Vershinina et al. (2015) were not available for revision. No specimens of I. I. Ma-karova (Makarova, Perfiljeva, 1984; Makarova, 1998) were found in LE. Among the studied specimens from Yakutia, R. archaea was not found. We also did not find R. archaea in the Far East of Russia (Sheard et al., 2017; Galanina, Ezhkin, 2019; Galanina et al., 2021a, b).

Ecology. Rinodina archaea is typically lignicolous but also occurs rarely on rough bark of deciduous and coniferous trees, and even more rarely on siliceous rocks in southern Europe (Mayrhofer, Sheard, 2007; Sheard, 2010).

Rinodina exigua (Ach.) Gray

Rinodina exigua is characterized by its Physcia-type spores, cortical atranorin, indistinct cortex, large areoles, and rugose thallus (Mayrhofer, Moberg, 2002).

Distribution. Europe (the species is common in western and central, scattered in southern areas), North Africa, Australasia, North America (California and the Sierra Nevada), and Russia (temperate zone). Our research has shown that it was misidentified in Russia. We think that this species is absent on the territory of Yakutia, as well as on the territory of the Far East (Sheard et al., 2017; Galanina, Ezhkin, 2019; Galanina et al., 2021a, b). Most of the speciments previously identified as Rinodina exigua and published for Yakutia (Makarova, 1998; Poryadina, 2001, 2003, 2006; Vershinina et al., 2015) have been revised and reidentified by us: two specimens as R. sibirica [15 VIII 2006, Poryadina, SASY; 13 VII 2002, Poryadina, SASY) with R. exiguella (Vainio) H. Magn.], one specimen as R. laevigata (3 IX 1993, Poryadina, SASY L-1993-09-03/1-1-2-3-4), two specimens as R. turfacea (2 VII 1988, Makarova, LE), and one as R. olivaceobrunnea (14 VII 1996, Poryadina, SASY L-1996-07-14/11-10-12). The material published by Vershinina et al. (2015) was not available for this study. Rinodina exigua was found neither among the collections from Yakutia nor from the Far East (Sheard et al., 2017; Galanina, Ezhkin, 2019; Galanina et al, 2021a, b).

Ecology. Rinodina exigua is typically corticolous growing on bark of deciduous and coniferous trees, rarely lignicolous (Mayrhofer, Moberg, 2002; Sheard, 2010).

Rinodina exiguella (Vainio) H. Magn.

Rinodina exiguella is characterized by its gray, thin thallus, dense, small, black apothecia, cellular cortex, I-, and the spores Physcia-type (Magnusson, 1947) and it is cited as a synonym of R. septentrionalis by Sheard (2010).

Distribution. Rinodina exiguella was previously recorded for Yakutia from Yano-Indigirsky Region (Poryadina, 2005). But this specimen (3 IX 1996, Poryadina, whith R. exigua, SASY L-1993-09-03/1-1-2-3-4) (Poryadina, 2005) appeared to belong to R. laevigata. As there are no other references stating this species as growing in Yakutia, we exclude it from the list of the area.

Ecology. Rinodina exiguella is a lignicolous species known from the bank of the Irtysh River (Magnusson, 1947).

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) ^m 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 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, 2013; Ohmura, Kashiwadani, 2018), North Africa (Kotlov, 2008), and North America (Greenland, Rocky Mountains, Sierra Nevada, Cascades and costal range) (Sheard, 2010). 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).

The specimen of Rinodina milvina (7 VII 1996, Poryadina, SASY) (Poryadina, 2001), was identified as Buellia sp. (brown hypothecia, no thallus margin, Buellia-type spores). Rinodina milvina was not found among the studied specimens from Yakutia.

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 sophodes (Ach.) A. Massal.

Rinodina sophodes is characterized by Milvina-type spores 13.0-18.0 x 7.0-9.0 |im with the well-developed torus, and Type A development. Rinodina sophodes has immersed to subimmersed apothecia, often confluent with plane dark brown disc and reddish-brown thallus on a black prothallus (Mayrhofer, Moberg, 2002).

