MOSSES AND LIVERWORTS OF STOLBOVOY ISLAND (NEW SIBERIAN ISLANDS ARCHIPELAGO, YAKUTIA) Текст научной статьи по специальности «Биологические науки»

Mosses and liverworts of Stolbovoy Island (New Siberian Islands Archipelago, Yakutia)

I. V. Czernyadjeva1, A. D. Potemkin1, E. I. Troeva2

'Komarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia 2Institute for Biological Problems of Cryolithozone of the Siberian Branch of the Russian Academy

of Sciences, Yakutsk, Russia Corresponding author. I. V. Czernyadjeva, [email protected], [email protected]

Abstract. The paper provides the data on mosses and liverworts of the Stolbovoy Island, situated in the New Siberian Islands of the Arctic Ocean, on the far north of the Republic of Sakha (Yakutia). The annotated list includes 140 species of mosses and 58 species of liverworts. For every species, data on localities, frequency, habitats are provided. For the first time, 119 species of mosses and 54 species of liverworts were revealed for Stolbovoy Island, and 40 species of mosses and 22 species, 5 varieties, and 2 forms of liverworts for the New Siberian Islands. Schistidiumgrandirete, Cephalozia hamatiloba, Lophozia cf. svalbardensis, Scapania cf. ligulifolia are reported for the first time for Yakutia. Bryo-erythrophyllum rubrum, Rhizomnium gracile, Sphagnum concinnum were revealed for the first time for Yakutia in our collections from Stolbovoy Island but published earlier. A new combination for Pseudotritomaria heterophylla f. anomala is published.

Keywords: Bryophyta, Marchantiophyta, bryophytes, flora, hepatics, mosses, Arctic, East Siberia, Republic of Sakha (Yakutia), Russia.

Мхи и печеночники острова Столбовой (архипелаг Новосибирские острова, Якутия)

И. В. Чернядьева1, А. Д. Потемкин1, Е. И. Троева2

1Ботанический институт им. В. Л. Комарова РАН, Санкт Петербург, Россия 2Институт биологических проблем криолитозоны СО РАН, Якутск, Россия Автор для переписки. И. В. Чернядьева, [email protected], [email protected]

Резюме. В статье представлены результаты изучения мхов и печеночников о. Столбового [архипелаг Новосибирские о-ва, Республика Саха (Якутия)]. Аннотированный список включает 140 видов мхов и 58 видов печеночников. Для каждого из них приводятся число находок, местонахождения и местообитания. Впервые для о. Столбового выявлены 119 видов мхов и 54 видов печеночников, а для архипелага Новосибирские о-ва — 40 видов мхов и 22 вида, 5 разновидностей и 2 формы печеночников. Schistidium grandirete, Cephalozia hamatiloba, Lophozia cf. svalbardensis, Scapania cf. ligulifolia приводятся впервые для Якутии. Bryoerythrophyllum rubrum, Rhizomnium gracile, Sphagnum concinnum выявлены впервые для Якутии в наших сборах с о. Столбового, но были опубликованы ранее. Обнародована новая комбинация для Pseudo-tritomaria heterophylla f. anomala.

Ключевые слова: Bryophyta, Marchantiophyta, мохообразные, флора, Арктика, Восточная Сибирь, Республика Саха (Якутия), Россия.

https\//doi.org/10.31111/nsnr/2021.55.2.439

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The New Siberian Islands Archipelago is located in the Arctic Ocean between the Laptev Sea and the East Siberian Sea, on the far north of the Republic of Sakha (Yakutia). The archipelago consists of three groups of islands: the Lyakhovsky Islands (Bolshoy Lyakhovsky, Maly Lyakhovsky, Stolbovoy, etc.), the Anjou Islands (Kotel-ny, New Siberia, etc.), and the De Long Islands (Zhokhov, Bennett, etc.). It is located within the subzone of High Arctic tundras and belongs to the Yano-Kolyma Province of the Arctic Floristic Region (Yurtsev et al, 1978).

This area is exceedingly difficult to access, and it remains almost uninhabited and still insufficiently explored. First known to us data on the moss flora of the New Siberian Islands were published by Brotherus (1910), who cites 27 species. A later study of the flora and vegetation of Kotelny Island was carried out in 1947 by Gorodkov (1956), who published 32 species of liverworts and 95 species of mosses. In 1956, the study of the flora of the archipelago was continued by a complex physical and geographical expedition of the Arctic Research Institute (at present Arctic and Antarctic Research Institute). The results of the study of collections, gathered by this expedition, were published by Abramov (1963), who reported 12 species of liverworts and 71 species of mosses for Stolbovoy, Kotelny, and Bennett islands. Aleksandrova (1963) reported 41 species of mosses and 2 liverworts for Bolshoy Lyakhovsky Island. Zhukova, Sumi-na (1976), and Zhukova (1982) reported 27 liverwort species for Kotelny Island and 55 species for Bolshoy Lyakhovsky Island. Stepanova (1986), in her generalizing work on the moss flora of tundras of Yakutia, mentioned 72 moss species for the archipelago but did not provide the geographical location for many species widespread in Yakutia. Later she published the list of liverworts and mosses of the Yakutian tundras in "Flora of tundra zone of Yakutia" (Egorova et al, 1991) with report of 52 liverwort species for Kotelny Island and Lyakhovsky Islands, without records for Stolbovoy Island. Based on the results of processing of small collections gathered in the eighties, 48 species and two varieties of mosses and 4 species of liverworts were published for Bennett Island (Abramov, Abramova, 1992), and 43 species of mosses for Zhokhov Island (Samarsky et al, 1997). Reports of separate species for the archipelago were published by Abramova and Abramov (1966), and by Sumina (1975, 1986). Afonina and Czernyadjeva (1995) listed 163 species for the New Siberian Islands summing up the results of the study of mosses of the Russian Arctic. Afonina (2015) recorded 69 species for the De Long Islands in her revision of the moss flora of the polar deserts. Published three volumes of the "Moss Flora of Russia" (Ignatov et al, 2017, 2018, 2020) list 57 moss species for the archipelago. Thus before our study about 205 species of mosses and 64 liverworts were known for the New Siberian Islands.

Stolbovoy Island is the smallest and the least investigated of the New Siberian Islands. It is situated 360 km North-East of Tiksi and stretches for nearly 50 km in the northwest direction (Fig. 1). Its latitudinal position is intermediate between the better investigated Bolshoy Lyakhovsky and Kotelny islands. Despite Stolbovoy Island is well studied in respect of geology and geomorphology (Voronkov, 1958; Sisko, 1968; etc.), its vegetation and flora remain hardly explored. There were only palynological

studies of the Tertiary sediments in the vicinity of the Stolbovaya River mouth (Sisko, 1968), as well as the records of 29 moss species and 5 liverwort species collected by R. K. Sisko during the physiographic research conducted by the Arctic Research Institute (Abramov, 1963; Stepanova, 1986).

Study area

The climate of Stolbovoy Island is harsh. According to Worldclim climatic data (1970-2000) (Fick, Hijmans, 2017), the average winter temperature is -33 °С (min. -38 °С), the average summer temperature does not exceed +4 °С (max. + 12 °С). The frost-free period lasts 12-14 days. Prevailing anticyclonic weather comes with low precipitation amounts (up to 150-170 mm per year), and the melting ice wedges in cryogenic landscapes of the tundra serve as a primary moisture source for plants. Permanent winds impede the development of continuous vegetation cover on most elevated and exposed to wind surfaces (Aleksandrova, 1977).

Fig. 1. Collecting places (1, 2, 3) on Stolbovoy Island and its location within the New Siberian

Islands.

There are three hypsometric levels on the island: lowland plains, flat ridges, and residual elevations (Fig. 2). The body of the island is represented by the mass of the Mesozoic turbidites with sandstones as prevailing facies. The Tertiary, Upper Quaternary, and modern deposits, mainly sandstones and argillites, are also characteristic of various terrace levels of the accumulation-denudation plain. The island features active thermoerosion processes resulting in specific landform development (Sisko, 1968).

Our study was conducted in the northern part of the island. The striped tundra landscape can be observed on the slopes of the most elevated flat ridges. The dwarf shrub-graminoid mossy tundra communities intermitted with stone agglomerations as medallions and stripes cover the ridge tops (ca. 100 m a. s. l.).

Fig. 2. The coastal part of Stolbovoy Island, view from a helicopter.

The medium level terraces are characterized by dissected topography. Hummocky tundra with bare spots of displaced ground is widespread there. Polygonal-fissured and baidjarakh (frost mound) complexes are common there also.

The lowland plains feature flat mound-hollow and baidjarakh complexes (Fig. 3, 5). Polygonal-ridged complexes with poorly developed ridges sometimes occur there (Fig. 4).

Fig. 3. Bryum cryophilum (1) and Calliergon giganteum var. sibiricum (2) from brook banks between

baidjarakhs (3).

