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

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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. exiguella, 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.
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Received: 5 April 2023 Accepted: 26 October 2023 Published: 7 November 2023
https://doi.org/10.31111/nsnr/2023.57.2.L49
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: Rinodi-
na 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, dierences from closely related species, and distribution in Rus-
sia 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
1Федеральный 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
Novosti sistematiki nizshikh rastenii 57(2): L49–L85. 2023
Резюме. Работа основана на изучении материала, собранного авторами в 1974–2022 гг. в
Республике Саха (Якутия). В результате исследования новый список лишайников рода Rino-
dina 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 близ-
ких 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 dicult 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 Rus-
sia, the genus Rinodina was actively studied in the last decade, but most of these inves-
tigations concern the south of the Far East (Galanina et al., 2011, 2018, 2021a, c; Ga-
lanina, 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 Rino-
dina 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 sum-
marise 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,
Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
L50
1983; Zhurbenko et al., 2005). The species R. archaea, R. bischoi (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. bischoi, 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 (Körb.) 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 conti-
nental, characterized by long winters and short summers. The temperature dierence
between the coldest month (January) and the warmest month (July) is 70–75 de-
grees. 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 tempera-
ture 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 Octo-
ber) — 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 500–
700 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 en-
tire 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 pla-
teaus (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.
Novosti sistematiki nizshikh rastenii 57(2): L49–L85. 2023
L51
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 dierent 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 lab-
oratory of Botany of the Federal Research Center for Biodiversity of the Far East
Branch of Russian Academy of Sciences. The investigation of anatomical and morpho-
logical features of lichens was made using the microscopes Zeiss Axioplan 2 and Stemi
2000-С. The study of spore structures and measurements were made using immersion
oil at 1000× magnification. Anatomical examination used hand-cut sections mounted
in water, and the following reagents: 10% KOH (K) and C6H4(NH2)2 (P). The proto-
col of Meyer and Printzen (2000) was used to characterize the epihymenium pigments.
Measurements of ascospores are presented as percentiles (5)25–75(95) µm length ×
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 (he xane: diethyl
ether: formic acid, 140:72:18), C (toluene: acetic acid, 170:30), and E (cyclohexane:
ethyl ace tate, 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 li-
terature 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; Tønsberg, 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; Aragón et al., 2004; Spribille et al., 2006; Mayrhofer, Sheard,
2007; Giavarini et al., 2009; Wirth et al., 2013), Asia (Vězda, 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., 2021с), 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 et al. The genus Rinodina in Republic of Sakha (Yakutia)
L52
Galanina, 2022), Yakutia (Andreev, 1983; Makarova, Perfiljeva, 1984, 1988; Ma-
karova, 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; Vershi-
nina 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; Gala-
nina, 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 Ame-
rica (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).
Novosti sistematiki nizshikh rastenii 57(2): L49–L85. 2023
L53
Results
In total, 19 species Rinodina were known for the Republic of Sakha (Yakutia) be-
fore the present study. As a result of our research, the list of Rinodina species of Yaku-
tia 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.exi-
gua, 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 Rinodina pyrina by Vershinina et al. (2012, 2015) re-
quires 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, some-
times 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 µm wide), with a cortex expanded to
20–60 µm wide below. Rinodina cinereovirens has Physcia-type spores of Type A deve-
lopment, (21.5)23.0–25.5(27.5) × (10.0)11.5–13.5(14.0) µm and a well developed torus.
Crystals absent in cortex and present in medulla (sphaerophorin). Spot tests all nega-
tive, the secondary metabolite sphaerophorin turns blue-white in longwave UV.
Rinodina cinereovirens is closely related to R. turfacea but diers by more broadly
ellipsoid spores with more bluntly rounded apices. Furthermore, R. cinereovirens in-
habits 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).
Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
L54
D i s t r i b u t i o n . Eurasia [Scandinavia (Norway, Sweden, and Finland), Rus-
sia (from Leningrad Region to Kamchatka Territory and Magadan Region)], North
America (USA: Alaska; Canada: northern Manitoba, northern Ontario, Newfound-
land, New Brunswick).
E c o l o g y . 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; To m -
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 per-
sistent, up to 100 µm wide, with a columnar cortex expanded to 30–50 µm wide be-
low. Rinodina confragosa has Physcia-type spores of Type A development, (16.5)19.0–
20.5(22.5) × (8.0)9.0–10.0(11.0) µm 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 morpho-
logy and spot tests, but the latter possesses Dirinaria-type spores (16.5)18.0–20.0(23.0)
× (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 diers in the presence
of pannarin located only in the epihymenium (Giralt, van den Boom, 1996).
