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29BRYOPHYTES - МОХООБРАЗНЫЕ
On the genus Oncophorus (Rhabdoweisiaceae, Bryophyta) in Russia
O. M. Afonina1, O. D. Dugarova2, V. E. Fedosov3, 4, D. Ya. Tubanova5
'Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia
2Branch of Federal Budgetary Institution "Roslesozashchita" "Forest Protection Center of the Republic of Buryatia", Ulan-Ude, Russia 3Lomonosov Moscow State University, Moscow, Russia 4Botanical Garden-Institute, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok,
Russia
5Institute of General and Experimental Biology, Siberian Branch of the Russian Academy
of Sciences, Ulan-Ude, Russia Corresponding author. O. M. Afonina, stereodon@yandex.ru
Abstract. Revision of the genus Oncophorus s. str. in Russia based on throughout study of extensive materials kept in LE, MAG, MHA, MW, IRK, KRABG, NSK, PTZ and UUH was carried out. The genus is represented in Russia by two species, O. integerrimus and O. virens. New subspecies, O. virens subsp. minor has been revealed and described based on integrative morphological and molecular (plastid rps4 gene with trnS-rps4 spacer, plastid, trnL-trnFregion and nuclear ITS1-2 region) studies. It differs from O. virens subsp. virens in very dense (vs. loose) tufts, smaller plant size (1-3 cm vs. (2)5-8 cm), smaller leaves [(1.3)1.7-2.1 x (0.4)0.6-0.7 mm vs. 2.4-4.0 x 0.6-1.0 mm)] and smaller mid-leaf cells [(4.3)5.0-9.5(15.0) x (7.6)8.9-11.4(13.0) |m vs. 8-17(20) x 6-10 |m]. Descriptions of O. integerrimus and both subspecies of O. virens together with their photomicrographs, graphic illustrations and a key for their differentiation are provided. Their distribution in Russia is discussed and mapped.
Keywords: Oncophorus integerrimus, Oncophorus virens, Oncophorus virens subsp. minor, distribution, mosses, taxonomy, Russia.
О роде Oncophorus (Rhabdoweisiaceae, Bryophyta) в России О. М. Афонина1, О. Д. Дугарова2, В. Э. Федосов3, 4, Д. Я. Тубанова5
1Ботанический институт им. В. Л. Комарова РАН, Санкт-Петербург, Россия 2Филиал ФБУ «Рослесозащита» Центр защиты леса Республики Бурятия, Улан-Удэ, Россия 3Московский государственный университет, Москва, Россия 4Ботанический сад-институт ДВО РАН, Владивосток, Россия 5Институт общей и экспериментальной биологии СО РАН, Улан-Удэ, Россия Автор для переписки. О. М. Афонина, stereodon@yandex.ru
Резюме. Проведена ревизия гербарных материалов по роду Oncophorus s. str. с территории России, хранящихся в бриологических гербариях LE, MAG, MHA, MW, IRK, KRABG, NSK, PTZ, UUH. В результате интегративного морфологического и молекулярного (пластидные
https://doi.org/10.31111/nsnr/2023.57.1.123 Received. 28 December 2022 Accepted: 14 March 2023
Published. 2 April 2023
маркеры rps4 и trnL-trnF, а также ядерный ITS1-2) изучения выявлен и описан новый подвид Oncophorus virens subsp. minor. Он отличается от O. virens subsp. virens очень плотными и низкими дерновинками, более мелкими размерами растений (1-3 см против (2)5-8 см), более мелкими листьями [(1.3)1.7-2.1 х (0.4)0.6-0.7 мм против 2.4-4.0 х 0.6-1.0 мм] и более мелкими размерами клеток листовой пластинки [(4.3)5.0-9.5(15.0) х (7.6)8.9-11.4(13.0) мкм против 8-17(20) х 6-10 мкм]. На основании полученных данных приводятся иллюстрированные микрофотографиями и графическими рисунками описания и ключ для определения O. integerri-mus и двух подвидов O. virens, обсуждено и закартировано их распространение в России.
Ключевые слова: Oncophorus integerrimus, Oncophorus virens, Oncophorus virens subsp. minor, мхи, распространение, таксономия, Россия.
Until recently, according to the Check-list of East Europe and North Asia (Ig-natov et al., 2006) the genus Oncophorus (Brid.) Brid. in the moss flora of Russia was represented by four species: O. compactus (Bruch et al.) Kindb., O. crispifo-lius (Mitt.) Lindb., O. virens (Hedw.) Brid. and O. wahlenbergii Brid. In 2005, one taxon — O. wahlenbergii var. elongatus I. Hagen was raised by L. Hedenas (2005) to the species level, however, in the Check-list of East Europe and North Asia this species was not included (Ignatov et al., 2006). Recently Hedenas (2017) revised Scandinavian Oncophorus based on molecular and morphological evidence and described a new species O. integerrimus Hedenas, which differs from the generitype (O. virens) in having mainly entire or almost entire upper leaf margins. A little later, Hedenas (2018) introduced one more species of genus Oncophorus, O. demetrii (Re-nauld et Cardot) Hedenas for one of the previously delimited phylogenetic lineages of O. wahlenbergii, which appeared to be distinct morphologically and already described, although in the genus Dicranum Hedw. In the course of our revision of herbarium materials in LE, MAG, MHA, MW, IRK, KRABG, NSK, PTZ, UUH Oncophorus integerrimus and O. demetrii were added to the moss flora of Russia (Sofronova et al., 2018, 2020a, b; Ellis et al, 2019, 2020). However, according to the recently published revision of Rhabdoweisiaceae Limpr. inferred from the molecular phylo-genetic data (Fedosov et al., 2021) the genus Oncophorus was split. Several species, including O. crispifolius and O. elongatus, were transferred to the genus Symbleph-aris Mont. A new genus Brideliella Fedosov et al. was established to accommodate O. wahlenbergii and O. demetrii. The narrow and accepted here concept of the genus Oncophorus includes only two species with gradually narrowed leaves and recurved leaf margins — O. virens and O. integerrimus.
In the course of our revision of the herbarium materials from Russia among the specimens previously identified as Oncophorus integerrimus and O. virens we found several specimens, which differ due to having rather small plants arranged in the very dense tufts. These plants possess entire or almost entire upper leaf margins and loosely twisted leaves that are similar to O. integerrimus and thus appeared problematic for identification. The goal of the present study is to check identity and phylogenetic affinities of these plants and to make the treatment of the genus Oncophorus in Russia.