Rinodina sophodes can be often confused with R. archaea, but the latter has Physco-nia-type spores and mainly lignicolous (Sheard, 2010). Specimens of R. sophodes may also be confused with R. sibirica, which differs by Physcia-Physconia-type spores and erumpent apothecia broadly attached at first and then becoming narrow at the base,

scattered, numerous with the dark brown to black disc becoming convex, sometimes hemisphaerical (Sheard, 2010; Galanina et al., 2021c).

Distribution. Rinodina sophodes is common in Europe, scattered in southern and central Scandinavia, rare in Denmark, widespread in the British Isles, less common in Germany, common in southern Europe and Macaronesia. Rinodina sophodes is widely distributed throughout Russia (Arctic, European part, Urals, Siberia, Far East). However, it was previously excluded from the list of species of the Far East (Sheard et al., 2017). As well as in Yakutia, the specimens previously identified as R. sophodes from the Far East appeared to be a mixture of other species. The species was recently excluded from the lichen list of North America (Sheard, 2010) as an old, misapplied name. Rinodina sophodes was reported in Yakutia for the Verkhoyansk Range (Poryadina, 1999b) and for the Sukharnaya River mouth on the East Siberian Sea coast (Andreev, 1984). The specimen (25 VII 1993, Poryadina, SASY L-1993-07-25/11-3-4) was reidentified as R. sibirica. Rinodina sophodes was not found among the studied collection of specimens from Yakutia.

Ecology. Rinodina sophodes is corticolous, on smooth bark, especially on twigs of deciduous trees (Mayrhofer, Moberg, 2002).

Key to the species of Rinodina from the Republic of Sakha (Yakutia)

1. On rock..................................................................................................................................................................2

— On other substrate .............................................................................................................................................6

2. Ascospores Physcia-type ................................................................................................................................... 3

— Ascospores different...........................................................................................................................................4

3. Ascospores averaging 19.0-20.0 |m, Physcia-type, thallus K+ yellow, P+ yellow, atranorin

present in cortex.................................................................................................................R. confragosa

— Ascospores averaging 15.0-17.0 |im, Physcia-Physconia-type, thallus K-, P-, atranorin absent ....

................................................................................................................................................ R. interpolata

4. Ascospores Bischoffii-type.................................................................................................................................5

— Ascospores Dirinaria-type...............................................................................................................R. jacutica

5. Ascospores averaging 16.0-18.0 |im, with unusually broad lumina canals during spore

development, which sometimes become excluded........................................................R. bischoffii

— Ascospores averaging 19.0-20.0 |im, with unusually narrow lumina canals during spore

development..........................................................................................................................R. calcigena

6. On bark or wood, ascospores of various types.......................................................................................... 11

— On decaying plant debris, bryophytes, or soil, ascospores Physcia- or Physcia-Physconia-type .... 7

7. Ascospores averaging 20.5-21.5 |im, Physcia-type...................................................R. olivaceobrunnea

— Ascospores averaging >21.5 |m, of various types......................................................................................8

8. On soil, ascospores averaging 22.5-24.0 |m, Physcia-Physconia-type..............................R. terrestris

— On decaying plant debris or bryophytes, ascospores averaging >24.0 |m, Physcia-type

or Physcia-like...........................................................................................................................................9

9. Ascospores averaging 24.5-25.5 |m, apothecia becoming convex, thalline margin becoming

excluded .................................................................................................................................R. mniaroea

— Ascospores averaging >27.0 |m, apothecia not becoming convex, thalline margin not becoming

excluded .................................................................................................................................................. 10

10. Ascospores averaging 27.5-29.5 |im, Physcia-type, sphaerophorin present....................R. turfacea

— Ascospores averaging 30.0-32.0 |im, Physcia-like (sometimes similar to Dirinaria-type),

sphaerophorin absent..............................................................................................................R. roscida