According to the geobotanical zonation (Aleksandrova, 1977), the island is situated within the northern stripe of the Arctic tundra (the New Siberian Islands District of the East Siberian Province). The zonal vegetation represents various patterns of dwarf shrub-graminoid moss and lichen-moss tundra, mainly on weakly gleyed humic soils (Desyatkin et al., 2009). The projection cover values range from 30% to 70%. The total height of communities does not exceed 5-10(30) cm (Fig. 6). The following species feature the highest constancy values: Salix polaris, Alopecurus alpinus, Luzula confusa, Dupontia fisheri, Papaver polare, Oxyria digyna, Saxifraga spp., and Draba spp. The first three species also co-dominate in zonal communities. The stony slopes bear forb-Racomitrium tundra. The gully slopes are subject to ther-moerosion and thus covered with loose vegetation predominated by forb species (Oxyria digyna, Papaver polare, species of Ranunculaceae). The widespread communities of Dupontia fisheri, Alopecurus alpinus, and Arctophila fulva are confined to hollows and run-off landforms. Occasionally, Carex stans and Eriophorum scheuchzeri coenoses were observed there.

Material and Methods

The present publication is based on the identification of bryophytes collected by I. V. Czernyadjeva and E. I. Troeva from 2 to 7 August 2019. All literature sources on bryophytes of Stolbovoy Island were considered (Abramov, 1963; Stepanova, 1986). The mosses and liverworts in three remote collecting places (Fig. 1) are noted as follows in the annotated list below: 1 — the northwest coast in the vicinity of the polar station (~ 74°10'N, 135°28'E); 2 — the northern tip of the island, in the vicinity of Cape Skalisty (~ 74°13'N, 135°27'E); 3 — the northwest coast in the vicinity Melkoe Lake (~ 74°09'N, 135°33'E). In total, over 800 moss and over 400 liverwort identifications we made.

Results

The nomenclature of the annotated list follows "Moss flora of Russia" (Ignatov et al, 2017, 2018, 2020) and Hodgetts, Lockhart (2020) for mosses and to "Liverworts and hornworts of Russia" (Potemkin, Sofronova, 2009) for liverworts with some updates from the recent literature. Annotation of each species includes literature data, the number of our records, mentioned above numbers of collection places, habitats, associated species, presence of structure associated with reproduction (S+ — sporo-phytes, gem. — gemmae, per. — perianths). For rare species found in 1-4 specimens, the collector numbers are indicated. Collections of Czernyadjeva have numeric numbers like 1-19, whereas collections of Troeva have numbers with letters te, like 10te19. The species reported for the first time for the New Siberian Islands are marked with an asterisk (*). All specimens are deposited in the Bryological Herbarium of the Komarov Botanical Inctitute (LE).

Division Bryophyta — Mosses

Aloina brevirostris (Hook. et Grev.) Kindb. — 2 records. 1: on bare soil on slope of baidja-rakh, 30-19; 2: in crack of polygonal tundra with Hennediella heimii var. arctica, 16-19; S+.

Fig. 4. Dicranella subulata (2) from bare loam in polygonal tundra (1).

Fig. 5. Pogonatum dentatum (2) from the slope of baidjarakh (1).

Fig. 6. Tetraplodon mnioides (2) from hummocky spotty Luzula-graminoid-lichen-moss tundra (1).

*Amphidium lapponicum (Hedw.) Schimp. — 2 records. 2: in herb-dwarf shrub-moss tundra, with liverworts, 20-19, S+, det. O. M. Afonina; in dwarf shrub-grass-moss rubble tundra, with Flexitrichum gracile, Scorpidium cossonii, 25tе19.

Andreaea papillosa Lindb. — 1 record. 2: in herb-dwarf shrub-moss tundra, with liverworts, 20-19, S+.

Aplodon wormskioldii (Hornem.) R. Br. — 2 records. 1: on soil of Dupontia meadow, 32-19; on bare soil of baidjarakh slope, with Bryum sp., Pohlia drummondii, 36-19.

Aulacomnium palustre (Hedw.) Schwägr. — 5 records. 1: in hummocky dwarf shrub-gram-inoid, dwarf shrub-grass-moss tundra; 3: dwarf shrub-Dupontia-Luzula-moss and Dupontia tundra, Dupontia-sedge-moss mire.

A. turgidum (Wahlenb.) Schwägr. — Abramov, 1963. 37 records. 1, 2, 3: in different types of tundra, Dupontia and herb-Luzula meadow, Dupontia-sedge-moss mire.

Bartramia ithyphylla Brid. — Abramov, 1963. 26 records. 1, 2, 3: in different types of tundra, Dupontia and herb-grass meadow, on bare loam, on bare soil of baidjarakh slope; S+.

Brachythecium cirrosum (Schwägr.) Schimp. — 2 records. 1: on spot of bare soil in Dupontia meadow, with Philonotis tomentella, Drepanocladus arcticus, 32-19; 2: on bare soil on top of baidjarakh, 15-19.

B. mildeanum (Schimp.) Schimp. ex Milde — 1 record. 1: in grass-Saxifraga-moss tundra, 9-19.

B. turgidum (Hartm.) Kindb. — 5 records. 1: in grass-Saxifraga-moss, dwarf shrub-grass-moss and dwarf shrub-Luzula spotty tundra and Dupontia meadow; 2: in dwarf shrub-grass-moss rubble tundra, with Aulacomnium turgidum, Drepanocladus arcticus, etc.

Bryoerythrophyllum ferruginascens (Stirt.) Giacom. — 1 record. 1: on bare soil in Luzu-la-dwarf willow spotty tundra, 33-19, gem.

B. recurvirostrum (Hedw.) P. C. Chen — 4 records. 1: on bare soil in dwarf shrub-grass-moss tundra, 8-19; in rock crevices on rocks by the sea, 12-19; on bare loam, 23-19; on bare soil of baidjarakh slope, 30-19; with Distichium capillaceum, D. inclinatum, Orthothecium chryseon; S+.

B. rubrum (Jur. ex Geh.) P. C. Chen — Ellis et al, 2021. 1 record. 1: dry terrace, on soil in grass-dwarf shrub-moss tundra, 9-19, S+.

Bryum cf. amblyodon Müll. Hal. — Abramov, 1963; Stepanova, 1986. 1 record. 1: on bare soil of baidjarakh slope, 36-19, S+.

Bryum cf. arcticum (R. Br.) Bruch et Schimp. — 2 records. 1: in grass-Saxifraga-moss tundra, 9-19; in grass-dwarf shrub-moss tundra, 9'-1, S+.

B. calophyllum R. Br. — 3 records. 1: on bare soil in dwarf shrub-grass-moss tundra, 8-19; in grass-Saxifraga-moss tundra, 9-19; in grass-dwarf shrub-moss tundra, 9'-19; with Ceratodon purpureus, Bryum sp.; S+.

B. cryophilum Märtensson (Fig. 3: 1) — Abramov, 1963; Stepanova, 1986. 6 records. 1: in Dupontia-moss meadows, graminoid-moss meadow in runoff hollow; 2: in moss community near snowfield and Equisetum-Dupontia meadows; with Pohlia wahlenbergii, Bryum sp., etc.

B. cyclophyllum (Schwägr.) Bruch et Schimp. — 4 records. 1: in dwarf shrub-grass-moss tundra, 8-19; on bank of stream, in splits between baidjarakhs, 31-19; 2: in moss community near snowfield, 19-19; in Dupontia meadow, 21-19; with Orthothecium chryseon, Tomentypnum involutum, Warnstorfia sarmentosa, etc.; S+.

*B. pseudotriquetrum (Hedw.) P. Gaertn. et al. — 1 record. 2: in moss community near snowfield, 19-19.

B. rutilans Brid. — 1 record. 1: on bare soil of baidjarakh slope, with Timmia norvegica, 30-19.

B. teres Lindb. — 3 records. 1: in grass-dwarf shrub-moss tundra, 9-19; in rock crevices on rocks by the sea, 12-19; on bare loam, 23-19; with Distichium sp., Bryum sp.

Calliergon giganteum (Schimp.) Kindb. var. sibiricum Ignatova et Czernyadjeva (Fig. 3: 2) — 5 records. 1: in dwarf shrub-grass-moss tundra, 8-19; in grass-Saxifraga-moss tundra, 9-19; on spot of bare soil in Dupontia meadow, 32-19; in hummocky dwarf shrub-graminoid tundra, 34-19; 2: in Equisetum-Dupontia meadow, 22-19.

C. richardsonii (Mitt.) Kindb. - 2 record. 1: in grass-moss tundra in depression, with Scor-pidium revolvens, 29-19; on bank of stream in splits between baidjarakhs, with Sarmentypnum sarmentosum, 31-19.

Campylium bambergeri (Schimp.) Hedenas et al. — 1 record. 2: in herb-dwarf shrub-moss tundra, 20-19.

Ceratodon heterophyllus Kindb. — 1 record. 1: in grass-dwarf shrub-moss tundra, with Bryum sp., Ceratodonpurpureus, Hymenoloma crispula, Psilopilum leavigatum, 9-19.

C. purpureus (Hedw.) Brid. — 4 records. 1: in dwarf shrub-grass-moss tundra, 8-19; in grass-Saxifraga-moss tundra, 9-19; on bare soil in grass-dwarf shrub-moss tundra, 9-19; on spot of bare soil in hummocky spotty Luzula-graminoid-lichen-moss tundra, 25-19; with Bryum sp., Hymenoloma crispula; S+.

Cinclidium arcticum (Bruch et Schimp.) Schimp. — Abramov, 1963. 1 record. 1: in grass-moss tundra in depression, with Loeskypnum badium, Oncophorus demetrii, 27-19.

C. latifolium Lindb. — 3 records. 1: in dwarf shrub-grass-moss tundra, 8-19; in hollow in hummocky dwarf shrub-graminoid tundra, 34-19; 2: in swampy herb-Sphagnum-moss tundra, 18-19; with Drepanocladus arcticus, Scorpidium revolvens, etc.