Materials and Methods
The specimens from LE, MW and UUH were included in the molecular phylo-genetic study. We obtained nucleotide sequence data of plastid rps4 gene with trnS-rps4 spacer and trnL-trnF region, according to the protocol, described by Fedosov et al. (2021). Plastid data was complemented by nuclear ITS1-2 region obtained according to the protocol considered in detail by Gardiner et al. (2005). Nine originally studied ingroup Oncophorus specimens largely originate from not previously sampled area of the North Asia. Specimens studied by Fedosov et al. (2021) and a suite of O. integerri-mus and O. virens accessions from Scandinavia, studied by Hedenäs (2017, 2019) were downloaded from GenBank based on the results of BLAST search; the selection of accessions aimed covering molecular variability of the genus. For the later specimens trnL-F sequences were not available. So, ingroup included 34 accessions. Seventeen outgroup specimens, representing the closely related genera Symblepharis, Brideliella, Glyphomitrium Brid. and Ripariella Fedosov et al. were added as outgroups. Trees were rooted on Glyphomitrium crispifolium Nog. The specimen vouchers and GenBank accession numbers are given in the electronic supplementary materials1.
Sequences were aligned using MAFFT v. 7.487 (Katoh, Standley, 2013) and then edited manually in Bioedit (Hall, 1999). Two datasets used for phylogenetic inferences corresponded plastid data (51 accessions, 1102 positions) and nuclear ITS1-2 (51 accessions, 852 positions). Indel data in all analyses were scored using simple indel coding approach (Simmons, Ochoterena, 2000) using SeqState 1.4.1. (Müller, 2005). Since no supported conflicts of topologies between nuclear and plastid data appeared in the ingroup, we analysed combined dataset (51 accessions, 1954 positions). Phylogenetic analyses were performed using Bayesian Inference and Maximum Likelihood. BI was calculated in MrBayes 3.2.7a (Ronquist et al., 2012) with each run consisted of six Markov chains and 5 million generations; the convergences between runs (accessed as split deviation frequency lower than 0.01) were reached after 0.8 — 1.5 million generations. The sampling frequency was one tree each 1 000 generations, the chain temperature was set at 0.03 in all analyses. GTR model with sampling across the GTR model space (model setting nst = mixed) was used for all-three datasets. Consensus trees were calculated after omitting the first 25% trees as burnin. Convergence of analyses was assessed based on average PSRF values (1.000 in both analyses) and ESS values, checked using Tracer v.1.7.2. (Rambaut et al, 2018) to be higher than 200. Best-scoring maximum likelihood (ML) trees were searched in RAxML 8.2.12 (Stamatakis, 2014) under the GTR model and otherwise default settings on the Cipres Science Gateway (Miller et al, 2010). Robustness of the nodes was assessed using the thorough bootstrapping algorithm with 1000 iterations.
Results and Discussion
The trees inferred from both, plastid and nuclear datasets have subidentical topologies, largely confirming those obtained by Fedosov et al. (2021). However, nuclear
1 Electronic supplement is available at the end of the article page on the journal website (https://doi. org/10.31111/nsnr/2023.57.1.123).
data suggest rather close affinity of the genera Ripariella and Brideliella while sequences of Glyphomitrium are remarkably different, at the same time for plastid data rooting at Ripariella according to the topology by Fedosov et al. (2021) looks appropriate. Due to a rather poor resolution, they are not considered in details. In the tree, inferred from the combined dataset (Fig. 1), accessions of Brideliella form a highly supported clade with those of Ripariella (PP = 1, BS = 100), which largely originates from the phylo-genetic signal of ITS1-2. Accessions of the genus Oncophorus form a highly supported clade (PP = 1, BS = 100), sister to the Symblepharis clade. Within the former, two clades appear, a well supported, fairly diverged (i. e., on a long branch) and little variable one, corresponding to O. integerrimus (PP = 1, BS = 100), and weakly diverged and remarkably variable one, corresponding to O. virens supported (PP = 0.79, BS = 73). Most Asian specimens, which represent typical O. virens based on morphology, appeared in the not statistically supported O. virens subclade (PP = 0.79, BS = 35) together with three specimens from Sweden. This subclade occupies a sister position to the polytomic one, which comprises the rest of O. virens accessions. Three accessions of the small-sized Oncophorus, the present study is intended to deal with, form a clade (PP = 1, BS = 0.71) in this polytomy, thus appearing deep nested within the O. virens clade and no European specimens from GenBank has fallen in it.
The topology of our phylogenetic tree largely corresponds to the previously published ones (Fedosov et al, 2021); however, the phylogenetic position of Brideliella, which formed a basalmost lineage in the R2 clade as delimited by Fedosov et al. (2021) but appeared in well supported clade in Brideliella, in our ITS1-2 based tree seems problematic. This phenomenon requires further studies with the involvement of coding nuclear markers. Our results prove that the small Oncophorus-like plants with entire leaf margins forming dense tufts actually represent one of the infraspecific lineages of O. virens. Molecular variation within O. virens was considered in detail by Hedenas (2017). According to the haplotype network, presented in that study, four molecular lineages might be delimited within O. virens in Scandinavia, although only two are considered in detail. The lineage, revealed here to comprise morphologically distinct Asian plants is no one of those four. The topology of the phylogenetic tree does not suggest considering it as a separate species, thus in the taxonomic species this newly revealed taxon is introduced as a new subspecies.
All three specimens of Oncophorus virens s. str. from the south part of Asian Russia fell in a clade with three Swedish accessions, which formed a separate group in the haplotype network, published by Hedenas (2017); apparently in Asia this group, is more common than in Europe. One more Asian typical O. virens specimen, RF110 from Novaya Zemlya Archipelago appeared in the other isolated and rather weakly represented in the Scandinavia haplotype group, which occurs in Norway and North Sweden. None of two the most common in Scandinavia haplotypes was proved to occur in Asian Russia. This data may indicate the presence of broad scale geographical patterns in the Eurasian distribution of O. virens haplotypes.
Glyphomitrium crispifolium isolate OK707 Primorsky Territory Glyphomitrium crispifolium isolate OK705 Primorsky Territory - Glyphomitrium crispifolium isolate QK706 Primorsky Territory
00j—
_1/100|— Riparieiia riparia isolate 1666 Norvay
^ Riparieiia riparia isolate 291 Yamal ■ Brideliella wahlenbergii isolate 417 Kamchatka
0-94/92!