D i s t r i bu ti o n . 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.
E co l o g y . 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 develop-
ment. 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 diers in
submuriform spores with Type A development. Rinodina intermedia can have rose-vi-
olet medullary reaction with KOH (Sheard, 2010).
D i s t ri b ut io n. Widespread in the world in both hemispheres, with a cold tem-
perate to oroarctic distribution, in Russia it is widespread from the Murmansk and
Leningrad regions to the Primorye Territory in the Far East.
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Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
L56
E c o l og y . 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 deve-
lopment. 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.
D i s t r i b u t i o n . 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 Is-
land in Lake Baikal, Buryatia (Volcano valley), Trans-Baikal Territory (Sokhondins-
kiy Nature Reserve), Khabarovsk Territory (Kukanskii Ridge).
E c ol og y. On bark of Larix gmelinii and on a dead tree in larch forests. More in-
formation 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) × (6)7–8(9) µm 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) × (8.0)10.5–11.5(14.0) µm (Sheard, 2010).
D i s t r i b u t i o n . Europe (central part, British Isles, Scandinavia, and Iceland),
Russia (Republic of Karelia, the Caucasus, the Urals, and southern Siberia), North
America.
E co l o g y . 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 — Bischoi-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 µm.
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Rinodina metaboliza Vain.
Rinodina metaboliza is characterized by its Dirinaria-type spores (13.5)17.5–
19.0(23.0) × (8.5)9.0–10.0(11.0) µm 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) × (8.5)9.0–
10.0(11.0)µm (Sheard, 2010).
D i st r i bu t io n . 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).
E co l o g y. On bark of Alnus sp., Betula sp., Picea sp., Populus sp., P. tre mulaL., Sa-
lix 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; To m -
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) × (6.0)8.0–8.5(9.5) µm 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) × (7.5)8.5–9.0(10.5) µm] 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).
D i s tr i b ut i o n. 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).
E co l o g y . On bark of Pinus pumila (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) × (7.5)8.5–9.0(10.5) µm of Type A development (Fig. 3 B, C). The species
Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
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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
dierences 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. ar-
chaea have more prolonged Physcia-type stage of development. Rinodina convexula is
synonymous with R. trevisanii (Mayrhofer, Sheard, 2007; Sheard, 2010).
D is tr ib u t i o n . 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.
E co lo gy . 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 bischoi (Hepp) A. Massal.
Rinodina bischoi is variable in morphology. It is characterized by an inconspi-
cuous, 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
bischoi has spores Bischoi-type of Type A development, the torus is absent, the
walls are not ornamented. The hymenium contains oil drops.
Rinodina bischoi 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 bischoi is similar to R. guzzinii, but the latter has a more
developed thallus and larger spores (21–22 µm length), and the spores of R. bis-
choi (16–18 µm) 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. bischoi. In the Arctic, R. calcigena can be confused with R. bischoi, but the
former has larger, broadly ellipsoid spores (19–20 µm length), and a rapidly disap-
pearing thin canal. A pigmented belt around the septum is visible only in mature
spores (Sheard, 2010).
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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 µm.
D i s t ri b u t io n . 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, Rino-
dina bischoi 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).
E co l o g y . 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 sub-
strate. 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, typi-
cally persistent or rarely disappearing. Rinodina calcigena has Bischoi-type spores of
Type A development (Fig. 2 B), (15.0)19.0–20.0(23.5) × (9.0)11.5–11.5(13.0) µm and
torus absent. The spores of R. calcigena have unusually narrow lumina canals during
Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
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development for the Bischoi-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. bischoi but diers in its longer spores
with unusually narrow lumina canals during development. Rinodina calcigena is
closely related to R. guzzinii but diers 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).
D i s t r i b u t i o n . Eurasia [Europe (Scandinavia), Russia (Polar Urals, Novaya
Zemlya, Southern Siberia, and Kamchatka)], North America (Arctic, Rocky Moun-
tains, Greenland). In Yakutia (Mayrhofer, 1984; Kotlov, 2008).
E co 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 of-
ten 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 con-
tinuous 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 territo-
ries, Sakhalin and Shikotan islands), Japan — Hokkaido Island], North America (the
Great Lakes region). Known from Yakutia (Chesnokov et al., 2016).
E c o l o g y . 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 fre-
quently becoming contiguous on small thalli, and relatively small, darkly pigmented
Physcia-type spores (12.0)15.0–16.0(18.5) × (6.0)7.5–8.0(9.0) µm of Type A develop-
ment, and a heavy torus.
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Rinodina freyi can be confused with R. septentrionalis, but the latter species has
copper-brown thallus consisting of small discrete verrucae (convex when moist), nar-
rowly attached and scattered apothecia.