C. subrotundum Lindb. — 2 records. 1: in Dupontia-moss community, 35-19; in Dupon-tia-moss mire, 56te19.

*Cnestrum schistii (F. Weber et D. Mohr) J. I. Hagen — 1 record. 3: in Luzula-dwarf willow-moss hummocky tundra, 48te19, S+.

Conostomum tetragonum (Hedw.) Lindb. — 13 records. 1, 3: in different types of tundras, herb-grass meadow, on bare soil in sedge-moss mire and on bare soil of baidjarakh slope; S+.

*Dichodontium pellucidum (Hedw.) Schimp. — 2 record. 1: the edge of terrace, on stony-gravelly area, with Encalypta alpina, 10-19, det. O. M. Afonina; on spot of bare soil of Dupontia meadow, 32-19.

Dicranella crispa (Hedw.) Schimp. — 11 records. 1, 3: on bare soil in grass-dwarf shrub-moss, dwarf shrub-Luzula spotty and hummocky dwarf shrub-graminoid tundras, on bare soil in herb-grass and herb-Luzula meadows and on baidjarakh slopes; with Bartramia ithyphylla, Dicranella subulata, Pohliaproligera, Psilopilum cavifolium, etc.; S+.

D. subulata (Hedw.) Schimp. (Fig. 4: 2) — 6 records. 1: on bare soil in Luzula-grass-lichen and hummocky spotty herb-moss tundra and on bare loam; 3: on bare soil in dwarf shrub-Lu-zula-moss-lichen, hummocky dwarf shrub-graminoid tundra, in herb-grass meadow; with Di-cranella crispa, Psilopilum cavifolium, Distichium inclinatum, etc.; S+.

*Dicranum acutifolium (Lindb. et Arnell) C. E. O. Jensen — 9 records. 1, 2, 3: in different types of tundra and herb-Luzula and Dupontia-moss meadows.

D. elongatum Schleich. ex Schwagr. — 18 records. 1, 2, 3: in different types of tundra, herbgrass meadow, sedge-moss mire.

D. laevidens R. S. Williams — 14 records. 1, 3: in different types of tundra, Dupontia-moss and Dupontia-sedge-moss mire and Dupontia-moss meadow.

*Dicranum majus Turner — 7 records. 1: in grass-moss, hummocky spotty herb-moss, dwarf shrub-Dupontia-Luzula-moss and dwarf shrub-grass-forb hummocky tundra; 2: in swampy herb-Sphagnum-moss tundra.

D. spadiceum J. E. Zetterst. — 6 records. 1: in dwarf shrub-Luzula spotty and dwarf shrub-graminoid tundra, graminoid-moss meadow in runoff hollow; 2: in dwarf shrub-grass spotty tundra; 3: in graminoid-moss-lichen nival tundra; S+.

Distichium capillaceum (Hedw.) Bruch et Schimp. — Abramov, 1963. 6 records. 1: in dwarf shrub-grass-moss and grass-Saxifraga-moss tundra, on bare soil in hummocky spotty herb-moss tundra and on baidjarakh slope; 2: in dwarf shrub-grass hummocky tundra; S+.

D. inclinatum (Hedw.) Bruch et Schimp. — 9 records. 1: in grass-Saxifraga-moss tundra, Dupontia-moss meadow, on bare soil of baidjarakh slope; 2: on spot in dwarf shrub-graminoid spotty tundra, on bare soil on top of baidjarakh, in fissure in polygonal tundra; with Bryoeryth-rophyllum recurvirostrum, Bryum sp., etc.; S+.

Drepanocladus aduncus (Hedw.) Warnst. — 2 records. 1: on spot of bare soil in Dupon-tia-moss meadow, 32-19; in hollow in hummocky dwarf shrub-graminoid tundra, 34-19.

D. arcticus (Williams) Hedenäs — Abramov, 1963. 15 records. 1, 2, 3: in swampy herb-Sphagnum-moss, grass-dwarf shrub-moss and dwarf shrub-grass spotty tundra, Dupontia and Dupontia-moss meadow, moss community near snowfield, on splits between baidjarakhs, stream bank and on the lakeshore.

D. brevifolius (Lindb.) Warnst. — 4 records. 2: in herb-dwarf shrub-moss tundra, with Drepanocladus turgescens, 20-19; dwarf shrub-grass hummocky tundra, 12te19; 3: Dupontia-cotton grass-moss meadow, 34te19; Dupontia tundra, 35te19.

D. turgescens (T. Jensen) Broth. — 5 records. 1, 3: in Dupontia meadow; 2: in moss community near snowfield, herb-dwarf shrub-moss and dwarf shrub-grass-moss rubble tundra; with Flexitrichum gracile, Drepanocladus brevifolius, etc.

*D. sordidus (Müll. Hal.) Hedenäs — 1 record. 1: in graminoid-moss meadow in runoff hollow, 13-19.

Encalypta alpina Sm. — 5 records. 1: on stony-gravelly area on the edge of terrace, on spots of bare soil in Dupontia and Dupontia-moss meadow 2: in dwarf shrub-grass-moss rubble and herb-dwarf shrub-moss tundra; with Disichium inclinatum, Flexitrichum flexicaule, F. gracile, Oncophorus demetrii, etc.; S+.

E. rhaptocarpa Schwägr. — 4 records. 1: in grass-Saxifraga-moss tundra, 9-19; on pebbles by the sea, 11-19; on bare soil on baidjarakh slope, 30-19; 2: on spot on polygonal tundra, 16-19; S+.

*Encalypta cf. vulgaris Hedw. — 1 record. 1: on soil in graminoid-moss meadow in runoff hollow, 13-19.

Eurhynchiastrum pulchellum (Hedw.) Ignatov et Huttunen — 2 records. 1: in Luzula-dwarf willow spotty tundra, 33-19; 3: in dwarf willow-grass moss hummocky tundra, 36te19.

*Fissidens bryoides Hedw. — 2 records. 1: on bare loam, with Pohlia cruda, Timmia austriaca, 23-19; on bare soil in hummocky spotty herb-moss tundra, with Pohlia drummondii, 24-19; S+.

Flexitrichum flexicaule (Schwägr.) Ignatov et Fedosov — Abramov, 1963; Stepanova, 1986. 16 records. 1, 2, 3: in different types of tundra, Dupontia meadow, moss community near snow-field, on pebbles by the sea and bare soil of baidjarakh slope.

F. gracile (Mitt.) Ignatov et Fedosov — Czernyadjeva et al, 2020a. 8 records. 1, 2, 3: in different types of tundra, herb-grass and herb-Luzula meadow.

Funaria arctica (Berggr.) Kindb. — Ellis et al, 2020. 2 records. 1: on bare soil in grass-dwarf shrub-moss tundra, with Bryum sp., Ceratodon purpureus, Dicranella crispa, Psilopilum leavi-gatum, 9-19; on bare soil on baidjarakh slope, with Hennediella heimii var. arctica, 36-19; S+.

*F. hygrometrica Hedw. — 1 record. 1: on bare loam, with Dicranella crispa, D. subulata, 23-19, S+.

*Gymnostomum aeruginosum Sm. — 1 record. 1: on bare soil in dwarf shrub-grass hummocky tundra, with Pohlia drummondii, 36te19, det. O. M. Afonina.

Hamatocaulis lapponicus (Norrl.) Hedenäs — 1 records. 1: edge of puddle in depression, 26-19.

Hennediella heimii (Hedw.) R. H. Zander var. arctica (Lindb.) R. H. Zander — 5 records. 1: on bare soil in grass-dwarf shrub-moss tundra and on baidjarakh slope, on bare loam, on rock crevices by the sea; 2: in crevices of polygonal tundra; with Bryum sp., Distichium capillaceum, Tortula muralis var. aestiva; S+.

Hygrohypnella polare (Lindb.) Ignatov et Ignatova — 1 record. 2: in moss community near snowfield, 19-19.

Hylocomium splendens (Hedw.) Schimp. — Abramov, 1963. 31 records. 1, 2, 3: in different types of tundra, Dupontia and Dupontia-moss meadow; Dupontia-sedge-moss mire.

Hymenoloma crispulum (Hedw.) Ochyra — Abramov, 1963. 4 records. 1: on stony-gravelly area on the edge of terrace, 10-19; in hummocky spotty herb-moss tundra, 24-19; on spot in Dupontia meadow, 32-19; 3: on bare soil in dwarf shrub-Luzula-moss-lichen tundra, 45te19; S+.

Hypnum cupressiforme Hedw. — 1 record. 1: on pebbles by the sea, 11-19.

Isopterygiella alpicola (Lindb.) Ignatov et Ignatova — 1 record. 2: in dwarf willow-grass-moss rubble tundra, with Flexitrichum gracile, 25te19.

I. pulchella (Hedw.) Ignatov et Ignatova — 4 records. 1: on stony-gravelly area on the edge of terrace, with Distichium inclinatum, Pohlia cruda, 10-19; in Luzula-dwarf willow spotty tundra, with Bartramia ithyphylla, Flexitrichum flexicaule, Timmia austriaca, 33-19; in hollow in dwarf willow-grass-forb hummocky tundra, with Bartramia ithyphylla, Flexitrichum gracile, Oncophorus demetrii, 10te19; 2: in dwarf willow-graminoid hummocky tundra, 12te19.