11. Vegetative propagules present.................................................................................................................... 12

— Vegetative propagules absent ....................................................................................................................... 13

12. Areolae plane, light gray with labriform soralia, P+ yellow........................................... R. subpariata

— Areolae convex to bullate, dark gray to brown, blastidia often present, P+ orange........R. excrescens

13. Ascospores Milvina-type..............................................................................................................R. sophodes

— Ascospores different........................................................................................................................................ 14

14. Ascospores Dirinaria-type........................................................................................................................... 15

— Ascospores different........................................................................................................................................ 17

15. Ascospores averaging <16.5 |im, of Type A or B development, slightly inflated at septum,

thickening is hardly noticeable in water and is better visible in KOH ................................... 16

— Ascospores averaging 17.5-19.0 |m, of Type B development, inflated at septum, thickening is

clearly visible in water, more so in KOH..................................................................... R. metaboliza

16. Ascospores averaging 12.0-14.0 |im, of Type B development, with Physconia-like lumina and

thin walls, apothecia broadly attached, often contiguous with convex disc .............. R. pyrina

— Ascospores averaging 15.0-16.5 |m, of Type A and B development, with Physcia-like lumina,

apothecia erumpent, becoming broadly attached, often scattered with plane or slightly convex disc.................................................................................................................................... R. oleae

17. Ascospores 3-septate or submuriform at maturity................................................................................ 18

— Ascospores 1-septate at maturity ................................................................................................................ 19

18. Ascospores 3-septate, of Type B development ........................................................................R. conradii

— Ascospores submuriform, of Type A development .............................................................. R. intermedia

19. Ascospores Physcia-type.............................................................................................................................. 20

— Ascospores Physconia- or Physcia-Physconia-type.................................................................................. 25

20. Thallus K+ yellow, atranorin present in cortex.........................................................................R. exigua

— Thallus K-, atranorin absent........................................................................................................................ 21

21. Ascospores averaging 23.0-25.5 |m................................................................................. R. cinereovirens

— Ascospores averaging <21.5 |m................................................................................................................... 22

22. Ascospores averaging 20.5-21.5 |m, apothecia with disc becoming convex to half-globose..........

.......................................................................................................................................................R. sibirica

— Ascospores averaging <19.5 |m, disc of apothecia plane to convex................................................... 23

23. Ascospores averaging 18.5-19.5 |m, apotecia with disc plane, sometimes becoming convex ........

...................................................................................................................................................R. laevigata

— Ascospores averaging <17.0 |m................................................................................................................... 24

24. Ascospores averaging 15.0-16.0 |m, thallus gray-green, continuous, apothecia frequently

becoming contiguous on small thalli.......................................................................................R. freyi

— Ascospores averaging 16.0-17.0 |m, thallus copper-brown, consisting of small discrete verrucae

(convex when moist), apothecia narrowly attached and scattered................R. septentrionalis

25. Ascospores Physcia-Physconia type, averaging 16.0-16.5 |im — Ascospores Physconia-type, averaging >18.0 |im........................

R. orculata ............... 26

26. Ascospores averaging 18.0-18.5 |im, apothecia scattered, with the disc frequently becoming convex, and the narrow thalline margin often becoming biatorine or excluded, typically

— Ascospores averaging 19.0-20.0 |m, apothecia broadly attached and contiguous, with the disc

rather persistently plane, with thalline margin to 0.1 mm wide, typically lignicolous .............

..................................................................................................................................................... R. archaea

Discussion

As a result of the study, the list of Rinodina species known from the Republic of Sakha (Yakutia) consists of 24 species, eight of which are reported for the first time for the region (Rinodina cinereovirens, R. confragosa, R. conradii, R. intermedia, R. interpolata, R. metaboliza, R. orculata, R. trevisanii). One more interesting record is Rinodina jacu-tica which has only recently been described from Northeast Asia (Galanina et al, 2022).