EF
IÍ92L в i
iP—
Brideliella wahlenbergii isolate 419 USA
Brideliella demetri isolate 493 Buryatia
Ol— йУт ~ll/100j^Si
■ Brideliella demetri isolate 414 Buryatia Brideliella demetri isolate RF34 Orulgan
Symblepharis elongata isolate RF24 Rybachij Peninsula • Symblepharis elongata isolate RF25 Dagestan 0.86/73|_j~~ Symblepharis elongata isolate P255 Sweden
• Symblepharis elongata isolate RF19 Sweden 1/1001- Symblepharis vaginata isolate RF127 Transbaikalia
• Symblepharis vaginata isolate RF150 USA Symblepharis vaginata isolate RF151 USA - Oncophorus integerrimus isolate 420 Kamchatka
• Oncophorus integerrimus isolate 488 Caucasus
• Oncophorus integerrimus isolate MSOv Europe 1/100|- Oncophorus integerrimus isolate On1561 Austria
Oncophorus integerrimus isolate 484 Kamchatka Oncophorus integerrimus isolate P275 Sweden Oncophorus integerrimus isolate P304 Sweden Oncophorus integerrimus isolate P303 Sweden Oncophorus virens isolate 482 Sakhalin Oncophorus virens isolate 489 Zabaikalsky Territory
- Oncophorus virens isolate 415 Buryatia
- Oncophorus virens isolate P284 Sweden
►j- Oncophorus virens isolate P295 Sweden L Oncophorus virens isolate P281 Sweden
Oncophorus virens isolate RF110 Novaya Zemlya Oncophorus virens isolate On1556 Sweden Oncophorus virens isolate P265 Sweden Oncophorus virens isolate P262 Sweden Oncophorus virens isolate P540 Sweden Oncophorus virens isolate P536 Sweden Oncophorus virens isolate P524 Sweden
0.99/62 1/69
0.02
- Oncophorus virens ssp. minor isolate 490 Kamchatka
— Oncophorus virens ssp. minor isolate 487 Franz Jos. Land Oncophorus^ virens ssp. minor isolate 485 Kamchatka_
— Oncophorus virens isolate P301 Sweden * Oncophorus virens isolate P305 Sweden
■ Oncophorus virens isolate P294 Sweden
■ Oncophorus virens isolate P285 Sweden
■ Oncophorus virens isolate P264 Sweden Oncophorus virens isolate P286 Sweden
Oncophorus virens isolate P282 Sweden Oncophorus virens isolate P280 Sweden Oncophorus virens isolate P279 Sweden Oncophorus virens isolate P278 Sweden
Fig. 1. Bayesian phylogenetic tree of Oncophorus and closely related genera inferred from the combined nuclear and plastid data with indels coded using single indel coding approach. Posterior probabilities / Bootstrap support values higher than 60 are shown above the branches. Dashed lines indicated the target taxon.
Taxonomy
1/
Genus Oncophorus Brid., Bryol. Univ.: 389, 1826. Type: Oncophorus virens (Hedw.) Brid.
Key to identification of Oncophorus in Russia 1. Plants small, in very dense tufts, leaves up to 2 mm long, upper lamina cells transverse-rectangular
to quadrate...........................................................................................................O. virens subsp. minor
— Plants lager, usually in loose tufts, leaves 2-4 mm long, upper lamina cells quadrate to short rectangular .....................................................................................................................................................2
2. Leaf margins in acumen entire or indistinctly and obtusely denticulate; acumens of dry leaves
weakly twisted................................................................................................................ O. integerrimus
— Leaf margins in acumen to mid-leaf dentate or coarsely denticulate with sharp and often some double teeth; acumens of dry leaves strongly twisted......................................................O. virens
Oncophorus virens (Hedw.) Brid., 1826, Bryol. Univ. 1: 399. subsp. virens = Dicranum virens Hedw., 1801, Sp. Musc. Frond. 142. = Cynodontium virens (Hedw.) Schimp., 1956, Coroll. Bryol. Eur. 12.
= Dicranum virens var. serratum Bruch et Schimp., 1847, Bryol. Eur. 1: 119. = Cynodontium virens var. serratum (Bruch et Schimp.) Schimp., 1856, Coroll. Bryol. Eur. 12. = Oncophorus virens var. serratus (Bruch et Schimp.) Braithw., 1883, Brit. Moss. Fl. 1: 116. (Figs. 2, 3, 6D-F, 7A, B)
Plants 2-5(8) cm high, dark green, yellow-green or brownish, sometimes almost black below, in loose tufts. Stems erect, weakly branching, with central strand; cortex of 1-3 layers of incrassate cells, hyalodermis locally hardly noticeable; rhizoids red-brown, smooth, branched, in leaf axils. Leaves 2.4-5.0 x 0.6-1.0 mm, when moist erect spreading to spreading, straight or screwed, when dry above sheathing loosely incurved, above strongly twisted, from ovate or rounded-oblong sheathing base tapering to narrowly triangular acumen, apex longly acuminate; margins broadly recurved in the middle part of leaves, below entire, upper margins in all or many leaves regularly to irregularly dentate or coarsely denticulate; teeth single or double; lamina unistra-tose, rarely bistratose, on margins always bistratose; costa strong, 75-110 jm wide near base, ending in leaf apex or shortly excurrent, in cross-section with dorsal and ventral epidermis, one layer of guide cells and 1-3 layers of ventral and dorsal stereids; lamina cells in acumen incrassate, subquadrate, rectangular, rarely transverse-rectangular, 6.5-15.3 x 6.8-12 jm, in mid-leaf incrassate, subquadrate, rectangular, 8.5-17 x 6.5-11.9 jm, and in sheathing lamina less incrassate, rectangular, longer and slightly porose, 20.5-85 x 6.5-17 jm; transition between mid-leaf and basal cells relatively sudden; alar cells rectangular, very weakly differentiated, decurrent. Monoicous. Spo-rophytes frequent, 1 (rare 2 from the same perichaetium); perigonia lateral on stem, perigonial leaves with acute apex; inner perichaetial leaves up to 6.0 mm long, from high vaginal base suddenly narrowed to long, narrow acumen. Seta long, 10-20 mm, yellow, becoming brown with age; capsule cylindric, 1.5-2.0 x0.5-0.8 mm, curved, strumose, furrowed when dry; peristome ruby red and vertically pitted-striolate, papillose, teeth united in short basal membrane, bifid at apex; spores 20-38 jm in diam.
Distinctions. Oncophorus virens subsp. virens is close to O. integerrimus and distinguished by dentate or denticulate upper leaf margins, whereas O. integerrimus has predominantly entire or almost entire leaf margins; also more weakly twisted acumens of dry leaves vs. more strongly twisted in O. virens; shorter cells in the upper part of leaf (subquadrate, transversely rectangular to short rectangular) vs. more elongated, short rectangular. Oncophorus virens subsp. virens differs from subsp. minor in size of plants.
Fig. 2. Oncophorus virens subsp. virens (Buryatia, 11 VII 2002, Tubanova 6(IX), UUH). A — basal leaf cells; B, C, D — leaf transverse section; E, F — leaves; G — upper leaf cells; H — habit, dry; I — leaf tip. Scale bars: 10 mm for H; 1 mm for E and F; 100 jm for A, B and C, D, G, I.
The latter is smallest among arctic Oncophorus and have short vegetative and loosely twisted leaves, and quadrate or short transverse-rectangular cells in the upper part.