Kiaeria glacialis (Berrgr.) J. I. Hagen — 7 records. 1: in dwarf shrub-Dupontia-Luzula-moss, hummocky spotty herb-moss and dwarf shrub-grass-forb hummocky tundra, Dupontia and Du-pontia-moss meadow, on bank of stream in splits between baidjarakhs; 2: in Racomitrium-lichen rubble tundra.

Leptobryum pyriforme (Hedw.) Wilson — 3 records. 1: in stony-gravelly area on the edge of terrace, with Distichium inclinatum, 10-19; on bare soil in graminoid-moss meadow in runoff hollow, 13-19; on bare loam, with Bryum sp., Distichium sp., 23-19; S+.

*Lewinskya iwatsukii (Ignatov) F. Lara et al. — 1 record. 1: on bare soil on baidjarakh slope, 30-19.

Loeskypnum badium (C. C. Hartm.) Paul. — 3 records. 1: in grass-moss tundra in depression, with Scorpidium revolvens, 27-19; in Dupontia-moss community, 35-19; 3: in Dupon-tia-moss meadow, 31te19.

Meesia triquetra (Jolycl.) Angstr. — 1 record. 1: in dwarf shrub-grass-moss tundra, 8-19.

M. uliginosa Hedw. — Czernyadjeva et al, 2020a. 3 records. 1: on bare soil in grass-Saxi-fraga-moss tundra, 9-19; in Dupontia-moss meadow, 31te19; in Dupontia tundra, 35te19; S+.

*Mnium lycopodioides Schwägr. — 3 records. 1: on bare soil on baidjarakh slope, with Sa-nionia uncinata, 36-19; in dwarf willow-Dupontia moss tundra, 55te19; 2: in Racomitrium-lichen rubble tundra, 26te19.

Myurella julacea (Schwägr.) Schimp. — 10 records. 1: on stony-gravelly area on the edge of terrace, in dwarf shrub-Luzula-tundra, in Dupontia-moss meadow; 2: in different types of tundra; with Distichium inclinatum, Flexitrichum flexicaule, F. gracile, etc.

M. tenerrima (Brid.) Lindb. — 3 records. 1: on bare soil in grass-Saxifraga-moss tundra, with Meesia uliginosa, 9-19; 2: in dwarf willow-graminoid moss tundra, 15te19; in dwarf shrub-grass-moss rubble tundra, with Flexitrichum gracile, 25te19.

*Niphotrichum elongatum (Frisvoll) Bednarek-Ochyra et Ochyra — 2 records. 1: in Du-pontia-moss meadow, 50te19; 2: in moss community near snowfield, with Drepanocladus arcticus, Orthothecium retroflexum, Philonotis sp., 19-19.

N. ericoides (Brid.) Bednarek-Ochyra et Ochyra — 1 record. 1: in Dupontia meadow, 32-19.

N. panschii (Müll. Hal.) Bednarek-Ochyra et Ochyra — 3 records. 1: on pebbles by the sea, 11-19; in hummocky spotty Luzula-graminoid-lichen-moss tundra, 25-19; in Luzula-dwarf willow spotty tundra, with Flexitrichum flexicaule, Timmia austriaca, 33-19.

*Oncophorus demetrii (Renauld et Cardot) Hedenäs — 10 records. 1, 2, 3: in different types of tundra; with Flexitrichum gracile, Philonotis fontana, Tomentypnum involutum, etc.; S+.

*O. elongatus (I. Hagen) Hedenäs — 8 records. 1, 3: in hummocky dwarf shrub-graminoid, dwarf shrub-grass hummocky and dwarf shrub-Dupontia-Luzula-moss tundra, Dupontia-moss community, Dupontia-moss meadow, Dupontia-sedge-moss mire; with Dicranum majus, Tomen-typnum involutum, etc.; S+.

*O. integerrimus Hedenäs — 3 records. 1: in dwarf shrub-grass-moss tundra, 8-19; on spot in Dupontia meadow, 32-19; 2: in herb-dwarf shrub-moss tundra, 20-19.

O. virens (Hedw.) Brid. — Abramov, 1963; Stepanova, 1986. 1 record. 2: in Dupontia meadow, 21-19, det. O. M. Afonina.

Orthothecium chryseon (Schwägr.) Schimp. — 14 records. 1, 2, 3: in different types of tundra, Dupontia and Dupontia-moss meadow, Dupontia-sedge-moss mire.

O. retroflexum Ignatov et Ignatova — 2 records. 2: in moss community near snowfield, with Sarmentypnum sarmentosum, 19-19; on spot in herb-dwarf shrub-moss tundra, 20-19.

O. strictum Lorentz — 1 record. 1: on bare soil on baidjarakh slope, with Distichium capilla-ceum, Pohlia cruda, 30-19.

Philonotis fontana (Hedw.) Brid. — 1 record. 1: in grass-moss tundra in depression, with Oncophorus demetrii, Tomentypnum involutum, 28-19.

P. tomentella Molendo — Abramov, 1963; Stepanova, 1986. 7 records. 1: in dwarf shrub-grass-moss tundra, graminoid-moss and Dupontia meadows in runoff hollow, on bank of stream; 2: in dwarf shrub-graminoid spotty tundra, moss community near snowfield, in Dupontia meadow.

Plagiomnium curvatulum (Lindb.) Schljakov — 3 records. 1: in dwarf shrub-grass-moss tundra, with Bryoerythrophyllum recurvirostrum, Cinclidium latifolium, 8-19; in grass-Saxifra-ga-moss tundra, with Bryum sp., 9-19; 2: in grass-Saxifraga-moss tundra, 18-19.

P. ellipticum (Brid.) T. J. Kop. — 2 records. 1: in grass-Saxifraga-moss tundra, 34-19; in grass-moss tundra in depression, with Aulacomnium turgidum, Scorpidium revolvens, Tomentypnum involutum, 28-19.

Plagiothecium berggrenianum Frisvoll — Ellis et al, 2021. 2 records. 1: in hummocky spotty Luzula-graminoid-lichen-moss tundra, with Hylocomium splendens, Tomentypnum involutum, 25-19; in dwarf shrub-Dupontia-Luzula-moss tundra, with Oncophorus elongatus, 54te19.

Pogonatum dentatum (Brid.) Brid. (Fig. 5: 2) — 2 records. 1: on bare soil on baidjarakh slopes, with Pohlia proligera, Psilopilum cavifolium, 30-19,36-19.

Pohlia beringiensis A. J. Shaw — Czernyadjeva et al., 2020b. 2 records. 1: on bare soil in Luzula-dwarf willow spotty tundra, 33-19; on bare soil of baidjarakh slope, 36-19; gem.

P. cruda (Hedw.) Lindb. — Abramov, 1963. 15 records. 1, 3: in different types of tundra, herb-grass meadow, on bare soil of baidjarakh slope and bare loam; S+.

P. drummondii (Müll. Hal.) A. L. Andrews — 13 records. 1, 3: in different types of tundra, on spots of bare soil in herb-grass and Dupontia meadow, on bare soil on baidjarakh slopes; with Bartramia ithyphylla, Distichium inclinatum, Pohlia proligera, Psilopilum cavifolium, etc.; gem.

*P. elongata Hedw. var. greenii (Brid.) A. J. Shaw — 3 records. 3: in graminoid tundra, with Dicranum elongatum, 30te19; in cotton grass-Dupontia moss tundra, 33te19; in Luzula-dwarf shrub tundra, with Hymenoloma crispula, 44te19.

*P. filum (Schimp.) Mârtensson — 1 record. 2: in moss community near snowfield, with Pohlia wahlenbergii, 19-19, gem.

P. nutans (Hedw.) Lindb. — 11 records. 1, 3: in different types of tundra, Dupontia-moss and herb-Luzula meadows; S+.

*P. nutans subsp. schimperi (Müll. Hal.) Nyholm — 1 record. 1: on spot in hummocky spotty Luzula-graminoid-lichen-moss tundra, 25-19.

P. proligera (Kindb.) Lindb. et Broth. — Abramov, 1963; Stepanova, 1986. 8 records. 1, 3: on bare soil in Luzula-grass-lichen, hummocky dwarf shrub-graminoid, dwarf shrub-Luzula-moss--lichen and dwarf shrub-grass-moss tundra, on bare soil in herb-grass and graminoid-moss meadows and on baidjarakh slope; with Dicranella crispa, Pohlia cruda, P. drummondii, Psilopilum cavifolium, etc.; gem.

*P. wahlenbergii (F. Weber et D. Mohr) A. L. Andrews — 2 records. 1: in graminoid-moss meadow in runoff hollow, 13-19; 2: in moss community near snowfield, with Philonotis tomentella, 19-19.

Polytrichastrum alpinum (Hedw.) G. L. Sm. — 8 records. 1, 2, 3: in different types of tundra, herb-Luzula and Dupontia meadows, on bare soil of baidjarakh slope; S+.

P. fragile (Bryhn) Schljakov — 8 records. 1, 3: in different types of tundra, Dupontia-moss community, Dupontia-moss mire.

P. septentrionale (Brid.) E. I. Ivanova et al. — Abramov, 1963; Stepanova, 1986. 26 records. 1, 2, 3: in different types of tundra, herb-grass and Dupontia-moss meadows, on bare soil of baidjarakh slope, in Dupontia-sedge-moss and sedge-moss mires; S+.

