The genus Rinodina in the Kuril Islands (Russian Far East)

Summary . The presented work is based on the study of extensive material collected by Ezhkin A. K. in 2014–2018 from the islands of Kunashir, Iturup, Paramushir, and Shikotan. As a result of the study, the new list of species of the genus Rinodina for the Kuril Islands consists of 17 taxa. One species, Rinodina herrei H. Magn., is new to northeast ern Asia and Russia, 14 taxa are new to the Kuril Islands. The genus Rinodina is represented by species with disjunc tive distribution by 35.3 % (6 species). They are divided into two groups: East Asian – West American ( R. herrei, R. megistospora – ( species for 17.6 % (3 species: R. hypobadia, subalbida and xanthophaea of number of Rinodina species of the Kuril Islands


Introduction
As well known, the south of the Russian Far East possesses a large flora of vascular plants and cryptogamic organisms (Kharkevich, 1985;Bulakh et al., 1990;Kozhevnikov, Probatova, 2006;Bogacheva et al., 2018). The genus Rinodina (Ach.) Gray remains unexplored in certain territories of Russia or the data about it is often erroneous. Species of Rinodina are relatively difficult to determine because of the wide variety of types of spores, which are distinguished by their development all stages.
The Russian Far East is one of the territories where the genus Rinodina remained poorly studied. Recently, a monograph of the genus Rinodina was published for North America and north of Mexico . This work facilitated an updating of our understanding of the genus in Northeast Asia (Sheard et al., 2017). This revision included parts of Russia and China, Korea and Japan. Many Rinodina species not previously recognized in the Russian Far East, were discovered. These include some species previously thought to be North American endemics. Close links were found with the Rinodina biota of both eastern and western North America (Sheard et al., 2017;Yakovchenko et al., 2018).
The Kuril Islands are located in the northeastern part of the Pacific Ocean, between Hokkaido (Japan) and the Kamchatka Peninsula (Russia). The climate is temperate, monsoon type with cold winters, but milder than in the mainland part with a cool rainy summers. The southern Kuril Islands (Kunashir, Iturup, Shikotan) are the part of the Sakhalin-Hokkaido province of the East Asian floristic region (Takhtajan, 1978). The flora of northern Kuriles (Paramushir) is enriched with arcticalpine elements including American flora representatives (Barkalov, 2002). The flora of South Kuriles is considered a part of the boreal zone with coniferous and mixed coniferous-deciduous forests but also includes tropical and subtropical elements.
The genus Rinodina in the lichen biota of the Kuril Islands remained unexplored due to its difficult taxonomy. For the Kuril Islands (Kunashir Island), six species of the genus Rinodina were cited (Bredkina et al., 1992;Tchabanenko, 1999;Galanina et al., 2018). These were Rinodina archaea (Ach.) Arnold, R. gennarii Bagl., R. megistospora Sheard et H. Mayrhofer, R. pyrina (Ach.) Arnold, R. sophodes (Ach.) A. Massal., R. xanthophaea Nyl. In our last study (Sheard et al., 2017), it was shown that these species, with the exception of R. gennarii, R. megistospora, and R. xanthophaea, were not found in the Russian Far East. We believe that these species are erroneously listed for the islands. Rinodina gennarii was not found by us, but there is a probability of it being present on coastal rock habitats.
The presented work is based on the study of extensive material collected from different islands (Kunashir, Iturup, Paramushir, and Shikotan). As a result, the list of species for the genus Rinodina in the Kuril Islands includes 17 taxa. One of them is the western North American species -Rinodina herrei H. Magn. -recorded for the first time from Northeast Asia and Russia, and there are additional 14 new records for the Kuril Islands.

Material and methods
Herbarium specimens collected by the second author (A. K. Ezhkin) during 2014-2018 from the Kuril Islands compose the core material for this study. Voucher specimens are deposited in the herbaria SAK and VLA. Morphological and anatomical characters were analyzed by applying standard light microscopical methods. Full label data of examined specimens are provided for every species. The map ( Fig. 1) shows the studied area. A few recent publications have been used to identify samples (Sheard, , 2018Sheard et al., 2017).