D i st r ib ut io n. European distribution of this species is poorly known although
originally it was described from Europe (Switzerland) (Magnusson, 1947); the spe-
cies is also published from Germany (Wirth et al., 2013). In Russia, R. freyi was pre-
viously 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).
E c ol o g y . On bark of Alnus sp., Picea sp., Salix sp. in dierent 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 fo rest in
a river valley.
Rinodina jacutica is well distinguished by its Dirinaria-type spores, (16.5)18.0–
20.0(23.0) × (7.5)8.5–10.0(11.5) µm (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 diers in its Pachysporaria-type spores (14.0)15.0–21.0(22.0)
× 8.0–12.0(14.0) µm, 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) µm 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) × (8.0)9.0–10.0(11.0) µm
(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)
× (7.5)8.5–9.0(10.0) µm. It can be distinguished by dark gray to ochraceous, areolate
thallus, lacking prothallus, and negative spot test reactions (Sheard, 2010).
D i s t r i bu t i o n . Russia (Yakutia, Oymyakonsky District) (Galanina et al., 2022).
E co l o g y . On stone in larch forest with mosses-lichen cover at 1179 m a. s. l.
Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
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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 µm. Photo from Galanina et al. (2022).
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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) × (7.0)8.5–9.0(10.5) µm 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 diers by a less developed
thallus, often its darker colour, smaller spores, and thick lower apothecial cortex. Of-
ten morphological and anatomical features of these two species have overlapping sizes,
therefore genetic analysis is necessary for better understanding.
D i s t r i b u t i o n . 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 un-
derstudied. In Yakutia, Rinodina laevigata was found on the Tukulan Mahatta in the
Vilyuysky District (Galanina, 2016a).
E c o l o g y . On bark of Alnus sp., Betula fruticosa Pall., Betula sp., Crataegus sp.,
Duschekia sp., Juniperus sp., Larix gmelinii, Larix sp., Picea sp., Pinus sylvestris, Populus
tremula in dierent 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.) Körb.
Rinodina mniaroea is characterized by its apothecia becoming convex with thal-
line margin becoming excluded, and large Physcia-type spores (20.5)24.5–25.5(30.0)
× (9.5)11.5–12.5(14.5) µm of Type A development.
Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
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Rinodina mniaroea is similar to R. turfacea, which diers 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 diers in its smaller apothecia and spores.
D is t r i b u t i o n . Eurasia [central European part, Iberian Peninsula, and Scandi-
navia, the Caucasus, Himalayas, Mongolia, China, Russia (widespread in the nor thern
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).
E c o l o g y . On mosses and plant debris in the birch and pine forests at 357–
727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) × (6.5)7.5–8.0(9.5) µm 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) × (8.5)9.0–10.0(11.0) µm.
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.
D i st r ib ut io n. 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 distribu-
tion. In Yakutia, Rinodina oleae was found on the Tukulan Mahatta in the Vilyuysky
District (Galanina, 2016a).
E c o l o g y . 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, Galani-
na, 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) × (8.0)9.5–10.0(12.0) µm of Type A development. The species has abun-
dant and small, narrowly attached apothecia.
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Rinodina olivaceobrunnea might be confused with R. turfacea, but diers 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 Vi-
lyuysky District (Tukulan Mahatta) (Samarskii et al., 1997; Galanina, 2016a).
E co lo gy . 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 Dist-
rict, 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 epihy-
menium (Mayrhofer, Moberg, 2002; Sheard, 2010, 2018; Sheard et al., 2011).
D i s t ri b u t i o n. The species is widespread worldwide: Europe, Asia, nor thern
Africa, North America, and Australasia. In Russia, the species is widely distribu-
ted 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 Yaku-
tia (Porya dina, 1999a, 2005; Vershinina et al., 2012, 2015). The specimens (Ver-
shinina 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 co 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 si-
milar to Dirinaria-type), (22.5)30.0–32.0(39.5) × (10.5)12.5–13.5(16.0) µm of Type A or
rarely B development, often with submucronate apices of spores and the lack of a torus.
Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
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Rinodina roscida can only be confused with R. turfacea and dierings 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 mountain-
ous regions of Northern Hemisphere: Arctic (from Scandinavia to Chukotka and from
Alaska to Greenland), Asia (Caucasus, Himalayas, Mongolia, China, Southern Sibe-
ria (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).
E c o l o g y . 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) × (6.5)7.5–8.5(9.5) µm with a well-developed torus and Type A develop-
ment. Rinodina septentrionalis has copper-brown thallus consisting of small discrete
verrucae (convex when moist), and narrowly attached and scattered apothecia.