In Yakutia, Rinodina is mainly represented by species typically occurring in the oro-arctic (R. mniaroea, R. roscida, and R. turfacea) and north temperate regions (R. cinereovirens, R. metaboliza, R. olivaceobrunnea, R. septentrionalis, and R. sibirica). Other noteworthy species are R. bischoffii, R. calcigena, R. confragosa, R. conradii, R. intermedia, R. interpolata, R. jacutica, R. orculata, R. pyrina, and R. trevisanii which, as yet, have been found only once. Some species are apparently rare because saxicolous species are poorly collected and therefore poorly represented in herbaria. Other species which require further attention are R. laevigata and R. sibirica, both being widespread in Russia. The distribution of R. sibirica was recently presented by Galanina et al. (2022), and it is clear that both species are close to each other in morphology and ecology. They often occur together and have often been confused because of their overlapping in size Physcia-Physco-nia-type spores, numerous scattered apothecia often with a convex dark brown to black disc, and thallus consisting of scattered grayish brown areoles. The distribution of R. lae-vigata and its differences from R. sibirica deserve further study using molecular methods.

One more species that needs to be mentioned is Rinodina excrescens. We previously reported the rediscovery of this species in Russia from new records in the Russian Far East from Sakhalin Island to the Mongolian border (Galanina et al., 2011). Rinodina excrescens was described from western Siberia by Vainio (1928) but since then has been reported as frequent from eastern North America (Great Lakes region), and from south-central and southern Europe, where it is rare (Giralt et al, 1993, 1994; Aragon et al. 2004; Spribille et al, 2006; Giralt, 2010; Sheard, 2010). We reported a major extension to the species' range in northeastern Asia and an apparent second centre of its distribution that had previously been overlooked (Galanina et al, 2011). In Yakutia and Siberia as a whole, we found confirmation of this, because R. excrescens is very often encountered here, as well as R. laevigata and R. sibirica. These species are abundant on the territory of Yakutia.

Five species were not found by us in Yakutia, but were previously reported for the region. These are Rinodina archaea, R. exigua, R. exiguella, R. milvina, and R. sophodes.

corticolous

R. trevisanii

Based on studies of Rinodina in Northeast Asia (Sheard et al., 2017; Galanina, Ezh-kin, 2019; Galanina et al., 2021a, b), we believe that the specimens stored under these names in Russian herbaria need to be verified. Rinodina pyrina was also often incorrectly identified in Yakutia. We reidentified all samples of R. pyrina from Yakutia, with one exception from the central part of the region.

Despite the presented data, the genus Rinodina in the Republic of Sakha (Yakutia) requires further study.

Acknowledgments

We are very grateful to Dr. J. W. Sheard, a monographer of the genus Rinodina in North America, for consultations in the study of Rinodina. The research of I. A. Galanina was carried out within the state assignment of Ministry of Science and Higher Education of the Russian Federation (theme No. 121031000117-9). The research of L. V. Poryadina was carried out within the framework of the state assignment of Ministry of Science and Higher Education of the Russian Federation (theme No. АААА-А21-121012190038-0), with the use of scientific equipment of the Shared core facilities of the Federal Research Center «Yakutsk Science Center SB RAS» within the framework of the implementation of activities under grant № 13.21.0016. The research of S. V. Chesnokov and L. A. Konoreva was carried out within the framework of the institutional research project "Flora and taxonomy of algae, lichens, and bryophytes in Russia and phytogeographically important regions of the world" (theme No. 121021600184-6) of the Komarov Botanical Institute of the Russian Academy of Sciences. Field work of I. A. Galanina (travel from Yakutsk to the middle reaches of the Linde River as part of a field team) in the Linde River basin was funded by the Russian Science Foundation grant No. 21-17-00054, https://rscf.ru/project/21-17-00054/.

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