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Turczaninowia 22 (4): 5-16 (2019) floristic element. Distributed in the coastal zone of California, Baja California, and British Columbia . The species is first recorded here for Russia and Northeast Asia, on the bark of Quercus crispula Blume in an oak grove on the Iturup Island. This is another species belonging to the group of East Asian-Western North American species.

New species for Kuril Islands
This species grows on the bark of coniferous and deciduous trees, it is found in mixed deciduous forests in Honshu in Japan, at an altitude of 380-1480 m, on Jeju Island in Korea, at an altitude of 750-1600 m, and in Gangwon Province, 380-1660 m,  and Primorye Territory -at low altitudes (Sheard et al., 2017). Rinodina ascociscana was previously considered an endemic species of eastern North America Lendemer et al., 2014). New for the Kuril Islands. The species grows on various deciduous trees in North America (Magnolia, Prosopsis, Prunus, Quercus, Sorbus) and is found on the coastal plain around the Gulf of Mexico, and in the Great Smoky Mountains National Park . It is found in tropics of South America, on the Caribbean Islands, the coast of Portugal, in India (Giralt et al., 1995;Giralt, 2001), in England (Smith et al., 2009. In the latest studies of the genus Rinodina in Northeast Asia (Sheard et al., 2017), the species Rinodina colobinoides was not recorded. It was noted that the species is listed for the south of the Russian Far East by Urbanavichus and Andreev (2010), and based on an earlier reference (Insarov, Pchelkin, 1984;Tchabanenko, 2002) to R. sorediata H. Magn. (as a synonym) on the bark of Quercus mongolica in the Sikhote Alin Mountains (Insarov, Pchelkin, 1984). New for the Kuril Islands.
The species was described from West Siberia by Vainio (1928) and is known from very few localities in Europe where it is scattered and very rare (Giralt et al., 1994;Spribille et al., 2006;Galanina et al., 2011). The species grows in Northern America on the bark of living and dead tree trunks (Betula, Abies, Picea, Larix, Thuja) (Sheard, 1995. Rinodina excrescens was found to be a new record for Japan where it occurs at elevations up to 720- 1030 m in Hokkaido (Sheard et al., 2017). It grows on bark of deciduous and coniferous trees (Betula, Alnus, Quercus, Larix, Pinus, Picea, Salix) in Altai Mts., in Dauria (Sokhondinskiy reserve) and in the Far East of Russia (Galanina et al., 2011;Davydov, Printzen, 2012). New for the Kuril Islands. The species is most often found on branches of bushes and deciduous trees but also on branches of conifers. In North America, it is found on a large number of tree species: Abies, Acer, Alnus, Betula, Fraxinus, Picea, Pinus, Quercus, Sorbus, Ulmus, and others, mainly in the southern boreal zone .

Rinodina freyi
Distribution of the species in Europe is poorly understood, although it was described from Europe (Switzerland) (Magnusson, 1947), and the species is also given for Germany (Wirth et al., 2013).  saw specimens of this species from the Alps and Scandinavia and indicated a confusion with the species R. septentrionalis from Europe (Giralt, Mayrhofer, 1995). Rinodina freyi is most common species of the genus in North America, being frequent in both the east and west of the continent . It has also been reported from western Mongolia (Hauck et al., 2013). Recently the species was found in Japan and Russia (Sheard et al., 2017). In Russia the species is reported for the Magadan Region, Kamchatka Peninsula, Sakhalin Island, Khabarovsk and Primorye Territory (Sheard et al., 2017). New for the Kuril Islands. On bark of deciduous and coniferous trees, often on Sorbus and Acer . On bark of Quercus, Fraxinus in Primorye Territory (Sheard et al., 2017) and on bark of Kalopanax septemlobus on Kunashir Island.