The dierences from the similar Rinodina freyi see under that species.
D i s t r i b u t i o n . Eurasia (Arctic, boreal zones and mountainous regions from
northern Scandinavia to Сhukotka and Kamchatka, and southbound to Georgia, Ka-
zakhstan, Altai Mts, Sikhote-Alin Mts, Japan); North America (Arctic, extending
southwards in the Rocky Mountains to Colorado). In Yakutia, Rinodina septentrio-
nalis was found in the Suntar-Khayata Ridge and Tukulan Mahatta (Poryadina, 2001;
Galanina, 2016a).
E c o l o g y . On the bark of Alnus sp., Larix sp., Populus tremula, Salix sp., twig
of Crataegus sp. and Picea sp. in dierent forest and moss-ledum-blueberry flowing
swamp at 31–579 m a. s. l.
Specimens examined: Zhigansky District, Galanina, VLA L-3153; Vilyuysky District, Galani-
na, 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) × (8.5)10.0–11.5(13.0) µm with a well-developed torus, the darkly
Novosti sistematiki nizshikh rastenii 57(2): L49–L85. 2023
L67
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.si-
birica 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 diers in its small Milvi-
na-type spores and European distribution (Sheard, 2010).
D i s t ri b u t io n . This species has a wide Amphiberingian range, only slightly ex-
tending to Europe in the Ural Mountains (Galanina et al., 2021с). 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. granu-
lans; Sheard, 2010). Rinodina sibirica was also reported from Yakutia (Kotlov, 2008).
E c o l og y. On the bark of coniferous and deciduous trees in dierent forest types
and flowing swamp at 62–1640 m a. s. l. More information about this species is in Ga-
lanina 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, Poryadi-
na, 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, Porya-
dina, 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,
Ko noreva SC-301, LE L-25140, 17 VII 2016, Konoreva SC-296, 299, LE L-25141, L-25142; Porya-
dina, 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,
Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
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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 Dist-
rict, 1 VII 2015, Konoreva SC-253, LE L-25148; Fes’ko, SASY L-1987-08-12/525; 11 VII 2015, Po-
ryadina, 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. eorescens
Malme, but the latter species has pannarin and its soralia do not contrast with the thal-
lus in colour (Sheard, 2010). Rinodina griseosoralifera 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).
D i s t r i b u t i o n . Eurasia [Scotland, Scandinavia, and Austria (as R. degeliana
Coppins)], Mongolia (Khentey Mts.), Russia (from the Caucasus to the Far East), Ja-
pan, 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).
E c ol o gy . On bark of Picea obovata Ledeb., Salix sp. in dierent forests at 110–
582 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) × (8.0)10.0–
10.5(12.5) µm of Type A or B development, without torus and often mucronate.
Rinodina terrestris can be confused with R. roscida; for the dierences see the de-
scription 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,
Novosti sistematiki nizshikh rastenii 57(2): L49–L85. 2023
L69
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 cha-
racteristic for dry steppes and desert-like sites.
Specimen examined: Oymyakonsky District, 27 VI 1976, Makarova, LE.
Rinodina turfacea (Wahlenb.) Körb.
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) × (10.5)12.5–13.5(15.5) µm of Type A de-
velopment.
Rinodina turfacea can be confused with R. cinereovirens; for the dierences see the
description of the latter species.
D i st ri bu ti on . It is mainly northern circumpolar species restricted to the Arc-
tic 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).
E c o l o g y . On mosses and plant debris in dierent 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 Rinodina exigua); 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).
Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
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D i s t ri b u t i o n. 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. laevi-
gata (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, in-
distinct cortex, large areoles, and rugose thallus (Mayrhofer, Moberg, 2002).
D i s t ri b u t i o n. Europe (the species is common in western and central, scat-
tered 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, Porya dina,
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, Porya dina,
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 col-
lections 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).
Novosti sistematiki nizshikh rastenii 57(2): L49–L85. 2023
L71
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 be-
long 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) × (7.5)9.5–10.5(12.0) µm of Type A development.
Young thalli of R. milvina can be confused with R. parasitica, which is well distin-
guished 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, Ka-
zakhstan, Mongolia, and Japan) (Wagner, Spribille, 2005; Kotlov, 2008; Byazrov, 2013;
Ohmura, Kashiwadani, 2018), North Africa (Kotlov, 2008), and North America (Green-
land, 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 × 7.0–9.0 µm
with the well-developed torus, and Type A development. Rinodina sophodes has im-
mersed 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 diers by Physcia-Physconia-type spores and
erumpent apothecia broadly attached at first and then becoming narrow at the base,
Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
L72
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 com-
mon 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 spe-
cies 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
stu died 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 dierent ........................................................................................................................................... 4
3. Ascospores averaging 19.0–20.0 µm, Physcia-type, thallus K+ yellow, P+ yellow, atranorin
present in cortex ................................................................................................................. R. confragosa
— Ascospores averaging 15.0–17.0 µm, Physcia-Physconia-type, thallus K–, P–, atranorin absent ....