Distribution. The revision of herbarium collections (LE, MW, MHA, KRABG, IRK, UUH and partly MAG) revealed that the ranges of Oncophorus virens subsp. virens and O. integerrimus largely overlap. Both taxa are rather widespread in the Arctic and grow in similar habitats. However, O. virens has a wider distribution in the southern regions of East Siberia, especially in the Republic of Buryatia and Trans-Baikal Territory, also in the Republic of Altai, Republic of Sakha (Yakutia); whereas O. integerrimus is much less common there, and is more common in the northern regions, including the Arctic. In the Far East O. virens is distributed from Chukotka to the south of the Pri-morye Territory. In European part of Russia its single finds are known from Franz Josef Land Archipelago, Novaya Zemlya, Murmansk Region, Republic of Karelia, Komi Republic, Perm Region; in West Siberia known in the Yamalo-Nenets Autonomous Area; in the Caucasus it was collected in Karachaevo-Cherkessian Republic (Fig. 3).
Ecology. Oncophorus virens subsp. virens often is dominant of vegetation on pebbles, also grows in polygonal and wet moss tundra, on boulder near water along streams, on wet rocks, and in floodplain willow thickets.
Fig. 3. Distribution of Oncophorus virens subsp. virens in Russia.
Selected specimens examined. EUROPEAN RUSSIA: Arkhangelsk Region: Franz Josef Land Archipelago, Scott Keltie Island, V. P. Savicz (В. П. Саввич) 885, LE B0022161; Alger Island, S. S. Kholod (С. С. Холод), LE B0022160; Novaya Zemlya, Zhelaniya Cape, 1 IX 2015, S. S. Kholod (С. С. Холод), LE; Murmansk Region: Lapponia tulomensis, Kola, V. F. Bro-therus, LE B0022165; (Regio kuusamoensis) Kuolajärvi, Kutsanjoki, Edvard af Hallström, H, LE B0022163; Chil'tak Massiv, Chil' Creek, O. A. Belkina (О. А. Белкина) 7038, KPABG; Kolvitskaya Gulf, Kolvitsa Settlemen, O. A. Belkina (О. А. Белкина) 10203, KPABG; Northeast of Kola Peninsula, Lumbovskii Gulf, Zapadnaya River, O. A. Belkina (О. А. Белкина) 19913, KPABG; Lumbovskii Gulf, left bank of Kamenka River, O. A. Belkina (О. А. Белкина) 20010, KPABG; Republic of Karelia: Rugorskaya Bay, Kindo Island, V. R. Filin (В. Р. Филин), MW 905058; Komi Republic: Pechoro-Ilychskii Reserve, A. A. Korchagin, V. S. Bakhtin (А. А. Корчагин, В. С. Бахтин), LE B0022159; Caucasus, Karachayevo-Circassian Republic: Teberda Reserve, M. S. Igna-tov, E. A. Ignatova, MW 9050566; canyon Uchkulan-ichi, V. G. Onipchenko, MW 9050576; ASIAN RUSSIA, West Siberia: Yamal-Nenets Autonomous Area: Yamal Peninsula, Eryakha River, E. N. Andrejeva (Е. Н. Андреева), LE B0022178; Junto Lake, I. V. Czernyadjeva (И. В. Чернядь-ева), LE B0022179; Gydan Peninsula, Khonorasale Cape, O. V. Rebristaya (О. В. Ребристая), LE B0022168; Khanty-Mansi Autonomous Area — Yugra: Berezovskii District, Pol'ya River, I. D. Kil'dyushevskii (И. Д. Кильдюшевский), LE B0022172; Altai Territory: Between Bartuldag and Kair rivers, P. N. Krylov (П. Н. Крылов), LE B0022191; mouth of Bogodai River, V. I. Vereschagin (В. И. Верещагин), LE B0022190; Republic of Altai: Teletskoe Lake, V. I. Vereschagin (В. И. Верещагин), LE B0022173; East Siberia: Krasnoyarsk Territory: Taimyr Dolgano-Nenetsk District: middle course of Novaya River, urochishe "Ary-Mas", B. N. Norin (Б. Н. Норин), LE B0022169; Puto-rana Plateau: Talnakh Settlement, I. V. Czernyadjeva (И. В. Чернядьева), LE B0022177; Ayan Lake, I. V. zernyadjeva (И. В. Чернядьева), LE B0022185; Evenki District, Central Siberian Reserve, Stol-bovaya River, S. S. Scherbina (С. С. Щербина), MW 9050615; Republic of Sakha (Yakutia): Novo-sibirskie Islands, Stolbovoi Island, I. V. Czernyadjeva, LE B0024324; Medvezh'i Ostrova Archipelago, Chetyrekhstolbovoi Island, T. M. Zaslavskaya (Т. М. Заславская), LE B0022176; low course of Lena River, Tiksi Bay, I. D. Kil'dyushevskii (И. Д. Кильдюшевский), LE B0022194; Tompo District,
Verkhoyanskie Mts., Tompo River basin, Sukhikh, V. B. Kuvaev, LE B0024325; Suntar-Khayata Range, Mus-Khaya Mt., M. S. Ignatov, E. A. Ignatova 11-3219, MW 9050630; Oimyakon District, M. S. Ignatov, E. A. Ignatova, MHA 9028724; Ust'-Aldansk District, M. S. Ignatov, E. A. Ignatova, MHA 9025446; Ust'-Maisk District, M. S. Ignatov, MHA 9018476; Kobyaisky District, Undyulyung River, E. V. Akimova (Е. В. Акимова), MW 9050628; Irkutsk Region: Baikal Lake, Bol'shye Koty, J. Vana, LE B0022193; Shore of Baikal, Murino station, Smirnov (Смирнов) 740, LE; East Sayan, Uda river basin, Belaya Durgomzha River, L. V. Bardunov (Л. В. Бардунов), LE B0022199; Republic of Buryatia: Dzhidinsky District, Malyi Khamar-Daban Range, Barun-Sikhokhta, S. G. Kazanovsky, LE B0022202; Baikal'skii Reserve, Temnik River, S. G. Kazanovsky, LE B0022201; Oka District, Oka River: Zhambolok Creek, Afonina, LE B0022241; ibid., Sorok River, Afonina, LE B0022227; Bargu-sinskii Reserve, Shumilikha River, I. V. Czernyadjeva, LE B0022195; Kurbinsky Range, Angirskii Sanctuary, I. V. Czernyadjeva, LE B0022196; Mukhorshibirskii District, Bolshoi Sibildui River, Afonina, LE B0022253; Kurumkan District, Dzherginskii Reserve, Tubanova (Тубанова) 1(V), UUH; Barguzin Ridge, L. Dorzhieva, Tubanova (Л. Доржиева, Тубанова) Ва160614, UUH; Trans-Baikal Territory: Kyra District, Sokhondinskii Reserve: Bukukun River, Afonina, LE B0022244; Agutsa River, I. V. Czernyadjeva, LE B0022238; Enda River, Afonina, LE B0022239; Ingoda River, I. V. Czernyadjeva 43-13, LE B0022210, ibid., Afonina, LE B0022229; Aleksandrovskii Zavod District, Malaya Borzya River, Afonina, LE B0022204; Dul'durga District, National Park 'Alkhanai", Afonina, LE B0022242; Gazimuro-Zavodskii District, Pryamoii Muldai River, Afonina, LE B0022206; Kras-nochikoii District, Atsinskii Sanctuary, Yugal River, I. V. Czernyadjeva, LE B0022215; Khilok District, Engorok River, Afonina, LE B0022252; Kalar District: Udokan Range, Naminga