Rinodina hypobadia
The species is known from Southern Europe (Giralt, 2001) and North America, where it has a scattered distribution . As a synonym for R. gennarii Bagl., it is reported for many regions of Russia (Urbanavichus, Andreev, 2010). The close relationship of these species has been discussed in several papers (Giralt, Mayrhofer, 1995;Trinkaus et al., 1999;Giralt, 2001), also using molecular methods in recent works (Helms et al., 2003;Kaschik, 2006). These species are currently treated as separate . Samples on mossy stones in seaside habitats are defined as Rinodina gennarii, while R. oleae occurs on bark and has smaller spores . In East Asia, the species is known from China and Japan (Sheard et al., 2017) and was recently recorded from Korea (Kondratyuk et al., 2013). In the Russian Far East, R. olea was first noted in the Khabarovsk Region (Yakovchenko et al., 2013) and then in Primorye Territory (Sheard et al., 2017). New for the Kuril Islands.  Kotlov, 2008;. Rinodina polyspora was found in central Europe, rarely in southern and central Sweden and in southern Finland (Mayrhofer, Moberg, 2002), in North America in the Great Lakes region and on the Pacific Coast . Rinodina polyspora has recently been reported from Korea (Joshi et al., 2013) and from Japan (Sheard et al., 2017). In Russia, it is found in Karelia, the Leningrad Region, Tatarstan, Western and Southern Siberia (Tyumen Region, Baikal Region, Sayan Mountains) (Kotlov, 2008). New for the Kuril Islands.
The species grows on mosses and plant debris, less often on wood, soil or animal horns, on calcareous substrates (soil) in arctic and highland areas (Mayrhofer, Moberg, 2002;Kotlov, 2008), more rarely on the litter of arctic hare and partridge .
The species is found in central Europe (Wirth et al., 2013), sparsely and rarely in Scandinavia, rarely in southern Europe (Mayrhofer, Moberg, 2002). In North America, it is common in the Arctic, especially in the west, and southwards in the Rocky Mountains . In Russia, it is widely distributed in the northern and mountainous regions: in the Arctic (Franz Josef Land, Novaya Zemlya, Taimyr, Severnaya Zemlya, Yakutia, Chukotka), Murmansk Region, Karelia, Leningrad Region, Arkhangelsk Region, Republic of Komi, Western Siberia (Krasnoyarsk Territory), South Siberia (Transbaikalia, Sayan Mountains), in Asia (Caucasus, Himalayas, Mongolia, China) (Makarova, Katenin, 1983;Kotlov, 2008 The species is found on bark of coniferous and deciduous trees (Abies, Alnus, Berberis, Fagus, Malus, Picea, Prunus, Quercus, Salix, Sorbus), often in mixed and deciduous forests (Sheard et al., 2017). A complete description of the species was first given recently (Sheard et al., 2017).
Rinodina subalbida was described from Japan and was recently found in South Korea and the southern part of far eastern Russia (Sheard et al., 2017). This is an East Asian species, new for the Kuril Islands.  (Sheard et al., 2017), in Russia often in mixed and deciduous forests. It has a wide range of distribution (Sheard et al., 2017). The species is known from eastern Asia with a distribution spanning Russia, Japan and the Korean Peninsula (Sheard et al., 2017). It is a widespread species in the south of the Far East of Russia and in the east of the North America (Great Lakes Region, Appalachian Mountains) . The distribution of Rinodina subminuta is similar to that of R. ascociscana, R. excrescens and R. subparieta (Galanina et al., 2011;Sheard et al., 2017). This is an exclusively East Asian-Eastern North American species, new for the Kuril Islands. The species is found on the bark of coniferous and broad-leaved trees (Abies, Alnus, Betula, Castanea, Chosenia, Picea, Prunus, Quercus, Salix, Taxus), in Russia it is found in deciduous and mixed forests, and same along river valleys. It occurs from the sea level to 2550 m (Sheard et al., 2017). It is known from Scotland (Giavarini et al., 2009), Scandinavia and Austria (Tonsberg, 1992;Mayrhofer, Moberg, 2002), from the Khentey region of Mongolia (Hauck, Javkhlan, 2006) and from eastern and western North America Spribille et al., 2010). The species was previously reported from eastern Asia with a distribution spanning Russia, Japan and Korea (Sheard et al., 2017). It is a widespread species in Russia from the Caucasus to the Far East (Urbanavichus, Andreev, 2010;Himelbrant, Stepanchikova, 2011;Galanina, 2013). New for the Kuril Islands. The species is found on the bark and branches of coniferous trees (Abies, Picea) as well as on mossy horizontal logs (Sheard et al., 2017). In the Kuril Islands it is found in mixed forests on coniferous and broad-leaved species (Alnus, Betula, Padus, Picea Quercus, Salix, Ulmus). The species formerly known from Great Lakes-Appalachian region in North America  and in Japan (Kurokawa, Kashiwadani, 2006). Although widespread in eastern North America, it is relatively infrequent Lendemer et al., 2014, as R. adirondackii H. Magn.). Most recently, Rinodina tenuis was found in Russia, where the species seems to have a narrow distribution in the boreal zone of Khabarovsk Region, Sakhalin Island, and has a couple records from Hokkaido (Japan) (Sheard et al., 2017). We found this species is quite frequent on the southern Kuril Islands (Kunashir and Shikotan), The genus Rinodina in the Kuril Islands (Russian Far East) the closest to Japan. This is an exclusively East Asian-Eastern North American species, new for the Kuril Islands. Sheard  The species was found on the bark and branches of coniferous trees (Abies, Picea) as well as on mossy horizontal logs (Sheard et al., 2017) and in the Kuril Islands it is found in mixed forests on coniferous and broad-leaved species (Alnus, Betula, Padus, Picea, Quercus, Salix, Ulmus).