Polytrichum hyperboreum R. Br. — 8 records. 1, 3: in different types of tundra; in herbgrass meadow, on bare soil of baidjarakh slope; S+.

P. juniperinum Hedw. — 6 records. 1: in hummocky dwarf shrub-graminoid, dwarf shrub-Du-pontia-sedge-moss and grass-moss tundra; 2: in swampy herb-Sphagnum-moss tundra; 3: in dwarf shrub-graminoid moss-lichen hummocky and graminoid-moss-lichen hummocky tundra; S+.

P. piliferum Hedw. — 1 record. 1: in Luzula-dwarf shrub tundra, 44te19.

P. strictum Brid. — Abramov, 1963. 2 records. 1: in hummocky spotty herb-moss tundra, 24-19; 3: in dwarf willow-graminoid-moss-lichen tundra, 41te19.

Pseudostereodon procerrimus (Molendo) M. Fleisch. — 1 record. 2: in dwarf willow-gram-inoid moss tundra, 15te19.

Psilopilum cavifolium (Wilson) I. Hagen — Abramov, 1963. 10 records. 1, 2, 3: on bare soil in dwarf shrub-grass-moss, Luzula-dwarf shrub and dwarf shrub-grass-moss tundra and on baidjarakh slopes, in forb-grass meadow, Racomitrium-lichen rubble tundra; with Dicranella sp., Distichium sp., Pohlia proligera, etc.; S+.

P. laevigatum (Wahlenb.) Lindb. — 1 record. 1: in grass-dwarf shrub-moss tundra, with Bryum sp., Dicranella sp., Pohlia proligera, 9-19, S+.

Racomitrium lanuginosum (Hedw.) Brid. — 14 records. 1, 2, 3: in different types of tundra and stony-gravelly area on the edge of terrace.

Rhizomnium andrewsianum (Steere) T. J. Kop. — Czernyadjeva et al., 2020a. 4 records. 1: on bare soil in dwarf shrub-grass-moss tundra, with Sarmentypnum sarmentosum, 8-19; in grass-Saxifraga-moss tundra, with Bryum sp., 9-19; in dwarf shrub-grass-moss hummocky tundra, 52te19; 3: in Dupontia moss tundra, 35te19.

R. gracile T. J. Kop. — Sofronova et al, 2021. 1 record. 1: in Dupontia-moss community, with Sarmentypnum sarmentosum, 35-19.

Saelania glaucescens (Hedw.) Broth. — 2 records. 1: in Luzula-dwarf willow spotty tundra, with Bartramia ithyphylla, Sanionia uncinata, Timmia austriaca, 33-19; 3: in forb-grass meadow, with Pohlia drummondii, 47te19.

Sanionia uncinata (Hedw.) Loeske — Abramov, 1963. 32 records. 1, 2, 3: in different types of tundra, Dupontia-moss, herb-Luzula and herb-grass meadows, on bank of stream in splits between baidjarakhs, on pebbles by the sea.

Sarmentypnum exannulatum (Schimp.) Hedenäs— 1 record. 2: in water in lake, 17-19.

S. sarmentosum (Wahlenb.) Tuom. et T. J. Kop. — Abramov, 1963. 14 records. 1, 2, 3: in different types of tundra, Dupontia, Equisetum-Dupontia and Dupontia-moss meadows, Dupon-tia-sedge-moss mire, on bank of stream in splits between baidjarakhs.

*Schistidium abrupticostatum (Bryhn) Ignatova et H. H. Blom — 1 record. 2: on soil in herb-dwarf shrub-moss tundra, 20-19, S+.

*S. grandirete H. H. Blom — 1 record. 2: on soil in herb-dwarf shrub-moss tundra, 20-19.

S. holmenianum Steere et Brassard — 1 record. 2: on soil in herb-dwarf shrub-moss tundra, 20-19.

S. papillosum Culm. — 2 records. 1: on stony-gravelly area in the edge of terrace, 10-19; on soil in Luzula-dwarf willow spotty tundra, 33-19; S+.

Scorpidium cossonii (Schimp.) Hedenäs — 7 records. 1: in graminoid-moss and Dupontia meadows, on bank of stream in splits between baidjarakhs; 2: in dwarf shrub-grass-moss spotty tundra, Dupontia meadow, in moss community near snowfield.

S. revolvens (Sw. ex anon.) Rubers — Abramov, 1963. 11 records. 1, 2, 3: in different types of tundra; in Dupontia-moss mire; in moss community near snowfield.

Seligeria polaris Berggr. — Abramov, 1963; Stepanova, 1986. 1 record. 2: in herb-dwarf shrub-moss tundra, 20-19, S+.

*Sphagnum beringiense A. J. Shaw et al. — 1 record. 2: in swampy herb-Sphagnum-moss tundra, 18-19, det. A. I. Maksimov.

S. concinnum (Berggr.) Flatberg — Sofronova et al, 2021. 5 records. 1: in grass-dwarf shrub-moss, hummocky dwarf shrub-graminoid tundra; 2: in swampy herb-Sphagnum-moss tundra; 3: in Dupontia tundra and Dupontia-sedge-moss mire.

S. orientale L. I. Savicz — 1 record. 2: in swampy herb-Sphagnum-moss tundra, 18-19, det. A. I. Maksimov.

S. squarrosum Crome — 2 records. 1: in grass-dwarf shrub-moss tundra, 9-19; 2: in swampy herb-Sphagnum-moss tundra, 18-19.

Splachnum vasculosum Hedw. — Abramov, 1963; Stepanova, 1986. 2 records. 1: in dwarf shrub-grass-moss tundra, with Ceratodon purpureus, 8-19; in hummocky dwarf shrub-gra-minoid tundra, 34-19.

*Stereodon holmenii (Ando) Ignatov et Ignatova — 2 records. 1: in Luzula-dwarf willow spotty tundra, 33-19.

Syntrichia ruralis (Hedw.) F. Weber et D. Mohr — 4 records. 1: on bare loam, 23-19; in dwarf shrub-Luzula spotty tundra, 33-19; 3: in Luzula-dwarf shrub tundra, 44te19; in herb-grass meadow, 46te19.

*Tayloria cf. tenuis (Dicks. ex With.) Schimp. — 1 record. 1: on spot in graminoid tundra, 8te19, S+.

Tetraplodon mnioides (Hedw.) Bruch et Schimp. (Fig. 6: 2) — 2 records. 1: on bare soil in dwarf willow-grass-moss tundra, with Rhizomnium andrewsianum, 8-19; in hummocky spotty Luzula-graminoid-lichen-moss tundra, 25-19; S+.

Timmia austriaca Hedw. — 29 records. 1, 2, 3: in different types of tundra, herb-grass and herb-Luzula meadows, sedge-moss mire, on bare soil in baidjarakh slopes; with Bartramia ithyphylla, Flexitrichum flexicaule, Pohlia cruda, etc.

T. norvegica J. E. Zetterst. — 5 records. 1: in grass-Saxifraga-moss tundra, graminoid-moss and Dupontia meadows, on bare soil of baidjarakh slope, on bare loam; with Distichium sp., Bryum sp., etc.

T. sibirica Lindb. et Arnell — Czernyadjeva et al, 2020a. 2 records. 1: in graminoid-moss meadow in runoff hollow, with Drepanocladus arcticus, Philonotis tomentella, 13-19; in Dupontia meadow, 32-19.

Tomentypnum involutum (Limpr.) Hedenäs et Ignatov — 17 records. 1, 2, 3: in different types of tundra, Dupontia and Dupontia-moss meadows, Dupontia-sedge-moss mire.

*Tortella arctica (Arnell) Crundw. et Nyholm — 4 records. 2: in herb-dwarf shrub-moss tundra, with Myurella julacea, 20-19; in dwarf shrub-grass hummocky tundra, 12te19; in dwarf willow-grass spotty tundra, 13te19; in dwarf willow-grass-moss spotty tundra, 22te19.

*Tortula cernua (Huebener) Lindb. — 2 records. 1: on bare loam, with Distichium sp., Dicranella sp., 23-19; 2: in crack of polygonal tundra, 16-19; S+.

*T. muralis Hedw. var. aestiva Hedw. — 3 records. 1: on bare loam, with Distichium sp., Bryum sp., 23-19; on bare soil of baidjarakh slopes, 30-19,36-19; S+.

Warnstorfia fluitans (Hedw.) Loeske — 1 record. 1: in hollow in hummocky dwarf shrub-graminoid tundra, 34-19.

*W. procera (Renauld et Arnell) Tuom. — 1 record. 1: in Dupontia-moss community, 35-19.

*W. pseudostraminea (Müll. Hal.) Tuom. et T. J. Kop. — 1 record. 1: in hollow in hummocky dwarf shrub-graminoid tundra, 34-19.

Excluded taxa

Andreaea rupestris Hedw. (Abramov, 1963) reidentified as A. papillosa by Czernyadjeva.

Calliergon giganteum (Schimp.) Kindb. (Abramov, 1963) reidentified as C. giganteum var. sibiricum by Czernyadjeva.

Dicranum congestum Brid. (Abramov, 1963) reidentified as D. acutifolium by Czernyadjeva.

Oncophorus virens var. serratus (Bruch et Schimp.) Braithw. (Abramov, 1963) reidentified as O. integerrimus by O. M. Afonina.

Oncophorus wahlenbergii Brid. (Abramov, 1963) reidentified as O. elongatus by O. M. Afonina.