Settlement, V. R. Filin (В. Р. Филин), LE B0022254; Kodar Range, Schan'go River, Afonina, LE B0022249; ibid, Srednii Sakukan River, Afonina, LE B0022251; ibid., Syul'ban River, Khadatkan Creek, Afonina, LE B0022250; Chukotka Autonomus Area: Iultin District: Wrangel Island, Somnitel'naya Bay, Afonina, LE B0023536; ibid., Rodzhers Bay, B. N. Gorodkov (Б. Н. Городков), LE B0023539; Getlya-nen River, Afonina, LE B0022257; Chukotka District: Chegitun River, Afonina, LE B0022261; Lorino Hot Springs, Afonina, LE B0022256; Lavrentia Gulf, Lavrentia Settlement, Afonina, LE B0023532; Lavrentia Gulf, Krauze Cape, Afonina, LE B0022276; Provideniya District: Provide-niya Bay, Ureliki Settlement, Afonina, LE B0023527; Chaplin Hot Springs, Afonina, LE B0022263; Gil'mimli Hot Springs, Afonina, LE B0022262; Nunligran Settlement, Afonina, LE B0023524; Achen Lake, Afonina, LE B0023529; Yanrakynnot Settlement, Afonina, LE B0022271; Egvekinot Settlement, Afonina, LE B0022258; Anadyr District: Pekul'nei Range, Yuzhnyi Pekul'neiveem River, Afonina, LE B0023533; Bezymyannoe Lake, Afonina, LE B0023535; Pekul'neii Range, Baran'e Lake, Afonina, LE B0022267; Velikaya River, Tamvatnei Creek, Afonina, LE B0022278; between Pekul'neyskoe and Yanrakoim lakes, E. Yu. Kuzmina, LE B0022272; Pekul'neiskoe Lake, Kautayam River, E. Yu. Kuzmina, LE B0023534; Bilibino District: Anyui Plateau, Bilibino Settlement, Afonina, LE B0022264; Magadan Region: Ola District: Oksa, L. S. Blagodatskikh (Л. С. Благодатских), MAG, LE B0023545; Luzhin Bay, L. S. Blagodatskikh (Л. С. Благодатских), LE B0023549; P'ya-gina Peninsula, L. S. Blagodatskikh (Л. С. Благодатских) 1207, MAG; Magadan District, Chernyi Klyuch, L. S. Blagodatskikh (Л. С. Благодатских), LE B0023544; Geptner Bay, vicinity of Magadan, L. S. Blagodatskikh (Л. С. Благодатских) 1205, MAG; Kamchatka Territory: Kamchatka Peninsula, Ushakovskii volcano "Klyuchevskoii dol", I. V. Czernyadjeva, LE B0023548; Sredinii Range, Esso Settlement, I. V. Czernyadjeva, LE B0023542; Kommander Islands, Bering Island, vicinity of Polovina Bay, 13 VIII 2010, Fedosov, MW; ibid, Fedosov 10-3-101, MW; Amur Region: Bureya River, V. S. Docturowskii (В. С. Доктуровский), LE B0023551; Zeisk Reserve, Tukuringra Range, S. V. Dudov, K. V. Kotel'nikova, MW 9050633; Khabarovsk Territory: Sovetskii District, upper course of Tutto River, Kolesnikov (Колесников), LE B0023543; Primorye Territory: Chuguevsk District, M. S. Ignatov, MHA 9018496; Lazovsk District, M. S. Ignatov, E. A. Ignatova, MHA 9018497; Shko-tov District, M. S. Ignatov, E. A. Ignatova, MHA 9018495; Sakhalin Region: Sakhalin Island, Uanga River, Vagis Mt., Tubanova, Tumurova (Тубанова, Тумурова) S14009/19, UUH.
Oncophorus virens subsp. minor Tubanova, Fedosov et Afonina, subsp. nov.
(Figs. 4, 5, 6G-I, 7F-K, 8)
Type: Russia, Arkhangelsk Region: Franz Josef Land Archipelago, Northbrook Island, 79°56'52.3"N, 50°06'08.9"E, sea terrace (18 m a. s. l.), plant community with Alopecurus alpinus subsp. borealis, on wet soil, 24 VIII 2012, S. S. Kholod (С. С. Холод) 74, (UUH — holotype, LE B0023679 — isotype).
Diagnosis: Oncophorus virens subsp. minor resembles O. virens subsp. virens, but differs in having very dense tufts vs. loose tufts, shorter leaves, up to 2 mm vs. 2.4-4 mm long, distal lamina cells shorter, transverse-rectangular vs. subquadrate, short rectangular; leaf margins entire or obtusely denticulate vs. regularly dentate and coarsely denticulate.
Etymology: The species name originates from its small size.
Description. Plants small, up to 1-3(5) cm high, usually in dense and compact tufts, green or dark-green with alive green parts of plants ca. 1 cm and lower dead brown very dense tuft, slightly glossy. Stems erect, weakly branching, with central strand, cortex of 1-2(3) layers of incrassate cells; moderately tomentose, rhizoids strongly branched, yellowish-brown, smooth, in leaf axils, extend to the upper part of the stem. Leaves ± dense, appressed, (1.3)1.7-2.15(2.4) x (0.4)0.6-0.7(0.9) mm, from sheathing base erect-patent, straight, when dry curved or slightly curled, from ovate-triangular or rounded-oblong base gradually tapered into rather long and narrowly triangular acumen, apex shortly acuminate; leaf margins distinctly recurved in the lower leaves, below entire, in the upper part weakly obtusely denticulate or entire, margins varying from unistratose to bistratose; costa strong, ending few cells below apex to in leaf apex, 75-100 |im wide near base, in cross section with a row of guide cells, dorsal and ventral epidermises, two stereid bands (1-2 layers of ventral stereids and 3-4 layers of dorsal stereids); lamina unistratose, rarely bistratose in margins, smooth; lamina cells in the upper part to the middle incrassate, transverse-rectangular, subquadrate, rarely short-rectangular (4.3)5.0-9.5(15.0) x (7.6)8.9-11.4(13.0) jm, in sheathing lamina rectangular, slightly incrassate and not porose (27.8)35.0-55.0(70.0) x (4.6)6.2-8.3(10.2) jim, transition between median and basal cells quite abrupt to relatively sudden, rare gradual; alar cells rectangular, not differentiated or slightly inflated and forming small delimited group of 1-4 cells wide and 2-6 cells long, slightly decur-rent. Monoecious. Sporophytes rare; perigonia lateral on stem, perigonial leaves with slightly blunt to acute apex; inner perichaetial leaves 1.5-1.8 mm long, from high vaginal base suddenly narrowed to short and slightly blunt acumen. Seta 8-12 mm, yellow, becoming light brownish with age; capsule cylindric, 0.9-1.1 x 0.3-0.4 mm, curved, strumose, smooth when dry; peristome red in the larger lower part (ca. 0.7 of the length) and yellow in the upper part, ca. 350 |im, vertically pitted-striolate, weakly papillose, teeth united in the short basal membrane, bifid at apex; spores 20-30 |im in diam., rough.