Rinodina willeyi
This species was described from North America (Sheard, 1995), and before our research in northeastern Asia (Sheard et al., 2017) was known only from North America. The species was noted in Japan (Hokkaido and Honshu islands) and Russia (Kamchatka peninsula) (Sheard et al., 2017). Now the species is found in the southern Kurils for the first time (Kunashir, Shikotan), close to Japan. This record is another addition to the list of Eastern Asia-Eastern North American disjunct species.

Species previously noted for the Kuril Islands
Rinodina gennari Bagl. The species is known from southern Europe mainly on a stony substrate (Giralt, 2001) and from North America frequent on maritime rocks. Not recorded outside coastal regions in North America . Recently Rinodina gennari was found in coastal Japan on mortar and on vertical rock 100 m from the seashore (Sheard et al., 2017). Rinodina gennari has been reported for many regions of Russia as a synonym for the species R. oleae Bagl. (Urbanavichus, Andreev, 2010). These species are treated as separate taxon by . See R. oleae below for more details. Rinodina gennari is reported for the Kuril Islands (Kunashir Island) (Bredkina et al., 1992;Chabanenko, 1999), but we did not find the species on the Kuril Islands although it may well be present. Sheard et  The species grows on bark of Quercus crispula. First described from a Quercus stand at the type locality in southern Oregon, in a relatively high rainfall region close to the coast (Sheard et al., 2011). Recent studies of the genus Rinodina in northeast Asia suggesting that Rinodina megistospora may be widespread but infrequent on the mainland (Sheard et al., 2017). Now the species is recorded also from near coastal localities at low elevations in Hokkaido, Japan and from 1610-1630 m in Kochi Prov., Shikoku Island. The species also occurs in the Sikhote Alinʹ Mountains, Primorye Territory, and in the Khomi Mountains, Khabarovsk Region (Sheard et al., 2017)