Sphagnum fimbriatum Wilson (Abramov, 1963) reidentified as S. concinnum by Czernyadjeva.

Stereodon callichroum (Brid.) Braithw. (Abramov, 1963) reidentified as S. holmenii by O. M. Afonina.

Tomentypnum nitens (Hedw.) Loeske (Abramov, 1963) reidentified as T. involutum by Czernyadjeva.

Division Marchantiophyta — Liverworts

Aneura pinguis (L.) Dumort. — 2 records. 2: on soil in swampy herb-Sphagnum-moss tundra, 18-19; on soil in Dupontia tundra, 35te19, with young coelocaules.

Anthelia juratzkana (Limpr.)Trevis. — 21 records. 1, 2, 3: on bare often clayey soil and loamy spots in different types of tundra, herb-grass, and herb-Luzula meadows, stony-gravelly area at the edge of terrace, on bare soil of baidjarakh slope; per., mature capsules.

*A. juratzkana f. elongata Jorg. — 1 record. 3: on wet clayey soil in Luzula-graminoid moss tundra, 30te19.

Arnellia fennica (Gottsche) Lindb. — 1 record. 1: on soil in stony-gravelly area at the edge of terrace, with Anthelia juratzkana, Blepharostoma trichophyllum var. brevirete, Marchantia romanica, Mesoptychia heterocolpos var. arctica, Odontoschisma macounii, etc., 10-19.

Barbilophozia barbata (Schmidel ex Schreb.) Loeske — Abramov, 1963. 3 records. 2: on wet clayey soil in dwarf willow-graminoid tundra, 12te19, mod. biloba; in dwarf shrub-grass-moss rubble tundra, 25te19; 3: on wet clayey soil in Luzula-graminoid-lichen-moss tundra, 39te19.

Blepharostoma trichophyllum (L.) Dumort. var. brevirete Bryhn et Kaal. — 23 records. 1, 2, 3: in swampy herb-Sphagnum-moss, hummocky spotty herb-moss, hummocky spotty graminoid-lichen-moss, hummocky dwarf shrub-graminoid, hummocky grass-willow, Racomitrium-lichen rubble, Luzula-graminoid moss, cotton grass-Dupontia moss, dwarf willow-sedge-Dupontia-moss tundra, Dupontia meadow, and on soil in stony-gravelly area at the edge of terrace.

*Cephalozia ambigua C. Massal. — 5 records. 1: in hummocky spotty herb-moss tundra, 24-19; in hummocky spotty Luzula-graminoid-lichen-moss tundra, among Bartramia ithyphylla, with Anthelia juratzkana, Trilophozia quinquedentata f. gracilis, etc., 25-19; 3: in Racomitrium-lichen rubble tundra, 26te19; 3: in Luzula-graminoid-moss-lichen hummocky tundra, 40te19.

C. bicuspidata (L.) Dumort. — 6 records. 1: in hummocky spotty herb-moss, dwarf willow-sedge-Dupontia-moss, dwarf willow-grass-moss hummocky tundras and on stream bank between baidjarakhs; 3: in herb-grass meadow.

C. pleniceps (Austin) Lindb. — 1 record. 3: in cotton grass-Dupontia moss tundra, with Sphenolobus minutus, Blepharostoma trichophyllum var. brevirete, Cryptocolea imbricata, Pla-giochila arctica, 33te19.

*C. hamatiloba Steph. — 1 record. 1: in Dupontia-moss mire, among Polytrichastrum fragile, Cinclidium subrotundum, Sphagnum sp., Oncophorus elongatus, Scorpidium revolvens, with Sca-pania paludicola, Cephalozia varians, 56te19.

Cephaloziella divaricata (Sm.) Schiffn. — 3 records. 1: in dwarf willow-Dupontia-Luzu-la-moss tundra, 54te19; 3: in Luzula-graminoid moss tundra, 30-19 (cf.), herb-grass meadow, 46te19.

*C. grimsulana (J. B. Jack ex Gottsche et Rabenh.) Lacout. — 4 records. 1: on soil in stony-gravelly area at the edge of terrace, 10-19; in hummocky spotty Luzula-graminoid-li-chen-moss tundra, 25-19; 2: on soil in Equisetum-Dupontia meadow, 22-19; in dwarf wil-low-graminoid tundra, 12te19.

C. rubella (Nees) Schiffn. — 2 records. 1: on loamy soil in hummocky spotty herb-moss tundra, 24-19, per.; on loamy spot in dwarf willow-grass hummocky tundra, 11te19, cf.

C. varians (Gottsche) Steph. — 45 records, more often in 1 and 3, rare in 2. Mostly in various moss tundras, occasionally in graminoid tundras and mire communities with Dupontia, on bare soil; rare with perianths and gemmae.

*Cryptocolea imbricata R. M. Schust. — 2 records. 2: on soil in swampy herb-Sphagnum-moss tundra among Blepharostoma trichophyllum var. brevirete, 18-19; 3; on soil in cotton grass-Dupontia moss tundra, 33te19.

Gymnomitrion corallioides Nees — 6 records. 1, 2, 3: usually on bare soil, rare among mosses in spotty, hummocky, herb-moss tundras.

Jungermannia polaris Lindb. — 6 records. 1, 2: on bare loamy soil in different communities, on baidjarakh slope; per., mature capsules.

Lophozia excisa (Dicks.) Dumort. — 12 records. Mostly in 1 and 3, once in 2. In graminoid and moss tundras and on baidjarakh slope, often on bare soil; per., gem.

*L. jurensis Meyl. ex Müll. Frib. — 4 records. 1: in herb-grass-moss tundra, 8-19; on loamy spot in graminoid tundra, 8te19, cf.; 3: in dwarf shrub-Luzula-moss-lichen tundra, 45te19, gem.

L. longiflora (Nees) Schiffn. — 2 records. 3: Luzula-dwarf willow-moss-lichen tundra, 45te19, cf., gem.; in Luzula-graminoid-moss-lichen hummocky tundra, 40te19, per.

L. murmanica Kaal. sensu Damsholt 2013 (= L. heteromorpha R. M. Schust.) — 3 records. 1: in swampy herb-Sphagnum-moss tundra, with Lophozia polaris, 18-19, and in Dupontia meadow among Hymenoloma crispulum with Blepharostoma trichophyllum var. brevirete, Orthocaulis quadrilobus, etc., 32-19, gem.

*L. pellucida R. M. Schust. var. pellucida — 3 records. 1: on soil in stony-gravelly area at the edge of terrace, with Anthelia juratzkana, Blepharostoma trichophyllum var. brevirete, Arnellia fennica, Marchantia romanica, Mesoptychia heterocolpos var. arctica, Odontoschisma macounii, etc., 10-19, mod. succulenta; on bare soil of baidjarakh slope, with Orthocaulis quadrilobus, etc. 30-19, gem., per.; 3: in Luzula-dwarf willow-moss-lichen tundra, 45te19, cf., mod. parvire-tis-gemmipara, cells 25-30 ^m, gemmae green to brownish, 25-37 ^m.

*L. pellucida var. minor R. M. Schust. — 1 record. 1: in hummocky spotty Luzula-graminoid-lichen-moss tundra, among Bartramia ithyphylla, 25-19, no gemmae.

*L. perssonii H. Buch et S.W. Arnell — 5 records. 1: on bare loam, 23-19; on bare soil of baidjarakh slope, with Aloina brevirostris, Encalypta rhaptocarpa, Jungermannia polaris, Lophozia pellucida, Trilophozia quinquedentata, 30-19 (mod. densifolia-colorata, parvifolia- and laxifolia-viridis), 36-19 (mod. parvifolia-densifolia); in dwarf willow-Dupontia moss tundra, with L. wenzelii s. l., Sphenolobus minutus, etc., 55te19; usually with gemmae and in 55te19 with perianths, in 36-19 abundant. Protected in Russia (Krasnaya ..., 2008).

L. polaris (R. M. Schust.) R. M. Schust. et Damsh. [= L. alpestris auct. non (Schleich.) Evans. — Zhukova, 1976, 1982] — 14 records. 1 (most records), 2, 3: swampy herb-Sphagnum-moss tundra, graminoid-moss tundras, Dupontia-sedge-moss mire, gem., male (18-19) and female (30te19) plants.

*Lophozia cf. svalbardensis Konstant. et al. — 4 records. 1: on bare soil of baidjarakh slope, with L. perssonii, L. pellucida, Jungermannia polaris, Scapania obcordata, etc., 30-19; on bare soil in spotty Luzula-dwarf willow tundra, with Lophozia polaris, L. excisa, etc., 33-19; 3: on soil of herb-grass meadows, with Nardia geoscyphus, etc., 46te19, 47te19; gem., per.

Distinct from L. svalbardensis in not fleshy habit, perianths mouth at most with 1(2)-celled teeth, with marginal cells slightly thick-walled and orientated along the margin and locally forming crenulate projections, perianth cells with acute trigones, mostly unistratose leaves and rounded to polygonal stellate green to violet and purplish 1-2-celled gemmae.

*L. savicziae Schljakov — 2 records. 2: in swampy herb-Sphagnum-moss tundra, among Blepharostoma trichophyllum var. brevirete, 18-19, cf.; 3: in grass-forb-Luzula-moss-lichen nival tundra, 43te19, gem.