Fig. 4. Oncophorus virens subsp. minor (24 VIII 2012, S. S. Kholod (С. С. Холод) 74,
UUH — holotype).
A, F — upper leaf cells; B — leaf tip; C, E — leaves; D — habit, dry; G, H, I — leaf transverse section;
J — stem transverse section; K — habit, weight; L — basal leaf cells.
Scale bars: A, B, F-J, L — 100 |m; C, E, K — 1 mm; D — 5 mm.
Distinctions. Oncophorus virens subsp. minor differs from O. virens subsp. virens by very dense and low tufts, small leaves, transversely rectangular lamina cells in acumen, entire leaf margins, loosely twisted leaves. Oncophorus virens subsp. minor is similar to O. integerrimus in leaf margins and slightly twisted leaves but differs in very dense and low tufts, small leaves, predominance of transversely rectangular lamina cells in acumen and the absence of well differentiated alar cells (Figs. 6, 7). Molecular phylogenetic analysis confirmed its similarity to O. virens. Oncophorus virens subsp. minor without sporophytes can be confused with Oreas martiana (Hoppe et Hornsch.) Brid. but differs in short, transversely rectangular cells in acumen while Oreas martiana has elongate cells in acumen. Subglobose, estrumose capsules readily differ Oreas martiana from Oncophorus species, which possess arcuate, distinctly strumose capsules.
Distribution. Oncophorus virens subsp. minor is currently known from European Arctic (Franz Josef Land Archipelago: Northbrook and Hooker Islands), Asiatic
Fig. 5. Oncophorus virens subsp. minor (Kamchatka Peninsula, 23 VIII 2005, I. V. Czernyadjeva s. n, LE). A — habit, wet; B — capsule; C — inner perichaetial leaf; D — perigonial leaf; E, F — basal leaf cells.
A-D — 1 mm; E, F — 100 |m.
.7.
Fig. 6. Tufts and leaf cells of Oncophorus. Oncophorus integerrimus (Karachaevo-Cherkessia, 3 IX 2005, M. S. Ignatov, E. A. Ignatova 159, MHA: A, B, C); O. virens subsp. virens (Buryatia, 11 VII 2002, Tubanova 6(IX), UUH: D, E, F), O. virens subsp. minor (holotype: G, H, I). A, D, G — tufts (scale bar: 1 cm); B, E, H — upper lamina cells (scale bar: 100 |im); C, F, I — basal lamina cells (scale bar: 100 |im).
Fig. 7. Leaves of Oncophorus. A — O. virens subsp. virens (Sakhalin, 17 VII 2014, Tubanova, O. Tumurova S14009, UUH); B — O. virens subsp. virens (Trans-Baikal Territory, Ingoda River, I. V. Czernyadjeva 43-13, LE B0022210); C — O. integerrimus (Kamchatka, 13 VIII 2003, I. V. Czernyadjeva 96, LE); D — O. integerrimus (Kamchatka, 21 VII 2003, I. V. Czernyadjeva 49(1), LE); E — O. integerrimus (Karachaevo-Cherkessia, 3 IX 2005, M. S. Ignatov, E. A. Ignatova 159, MHA); F — O. virens subsp. minor (Kamchatka, 23 VIII 2005 I. V. Czernyadjeva s. n., LE); G — O. virens subsp. minor (Franz Josef Land Archipelago, V. P. Savicz 1488, LE B0023657); H — O. virens subsp. minor (Putorana Plateau, I. V. Czernyadjeva s. n., LE B0023668); I — O. virens subsp. minor (Franz Josef Land Archipelago, V. P. Savicz 1421, LE B0023658); J — O. virens subsp. minor (Polar Urals, 11 VII 1988, I. V. Czernyadjeva 34, LE); K — O. virens subsp. minor (Chukotka, Afonina s. n., LE B0023669).
Scale bars: 1 mm.
Arctic (Putorana Plateau, Wrangel Island, Anadyr River Basin), also in Khasyn District of the Magadan Region and in the Kamchatka Peninsula (Fig. 8).
Ecology. Oncophorus virens subsp. minor grows in goltsy (alpine tundra) on wet clay; in moss community on rocks; on wet fine earth in community with Alopecurus al-pinus subsp. borealis; in herb-moss-dwarf-shrub, sedge-moss, dwarf shrub-lichen- and herb-dominated spotty tundras, in nival meadows along streams.
Specimens examined. RUSSIA, European Arctic: Arkhangelsk Region, Franz Josef Land Archipelago, Hooker Island, Cape Sedov, V. P. Savicz (B. n. CaBuu) 1421, LE B0023658; ibid., V. P. Savicz (B. n. CaBuu) 1488, LE B0023657; Asian Arctic: Yamal-Nenets Autonomous Area, Polar Urals, Middle Sob' River, 11 VII 1988, I. V. Czernyadjeva 34, LE; Krasnoyarsk Territory, Taimyr Peninsula, Dolgano-Nenetsk District, Putorana Plateau, Talnakh Settlement, I. V. Czernyadjeva (M. B. HepnadbeBa), LE B0023668; Chukotka Autonomus Area, Wrangel Island, Upper course of V'yuchny Creek, Afonina LE B0023655; Somnitel'naya Bay, Afonina, LE B0023664; ibid., Afonina, LE B0023666; Anadyr District, Anadyr River Basin, upper course of Yablon River, Afonina, LE B0023669; Magadan Region, Khasyn District, watershed of Ola and Bolshaya Khaya
Fig. 8. Distribution of Oncophorus virens subsp. minor in Russia.
rivers, O. Yu. Pisarenko (0. №. nucapeHKo) 7076, LE B0023667; Kamchatka Territory: Kamchatka, Kl'yuchevskaya group of volcanoes, I. V. Czernyadjeva 49(2), LE; ibid., 23 VIII 2005, I. V. Czernya-djeva s. n, LE.
Oncophorus integerrimus Hedenas, 2017, Eur. J. Taxon. 315: 26, 10 = Oncophorus virens var. elongatus Limpr., 1886, Laubm. Deutschl. 1: 309. = Cynodon-tium virens var. elongatum (Limpr.) Monk., 1927, Laubm. Eur. 195.