................................................................................................................................................ R. interpolata
4. Ascospores Bischoi-type ................................................................................................................................. 5
— Ascospores Dirinaria-type ............................................................................................................... R. jacutica
5. Ascospores averaging 16.0–18.0 µm, with unusually broad lumina canals during spore
development, which sometimes become excluded ........................................................ R. bischoi
— Ascospores averaging 19.0–20.0 µm, 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 µm, Physcia-type ................................................... R. olivaceobrunnea
— Ascospores averaging >21.5 µm, of various types ...................................................................................... 8
8. On soil, ascospores averaging 22.5–24.0 µm, Physcia-Physconia-type .............................. R. terrestris
— On decaying plant debris or bryophytes, ascospores averaging >24.0 µm, Physcia-type
or Physcia-like ........................................................................................................................................... 9
9. Ascospores averaging 24.5–25.5 µm, apothecia becoming convex, thalline margin becoming
excluded ................................................................................................................................. R. mniaroea
Novosti sistematiki nizshikh rastenii 57(2): L49–L85. 2023
L73
— Ascospores averaging >27.0 µm, apothecia not becoming convex, thalline margin not becoming
excluded .................................................................................................................................................. 10
10. Ascospores averaging 27.5–29.5 µm, Physcia-type, sphaerophorin present .................... R. turfacea
— Ascospores averaging 30.0–32.0 µm, 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 dierent ........................................................................................................................................ 14
14. Ascospores Dirinaria-type ........................................................................................................................... 15
— Ascospores dierent ........................................................................................................................................ 17
15. Ascospores averaging <16.5 µm, 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 µm, 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 µm, 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 µm, 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 µm ................................................................................. R. cinereovirens
— Ascospores averaging <21.5 µm ................................................................................................................... 22
22. Ascospores averaging 20.5–21.5 µm, apothecia with disc becoming convex to half-globose ..........
....................................................................................................................................................... R. sibirica
— Ascospores averaging <19.5 µm, disc of apothecia plane to convex ................................................... 23
23. Ascospores averaging 18.5–19.5 µm, apotecia with disc plane, sometimes becoming convex ........
................................................................................................................................................... R. laevigata
— Ascospores averaging <17.0 µm ................................................................................................................... 24
24. Ascospores averaging 15.0–16.0 µm, thallus gray-green, continuous, apothecia frequently
becoming contiguous on small thalli ....................................................................................... R. freyi
— Ascospores averaging 16.0–17.0 µm, thallus copper-brown, consisting of small discrete verrucae
(convex when moist), apothecia narrowly attached and scattered ................ R. septentrionalis
Galanina et al. The genus Rinodina in Republic of Sakha (Yakutia)
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25. Ascospores Physcia-Physconia type, averaging 16.0–16.5 µm ............................................ R. orculata
— Ascospores Physconia-type, averaging >18.0 µm ..................................................................................... 26
26. Ascospores averaging 18.0–18.5 µm, apothecia scattered, with the disc frequently becoming
convex, and the narrow thalline margin often becoming biatorine or excluded, typically
corticolous ............................................................................................................................. R. trevisanii
— Ascospores averaging 19.0–20.0 µm, 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.cinereo-
virens, R. metaboliza, R. olivaceobrunnea, R. septentrionalis, and R. sibirica). Other note-
worthy species are R. bischoi, 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 fur-
ther attention are R. laevigata and R. sibirica, both being widespread in Russia. The dis-
tribution 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 eco logy. They often occur to-
gether 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 dierences from R. sibirica deserve further study using molecular methods.
One more species that needs to be mentioned is Rinodina excrescens. We pre viously re-
ported 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 exc rescens
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; Aragón 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 con-
firmation of this, because R. excrescens is very often encountered here, as well as R. laevi-
gata 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.
Novosti sistematiki nizshikh rastenii 57(2): L49–L85. 2023
L75
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 incor-
rectly 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 Rino-
dina in North America, for consultations in the study of Rinodina. The research of
I. A. Gala nina was carried out within the state assignment of Ministry of Science and
Higher Education of the Russian Federation (theme No. 121031000117-9). The re-
search of L. V. Poryadina was carried out within the framework of the state assign-
ment 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 frame-
work 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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