L. wenzelii (Nees) Steph. s. l. — 18 records. 1 and 3 (most records), 2: in graminoid, herb-moss, and various graminoid-moss tundras, gem.

*Marchantia romanica (Radian) D. G. Long et al. (= Bucegia romanica Radian) — 1 record. 1: on soil in stony-gravelly area at the edge of terrace, with Anthelia juratzkana, Arnellia fennica, Blepharostoma trichophyllum var. brevirete, Mesoptychia heterocolpos var. arctica, Odontoschisma macounii, etc., 10-19, young female receptacles.

Mesoptychia badensis (Gottsche ex Rabenh.) L. Söderstr. et Vana var. — 1 record. 1: separate shoots on bare soil in fissure of polygonal tundra, 16-19, mod. laxifolia-fusca: cuticle of the stem striolate, dorsal cortical cells ca. 25-30 ^m wide, with rudimentary underleaves.

*M. badensis var. apiculata (R. M. Schust.) Potemkin — 1 record. 1: separate shoots on bare soil of baidjarakh slope, among Bryoerythrophyllum recurvirostrum with Lophozia perssonii, 30-19, mod. angustifolia-colorata.

M. collaris (Nees) L. Söderstr. et Vana — 3 records. 2: in dwarf willow-graminoid tundra, with Cephaloziella grimsulana, Scapania brevicaulis, 12te19; in dwarf willow-graminoid moss tundra, 15te19; in dwarf willow-grass-moss rubble tundra, 25te19.

*M. heterocolpos (Thed. ex Hartm.) L. Söderstr. et Vana var. arctica (S.W. Arnell) L. Söderstr. et Vana — 2 records. 1: on soil in stony-gravelly area at the edge of terrace, with calciphilous hepatics, 10-19, gem.; 2: in herb-dwarf shrub-moss spotty tundra, with Anthelia juratzkana, Gymnomitrion corallioides, 20-19, mod. lepto- vel subpachyderma-viridis vel fusca.

M. sahlbergii (Lindb. et Arnell) A. Evans — 1 record. 2: among mosses in swampy herb-Sphagnum-moss tundra, 18-19.

*Nardia geoscyphus (De Not.) Lindb. — 20 records. 1 (most records), 3: on bare soil in various types of moss tundras, on baidjarakhs slopes, on moss-herb meadows; per., mature capsules.

Odontoschisma macounii (Austin) Underw. — 1 record. 1: on soil in stony-gravelly area at the edge of terrace, with calciphilous hepatics, 10-19, impoverished plants.

Orthocaulis binsteadii (Kaal.) H. Buch — 1 record. 1: in dwarf willow-sedge-Dupontia-moss tundra, with Sphenolobus minutus, Cephalozia bicuspidata, etc., 53te19.

*O. hyperboreus (R. M. Schust.) Konstant. — 1 record. 1: in dwarf willow-grass-forb hummocky tundra, with Ptilidium ciliare, Trilophozia quinquedentata, Cephalozia varians, 10te19, mod. parvifolia, marginal cells 25-25 ^m long, oil bodies 5-8 per cell.

*O. hyperboreus f. paradoxa (R. M. Schust.) Potemkin — 1 record. 1: in dwarf willow-grass spotty tundra, with Trilophozia quinquedentata, Cephalozia varians, Plagiochila arctica, 13te19, oil bodies (3)5(6) per cell, leaf surface papillose at leaf base only, cells with bulging trigones.

O. quadrilobus (Lindb.) H. Buch — 5 records. 1, 2: in Dupontia meadows, dwarf willow-grass-moss and dwarf willow-graminoid spotty tundras, in the latter communities (14te19, 22-24te19) phenotypes transitional to 'var. collenchymatica' occur, with marginal cells 1517 ^m wide or larger, with bulging trigones, oil bodies (3)5 per cell about 5 ^m, coarsely granulate, spherical, cuticle papillose.

*O. quadrilobus var. glareosus (Jorg.) S. W. Arnell — 1 record. 1: on bare soil on baidjarakh slope among Mnium cf. lycopodioides, 36-19.

Plagiochila arctica Bryhn et Kaal. — 5 records. 1: in dwarf willow-grass-moss hummocky tundra, 52te19; 2: in dwarf willow-graminoid tundra, 12te19; dwarf willow-grass spotty tundra, 13Te3: dwarf willow-grass moss hummocky tundra, 36te19; Luzula-graminoid-moss-lichen hummocky tundra, 40te19.

*Prasanthus suecicus (Gottsche) Lindb. — 2 records. 1: on bare soil in hummocky spotty herb-moss tundra, with mature capsules, 24-19.

*Pseudotritomaria heterophylla (R. M. Schust.) Konstant. et Vilnet (= Trilophozia heterophylla R. M. Schust.) — 3 records. 1: on spots and among mosses in hummocky spotty herb-moss tundra, 24-19, mod. colorata with bleached leaf margins, acute to nearly rounded lobes, and mod. viridis among mosses; on bare loam with Conostomum tetragonum and on spots in hummocky spotty Luzula-graminoid-lichen-moss tundra, 25-19, mod. fusca et viridis-mesoderma, mostly without vinaceous leaf bases and often with blunt lobes, with perianths and

gemmae. The phenotypes with blunt lobes should be attributed to Pseudotritomaria heterophyl-la f. anomala (Potemkin) Potemkin, comb. nov. (Basionym: Trilophozia heterophylla f. anomala Potemkin, 1990, Botanicheskii zhurnal 75: 1746. f. 1: 13-21).

Ptilidium ciliare (L.) Hampe — Abramov, 1963. 7 records. 1, 3: in various types of moss hummocky tundra, often with Dupontia, once in Dupontia-sedge-moss mire.

Scapania brevicaulis Taylor (S. degenii Schiffn. ex Müll. Frib. phenotype) — 1 record. 2: in dwarf willow-graminoid tundra, with Cephaloziella grimsulana, Mesoptychia alpestris, 12te19, with preserved oil-bodies, 2-5 per cell, occluding most of the cell lumen, no gemmae seen.

S. curta (Mart.) Dumort. — 2 records. 3: in Luzula-dwarf willow-moss-lichen tundra, 45te19, gem., male plants, cf.; in herb-grass meadow, 46te19, gem.

*S. curta var. grandiretis R. M. Schust. — 1 record. 1: on bare loam among Dicranella su-bulata, 23-19, gem. The plants represent mod. immarginata, which was described by Potemkin (1999).

S. gymnostomophila Kaal. — 1 record. 2: in dwarf willow-graminoid moss tundra, 15te19, gem.

S. irrigua (Nees) Nees — 1 record. 2: in dwarf willow-grass-moss rubble tundra, 25te19, mod. meso- vel pachyderma-fusca-egemmipara.

*Scapania cf. ligulifolia R. M. Schust. — 2 record. 3: on soil in Luzula-dwarf willow-moss-lichen tundra among Polytrichum hyperboreum and Polytrichastrum alpinum, 44te19; in Luzu-la-dwarf willow-moss hummocky tundra, 48te19; gem.

Despite both specimens are similar in habit to S. ligulifolia, oil bodies have not been preserved in leaf cells despite recent collection. Brownish gemmae being variable in shape and size occasionally have a reddish tinge. They are rounded 3-4-gonous to rhomboid and oval, 17-23 to 30-37 ^m long. Similar plants might be reported earlier for the New Siberian Islands as S. calcicola and were listed with a question mark and supposed to be S. ligulifolia by Konstanti-nova and Potemkin (1996).

*S. mucronata H. Buch subsp. praetervisa (Meyl.) R. M. Schust. — 4 records. 1: among Dicranum on bare soil of spots in hummocky spotty Luzula-graminoid-lichen-moss tundra, 25-19; on bare soil among Polytrichastrum septentrionale in Luzula-dwarf willow spotty tundra, 33-19; on bare soil among Pohlia proligera on baidjarakh slope, 36-19; 3: in forb-grass meadow, 46te19; gem.

S. obcordata (Berggr.) S. W. Arnell — 26 records. 1 (most records): on bare soil in various types of tundra and baidjarakh slopes; 3: on soil in forb-grass meadow; usually with gemmae.

S. paludicola Loeske et Müll. Frib. — 1 record. 1: among Polytrichastrum fragile, Cinclidium subrotundum, Sphagnum sp., Oncophorus elongatus, Scorpidium revolvens, in cotton grass-moss mire, 56te19.

S. scandica (Arnell et H. Buch) Macvicar —1 record. 3: on old rotten stem of Polytrichum sp. var. curta in Dupontia-sedge-moss mire, 37te19, mod. diplophylloidea-serrata.

*S. zemliae S. W. Arnell — 1 record. 1: on bare loam among Dicranella subulata, 23-19, gem.

*Schistochilopsis elegans (R. M. Schust.) Konstant. — 1 record. 1: on soil in stony-gravelly area at the edge of terrace, with calciphilous hepatics, 10-19, gem.

S. incisa (Schrad.) Konstant. — 3 records. 1: in dwarf willow-Dupontia moss tundra, 55te19; 2: in swampy herb-Sphagnum-moss tundra, 18-19, mod. laxifolia; 3: in graminoid-dwarf willow-lichen-moss hummocky tundra, 42te19; gem.

*Solenostoma hyalinum (Lyell) Mitt. — 2 records: 3: on soil in Luzula-dwarf willow-moss-lichen tundra, 45te19, mod.parvifolia-parviretis-leptoderma-subcolorata (fusca); in Luzula-dwarf willow-moss hummocky tundra, 48te19.