(Fig. 6A-C, 7C-E, 9, 10)
Plants 1-5(7) cm long, dark green, yellow-green or brownish, in loose tufts. Stems erect, weakly branching, with large central strand; cortex of 1-3 layers of incrassate cells; rhizoids red-brown, smooth, branched, in leaf axils. Leaves 2.4-3.2(4.0) x 0.61.0 mm, from ovate or rounded-oblong base tapering to narrowly triangular acumen, apex longly acuminate, leaves loosely incurved and curled or weakly twisted when dry, and erect spreading when moist; margins broadly recurved in the middle part of leaf, mostly entire or slightly obtusely denticulate in the upper part; lamina unistratose, rarely bistratose, on margins always bistratose; costa strong, 75-145 jm wide near the base, ending in leaf apex or shortly excurrent, in cross-section with dorsal and ventral epidermis, one layer of guide cells and 2-4 layers of dorsal and ventral stereids; lamina cells in acumen incrassate, subquadrate and rectangular, 5.5-25 x 5-11 jm, in mid-leaf incrassate, 6.5-30 x 6-10 jm, and in sheathing lamina less incrassate, longer and slightly porose, 16-65 x 6-11 jm; transition between the median and basal cells gradual; alar cells oblong rectangular, slightly inflated, forming delimited group. Monoecious. Sporophytes often, 1 (rare 2 from the same perichaetium); perigonia lateral on
stem, perigonial leaves with acute apex; inner perichaetial leaves up to 6.0 mm long, from high vaginal base suddenly narrowed to long, narrow acumen. Seta long, 1018 mm, yellow, becoming brown with age; capsule cylindric, 1.5-2.0 x 0.5-0.8 mm, curved, strumose, furrowed when dry; peristome ruby red and vertically pitted-strio-late, papillose, teeth united in short basal membrane, bifid at apex; spore 18-28 |im.
Fig. 9. Oncophorus integerrimus (Caucasus, Karachaevo-Cherkessian Republic, 3 IX 2005, M. S. Ignatov, E. A. Ignatova 159, MHA).
A — basal leaf cells; B, C, H — leaf transverse section; D, E — leaves; F — leaf tip; G — upper leaf cells; I — habit, dry. A-C, F, G, H — 100 |m; D, E — 1 mm; I — 10 mm.
Distinctions. Oncophorus integerrimus is close to O. virens subsp. virens, and distinguished by entire or almost entire leaf margins, whereas O. virens subsp. virens has predominantly dentate or denticulate upper leaf margins; from O. virens subsp. minor it distinguished by larger plants (1-5 cm vs. 1-3 cm) and leaves (2.4-3.2 mm vs. 1.7-2.15 mm). Moreover, cells in the upper part of vegetative leaves in O. integerrimus quadrate or short rectangular whereas in O. virens subsp. minor shorter or transversely rectangular.
Distribution. Oncophorus integerrimus was described relatively recently and its distribution is far from being fully revealed. According to the revision of the herbarium collections from Russia, this species is rather common in the Arctic (Islands of Franz Jozef Land Archipelago, Murmansk Region, Novaya Zemlya Archipelago, Severnaya Zemlya, Taimyr Peninsula, Novosibirskie Islands, lower course of Lena River, common in Wrangel Island, widespread in the Chukotka Peninsula, and South Chukotka). On-cophorus integerrimus is less common in the southern regions, where it occurs usually in
mountains at an altitude of more than 1000 m a. s. l.; few finds of this species are known from the Republic of Karelia, North Urals, Russian Caucasus, Tomsk Region, Altai, Republic of Yakutia, Buryatia, Magadan Region, Kamchatka Peninsula, Commander Islands (Fig. 9). Outside of Russia O. integerrimus is known from Scandinavia (Hedenas, 2017) and the North American Arctic, namely from Seward Peninsula, Alaska, USA; Ellef Ringens Island, Nunavut and Prince Patrick Island (Queen Elizabeth Islands), Northwest Territories, Canada (Ellis et al, 2020). These North American records are based on our revision of kept in LE herbarium materials on the genus Oncophorus from North America. Thus, based on the received data, O. integerrimus can be characterized as an arctomontane circumpolar species with a predominant distribution in the Arctic.
Fig. 10. Distribution of Oncophorus integerrimus in Russia.
Ecology. In tundra zone, this species often occurs in wet low bush-moss tundra and wetlands, also growing in moss communities along streams and banks of lakes, on wet rocks and also in floodplain willow and alder thickets.
Selected specimens examined. EUROPEAN RUSSIA: Arkhangelsk Region, Franz Jo-zef Land Archipelago: Hooker Island, Sedov Cape, V. P. Savicz (В. П. Савич), LE B0021799; Nansen Island, S. S. Kholod, LE B0021796; Meibl Island, S. S. Kholod, LE B0021794; Jackson Island, S. S. Kholod, LE B0021793; Wiener Neustadt Island, S. S. Kholod, LE B0021792; Novaya Zemlya Archipelago: Zhelaniya Cape, S. S. Kholod, LE B0024348; Mashigina Bay, I. V. Palibin, LE B0021825; South Island, Moller Bay, S. V. Sidorenko, I. V. Czernyadjeva, LE B0021835; Tsvetnoi Island, S. V. Sidorenko, I. V. Czernyadjeva, LE B0021770; Solovki Archipelago: Bolshoi Muksal-ma Island, T. A. Maksimova (Т. А. Максимова) Б-02/II, PTZ; Murmansk Region: Kola Peninsula, Rybachii Peninsula, Tsipnavlok Settlement, Bomansson, LE B0021815; ibid, valley of Srednii Creek, K. B. Popova (К. Б. Попова), MW 9050546; Orlov Cape, V. P. Savicz (В. П. Савич), LE B0021803; Gav-rilov Bay, V. P. Savicz (В. П. Савич), LE B0021813; Turii Peninsula, R. N. Schljakov (Р. Н. Шляков), LE B0021797;ibid, O. A.Belkina(О. А.Белкина)8043, KPABG; Laplandskii Reserve, E.N. Andrejeva
(Е. Н. Андреева) 2A050884, LE B0021832; Lovozerskie Mts., Lovozero, O. A. Belkina (О. А. Белкина) 74/7, KPABG; Kandalakshskii Gulf, Olenii Island, O. A. Belkina (О. А. Белкина) 265, KPABG; Kandalaksha Reserve, Monchegorsk District, Chuna-tundra, O. A. Belkina (О. А. Белкина) 19468, KPABG; Khibin Mts., Vud'yavrchorr, R. N. Schljakov (Р. Н. Шляков), LE B0021807; Pechenga District, "Pasvik" Reserve, M. A. Boichuk (М. А. Бойчук) 12936, PTZ; Republic of Karelia: Loukhskii District, Kuonsu-yarvi, L. A. Volkova, (Л. А. Волкова), LE B0021828; "Keret'skii" Reserve, A. I. Mak-simov, T. A. Maksimova (А. И. Максимов, Т. А. Максимова) 141-48, PTZ; Natonal Park "Paanayarvi", T. A. Maksimova (Т. А. Максимова) naa-88/3, PTZ; Nenets Autonomous Area: Kanin Peninsula, Kanin Nos, F. I. Ruprecht, LE B0023659; Kolguev Island, F. I. Ruprecht, LE B0023660; Pes-chanka River, O. V. Lavrinenko (О. В. Лавриненко), LE B0022158; Malozemel'skaya tundra, Malaya Labagei-sede, R. N. Schljakov (Р. Н. Шляков), LE B0021695; Bolshezemel'skaya tundra, Vangureimusyur hill, O. V. Lavrinenko, LE B0022156; Coast of Barents Sea, Bolshaya Dvoi-nichnaya River, A. Kochergina, LE B0021823; Komi Republic:150 km W of Workuta, O. V. Ivanov,
D. G. Donskov, MHA 9018463; Caucasus: Karachayevo-Circassian Republic: Klukhor District, Te-berda Nature Reserve: A. L. Abramova, 1.1. Abramov (А. Л. Абрамова, И. И. Абрамов), LE B0021837; Kabardino-Balkarian Republic: Adyrsu canyon, G. Ya. Ukrainskaya (Г. Я. Украинская) 13959, LE B0022088; Republic of Daghestan: Dagestania superior montosa ad fl. Ilan-chewi, F. I. Ruprecht, LE B0021839; ASIAN RUSSIA, West Siberia, Yamal-Nenets Autonomuos Area: Sob' River, I. V. Czernyadjeva (И. В. Чернядьева), LE B0023662; Khanty-Mansi Autonomous Area: Lyapi-na River, V. B. Sochava (В. Б. Сочава), LE B0021830; Berezovsk District, Malaya Pol'ya River, I. D. Kil'dyushevskii (И. Д. Кильдюшевский), LE B0021827; Tomsk Region: Topchugan River,
0. L. Kuznezow 2734, LE B0022092; Republk of Altai: Sailyugem Range, Tarkhata River, 1926, V. Baranov (В. Баранов), LE B0022090; East Siberia. Krasnoyarsk Territory: Severnaya Zemlya Archipelago: Oktyabrskoi Revolutsii Island, I. N. Safronova (И. Н. Сафронова), LE B002209; Bol'shevik Island, N. V. Matveeva (Н. В. Матвеева), LE B0022097; Taimyr Dolgano-Nenetsk District: Mamontovaya River, B. A. Tikhomirov, Uvarov, (Б. А. Тихомиров, Уваров), LE B0022094; Pyasina River, Tareya Settlement, L. S. Blagodatskikh (Л. С. Благодатских), LE B0022100; Marii Pronchishshevoi Bay, N. V. Matveeva, O. I. Sumina (Н. В. Матвеева, О. И. Сумина), LE B0022096; Ust'-Avamskaya Tundra, Kystyktakh river valley, E. D. Lapshina (Е. Д. Лапшина) 324E/2-21, LE B0025403; Putorana Plateau, Ayan Lake, I. V. Czernyadjeva (И. В. Чернядьева) LE B0022188; Lama Lake, I. V. Czernyadjeva (И. В. Чернядьева) 41, LE B022099; Republic of Sakha (Yakutia): New Siberian Islands (Novosibirskie Islands), Stolbovoi Island: R. K. Sisko, LE B0022102; ibid.,
1. V. Czernyadjeva (И. В. Чернядьева), LE B0022106; Lower course of Lena River, Tiksi Settlement, A. E. Katenin (А. Е. Катенин), LE B0022101; Moma District, Ulakhan-Chistai Range, M. S. Ignatov,
E. A. Ignatova, MHA 9028383; Repubic of Buryatia: Tunka District, Sayan Mts., Ikhe-Ukhgun River, A. A. Elenkin (А. А. Еленкин), LE B0022103; Severo-Baikal'sk District, Barguzin Reserve, Afonina s. n., LE B0022109; Khamar-Daban Range, Baikal'skii Reserve, S. G. Kazanovsky (С. Г. Казанов-ский), LE B0022105; Kurumkan District, Barguzin Ridge, L. Dorzhieva, Tubanova (Л. Доржиева, Тубанова) Ba161004, UUH. Chukotka Autonomous Area: Wrangel Island, Somnitel'naya Bay, Afonina, LE B0022140; Lednikovaya Creek, Afonina, LE B0022132; Berry Mt., B. N. Goro-dkov (Б. Н. Городков), LE B0022141; Central Mts., R. P. Obabko, LE B0022154; Chukotka District: Chegitun River, Afonina, LE B0022115; Lorino Hot Springs, Afonina s. n., LE B0024321; Lavrentia Bay, Lavrentia Settlement, Afonina, LE B0022126; ibid, Lavrentia Bay, Krause Cape, Afonina, LE B0022121; Provideniya District: Chaplin Hot Springs, Gavrilyuk, Gagarin (Гаври-люк, Гагарин), LE B0022123; Yanrakynnot Settlement, Afonina, LE B0022114; Gilmimliveem Hot Springs, Afonina, LE B0022128; Penkignei Bay, Afonina s. n., LE B0024322; Cross (Krest) Gulf, Egvekynnot Settlement, Afonina, LE B0022129; Anadyr District: Pekul'nei Range, Yuzhny Pekul-nei River, Afonina, LE B0022117; Anadyr River basin, Tamvatnei River, Afonina, LE B0022135; Pekulnei Lake, Kakanaut Bay, E. Yu. Kuzmina, LE B0022133; vicinity of Meinypil'gyno Settlement, E. Yu. Kuzmina, LE B0022138; Emravaam River, E. Yu. Kuzmina, LE B0022134; Magadan Region:
Ola District, P'yagin Peninsula, L. S. Blagodatskikh (Л. С. Благодатских), LE B0022150; Kamchatka Territory: Kamchatka Peninsula, Sredinnii Range, vicinity Esso Settlement, I. V. Czernya-djeva, LE B0022149; Ushakov volcano "Klyuchevskoy dol", I. V. Czernyadjeva, LE B0022147; Ostryi Tolbachek volcano, I. V. Czernyadjeva, LE B0022145; Commander Islands, Bering Island, Fedo-sov 10-3-615, MHA.
Acknowledgments
The authors are grateful to curators of herbaria MW, MHA, KPABG, IRK, PTZ, NSK for loan of specimens. We also thank Jan Kucera (Ceske Budejovice) for providing the ITS sequences of On1556 and On1561. The work of O. M. Afonina was conducted within framework of institute theme «Flora and taxonomy of algae, lichens and bryophytes in Russia and phytogeographically important regions of the world» № 121021600184-6. The work of V. E. Fedosov and O. D. Dugarova was partly supported by RSF Grant 18-14-00121-П. The work of V .E. Fedosov was carried out within framework of the Botanical Garden-Institute FEB RAS scientific project 122040800088-5.
The work of D. Ya. Tubanova was carried out within framework of IGEB SB RAS №121030900138-8.
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