*Solenostoma pusillum (C. E. O. Jensen) Steph. (= Jungermannia gracillima Sm. subsp. arctica Damsh.) — 1 record. 1: on spots in hummocky spotty herb-moss tundra, 24-19.

S. subellipticum (Lindb. ex Heeg) R. M. Schust. [= Plectocolea obovata (Nees) Mitt. var. minor (Carrington) Schljakov] — 15 records. 1: on bare soil in spotty herb-moss and graminoid tundras, on bare soil and loam of baidjarakh slopes; per., mature capsules.

*S. sphaerocarpum (Hook.) Steph. var. nanum (Nees ex Flot.) Müll. Frib. — 1 record. 1: on bare soil in Luzula-dwarf willow spotty tundra, 33-19.

Sphenolobus minutus (Schreb. ex Cranz) Berggr. — Abramov, 1963. 29 records, the most common species. 1 and 3: among mosses and on bare soil in various types of moss tundra; 2: in Racomitrium-lichen rubble tundra only.

Trilophozia quinquedentata (Huds.) Bakalin [= Trilophozia quinquedentata (Huds.) H. Buch] — Abramov, 1963. 24 records. 1, 2, 3: among mosses and on bare soil in various types of dwarf willow tundra and moss tundra.

T. quinquedentata f. gracilis (R. M. Schust.) Konstant. — 20 records. 1, 3: On bare soil and among mosses in various types of moss tundra and graminoid communities.

Discussion

The study of our collection resulted in the identification of 140 species and 1 variety of mosses and 58 species, 4 varieties, and 2 forms of liverworts. Of these, 119 species and 1 variety of mosses and 54 species, 4 varieties, and 2 forms of liverworts are new to Stolbovoy Island. Among them, 40 species and 2 varieties of mosses were found for the first time for the New Siberian Islands. Schistidium grandirete, Cephalozia hama-tiloba, Lophozia cf. svalbardensis, Scapania cf. ligulifolia are reported for the first time for Yakutia and Bryoerythrophyllum rubrum, Rhizomnium gracile, Sphagnum concinnum were revealed for the first time for Yakutia in Stolbovoy Island and published earlier. Ten moss species have already been reported in the «New records» papers (Czernyadjeva et al., 2020 a, 2020b; Ellis et al., 2020, 2021; Sofronova et al., 2021). For comparison, for Kotelny and Bolshoy Lyakhovsky islands, which are the most investigated of the New Siberian Islands, 129 and 64 species of mosses are known, respectively. Thus, taking into account our data, 245 species of mosses and 88 species of liverworts are known for the New Siberian Islands.

Fourteen species of mosses are widespread on the island, where they represent an important component of the vegetation cover. They are Aulacomnium turgidum, Dicra-num elongatum, D. laevidens, Drepanocladus arcticus, Flexitrichum flexicaule, Hylocomi-um splendens, Oncophorus demetrii, Orthothecium chryseon, Pohlia cruda, Polytrichas-trum septentrionale, Racomitrium lanuginosum, Sanionia uncinata, Timmia austriaca, Tomentypnum involutum. Bartramia ithyphylla, Conostomum tetragonum, Distichium in-clinatum, Myurella julacea, Pohlia nutans are common, although occur in admixture to other species and small amount. Less common but abundant in some sites species are Dicranum acutifolium, D. majus, D. spadiceum, Flexitrichum gracile, Kiaeria glacialis, Oncophorus elongatus, Polytrichastrum alpinum, P. fragile, Polytrichum hyperboreum, Scorpidium cossonii. Dicranella crispa, Pohlia drummondii, P. proligera, Psilopilum cavi-folium are frequent on bare soil. Typical species of waterlogged habitats are Philonotis tomentella, Sarmentypnum sarmentosum, Scorpidium revolvens.

Taking into account that liverworts were not specially gathered on the island, only their frequency in collections may be evaluated. The most frequently collected species are Anthelia juratzkana (21 records), Cephaloziella varians (45), Lophozia polaris (14), L. wenzelii s. l. (18), Nardia geoscyphus (20), Scapania obcordata (26), Sphenolobus mi-nutus (29), Solenostoma subellipticum (15), Trilophozia quinquedentata (44 records). However, despite Sphenolobus minutus was gathered 29 times, it was found only once in the northern tip of the island, in the vicinity of the Cape Skalisty, whereas Scapania obcordata was gathered mostly in the northwest coast in the vicinity of the polar station, and not found in the northern tip of the island. The species composition and frequency mentioned above recalls much with that known for the liverworts of Bolshoy Lyakhovsky Island (Zhukova, 1982). However, Blepharostoma trichophyllum, Orthocaulis quadrilobus, Plagiochila arctica, Ptilidium ciliare, Scapania irrigua are much less frequent in Stolbovoy Island whereas rather frequent in Bolshoy Lyakhovsky Island. Diplophyllum taxifolium (Wahlenb.) Dumort., Marsupella arctica (Berggr.) Bryhn et Kaal., Orthocaulis kunzeanus (Huebener) H. Buch, Radula prolifera Arnell, Scapania simmonsii Bryhn et Kaal., Tet-ralophozia setiformis (Ehrh.) Schljakov (as Chandonanthus setiformis (Ehrh.) Lindb.), Trilophozia scitula (Taylor) Jorg. were not found in our collections from Stolbovoy Island.

Several rare and new species for Yakutia are phytogeographically remarkable.

Bryoerythrophyllum rubrum was first collected on the territory of Yakutia (Ellis et al., 2021). In Russia, its isolated finds are known in the Caucasus, Taimyr Peninsula, Severnaya Zemlya Archipelago, Wrangel, and Sakhalin islands (Afonina, 2002, 2004; Fedosov, Ignatova, 2008; Bakalin et al., 2012). This species was found mostly in the mountainous regions of central and southern Europe (Fedosov, Ignatova, 2008; Hodgetts, Lockhart, 2020) and reported from China (Li et al., 2001). The find of B. rubrum in Stolbovoy Island is northernmost.

Sphagnum concinnum is an arctic circumpolar species. Until 2007, it was known in Russia from collections from Novaya Zemlya (Flatberg, 2007) only. Recently, S. concinnum was found in the Asian part of Russia in the Yamal-Nenets Autonomous District (Sofronova et al., 2019; Beldiman et al., 2020) and the Taimyr Peninsula (Fedosov et al., 2020). This record of S. concinnum is easternmost in Russia.

Sphagnum beringiense is an Asian-American species. In Russia, it occurs in the Yamal and Tazovsky peninsulas, in the eastern Taimyr Peninsula, Yakutia, and the Russian Far East from Chukotka to the Primorsky Territory (Maksimov et al., 2016). In North America, its distribution area is restricted to the western coastal area (Alaska, Washington, and British Columbia) (Shaw et al., 2008). The present location of S. beringiense is at the northern limit of its distribution.

Rhizomnium gracile is widespread in the north of North America. Until present, in Russia, it was known from the northern suboceanic territories, from in the northern Far East (Chukotka, Kamchatka, Commander Islands, Magadan Region) mostly, and a few finds in the Murmansk Region (Ignatov et al., 2018).

The find of Cephalozia hamatiloba in Stolbovoy Island is westernmost in Russia. Its main range is in Pacific Asia and Appalachian Mts. in eastern North America. More-

over, it is known from China, Bhutan, S. India, and Thailand (Potemkin, Sofronova, 2013). Considerable morphological variability of this species and the very broad range presumes its possible genetic diversification, which needs a special study.

Funaria arctica is a rare arctic species. This species was recorded in North America from Alaska, Nunavut, Yukon, Northwest Territories, Greenland (Miller, 2007); in Europe - from Svalbard and one site in Swedish Scandes mountains; in Russia, F. arctica is known from Asian Arctic - the Yamal Peninsula, Taimyr Peninsula, Anabar Plateau, Ust'-Lensky Reserve (Yakutia), Srednekolymsky District (Yakutia), Beringian Chu-kotka (Fedosov, Ignatova, 2005; Ivanova, 2009; Fedosov et al., 2011; Ignatov et al, 2017).

Obtained data on mosses and liverworts of Stolbovoy Island complement the knowledge of bryophyte flora of the New Siberian Islands and provide a comparable representation of the flora mosses and liverworts of Stolbovoy Island. Despite the limited time of collecting and only three visited localities in the northern part of the island its bryophyte flora now is comparable with that of much better investigated Kotelny and Bolshoy Lyakhovsky islands.

Acknowledgments

The authors are very grateful to O. M. Afonina, M. S. Ignatov, E. A. Ignatova. A. I. Maksimov for help in the identification of the problematic moss specimens. The work of I. V. Czernyadjeva and A. D. Potemkin was supported by Russian Foundation for Basic Research (grant 18-05-60093 Арктика) and it was carried out within the framework of the institutional research project (no. 121021600184-6) of the Koma-rov Botanical Institute of the Russian Academy of Sciences. I. V. Czernyadjeva and A. D. Potemkin thank Ministry of Science and Higher Education of Russia for support of CCU "Herbarium MBG RAS". The expedition to Stolbovoy Island was part of the 4th complex expedition 'The Novosibirskie Islands as an outpost of Russia in the Arctic. Polar expeditions of trailblazers, scientists and hunters' arranged by the Russian Geographical Society and the Academy of Sciences of the Republic of Sakha (Yakutia).

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