Novosti Sistematiki Nizshikh Rastenii 54(1): 211–241

Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 54(1): 211–241. 2020

The mosses of the Khulga River Basin (Subpolar Urals, Khanty-Mansi Autonomous Area — Yugra)

E. D. Lapshina1, E. A. Ignatova2, M. S. Ignatov2, 3, G. N. Ganasevich1, Yu. V. Skuchas4

1Yugra State University, Khanty-Mansiysk, Russia 2Lomonosov Moscow State University, Moscow, Russia 3N.V. Tsitsin Main Botanical Garden of the Russian Academy of Sciences, Moscow, Russia 4Museum of Nature and Man, Khanty-Mansiysk, Russia Corresponding author: E. D. Lapshina, [email protected]

Abstract. Result of an extensive moss study of earlier nearly unexplored areas of the northeastern part of the Subpolar Urals, namely Khulga River Basin is presented. An annotated list of mosses is compiled based on the identification of 1200 specimens collected in the Khulga River Basin (60.45–65.20°N, 60.42–62.20°E). The river starts at the border between the Polar and Subpolar Urals and flows down southwards along the eastern slope of the Ural Mountains to the confluence with the Severnaya Sosva River near the village Saranpaul (Berezovo District, Khanty-Mansi Au- tonomous Area — Yugra). The list includes 210 moss species, of which 56 species are new to the Khanty-Mansi Autonomous Area — Yugra. Five species are new to the Urals: Didymodon perobtusus, Lewinskya iwatsukii, Schistidium frisvollianum, Sphagnum mirum, Warnstorfia tundrae. Further 9 species were found in the Subpolar Urals for the first time: Dicranum bardunovii, D. laevidens, D. schljakovii, Encalypta pilifera, Flexitrichum gracile, Grimmia elatior, G. tergestina, Orthotrichum hyperboreum, Sphagnum annulatum. The data on occurrence of many moss species, which were previously known from single finds in the mountainous part of the area, is significantly expanded. Ecology, phytoceno- tic preference and distribution of the recorded species are discussed. Keywords: bryoflora, biogeography, ecology, distribution, phytocenology, Subpolar Urals, Russia.

Мхи бассейна р. Хулга (Приполярный Урал, Ханты-Мансийский автономный округ — Югра)

Е. Д. Лапшина1, Е. А. Игнатова2, М. С. Игнатов2, 3, Г. Н. Ганасевич1, Ю. В. Скучас4

1Югорский государственный университет, Ханты-Мансийск, Россия 2Московский государственный университет им. М. В. Ломоносова, Москва, Россия 3Главный ботанический сад им. Н. В. Цицина РАН, Москва, Россия 4Музей Природы и Человека, Ханты-Мансийск, Россия Автор для переписки: Е. Д. Лапшина, [email protected]

Резюме. Представлены результаты исследования мхов ранее почти не изученной севе- ро-восточной части Приполярного Урала, бассейна р. Хулга. Аннотированный список мхов со- ставлен на основе определения 1200 образцов, собранных в бассейне р. Хулга (60.45–65.20°N, 60.42–62.20°E), берущей начало от границы Полярного и Приполярного Урала и стекающей вдоль восточного склона Уральских гор к югу до впадения в р. Северная Сосьва в районе https://doi.org/10.31111/nsnr/2020.54.1.211 211 Lapshina et al. The mosses of the Khulga River Basin (Subpolar Urals)

пос. Саранпауль (Березовский р-н, Ханты-Мансийский автономный округ). Список включа- ет 210 видов, из них 56 видов мхов являются новыми для Ханты-Мансийского автономно- го округа. Приводится 5 видов новых для Урала: Didymodon perobtusus, Lewinskya iwatsukii, Schistidium frisvollianum, Sphagnum mirum, Warnstorﬁa tundrae. Еще 9 видов (Dicranum bardunovii, D. laevidens, D. schljakovii, Encalypta pilifera, Flexitrichum gracile, Grimmia elatior, G. tergestina, Orthotrichum hyperboreum, Sphagnum annulatum) впервые найдены на Приполярном Урале. Су- щественно расширены представления о частоте встречаемости многих видов мхов в горной ча- сти территории округа, известных ранее по единичным находкам. Обсуждаются особенности экологии, фитоценотической приуроченности и характер распространения видов мхов. Ключевые слова: бриофлора, биогеография, экология, распространение, фитоценология, Приполярный Урал, Россия.

The Subpolar Urals is the widest and most elevated part of the Ural Mountains, about 300 km in length from the headwaters of the Khulga River, from 65°40'N in the north up to 64°N in the south, including the foothill plains adjacent from the east and west (Gorchakovskiy, 1975). The first list of mosses of the Subpolar Urals was compiled by I. D. Kildyushevsky (1956). It includes 233 species with references to earlier published records by Pohle (1915), Ivanovskiy (1917), Gorodkov (1932), and Sochava (1930). Later, this list was supplemented by A. P. Dyachenko (1995) according to the literature data mainly on species identified during studies of the vegetation of the Subpolar Urals (Gorchakovs- kiy, 1955, 1958; Kuvaev, 1970, 1980; Nepomilueva et al., 1976; Nepomilueva, 1978). Thus, the total number of moss species known for the Subpolar Urals and adjacent plains was increased to 265. Though mosses of the Subpolar Urals were studied for nearly a century, knowledge on the moss species diversity, their habitats, frequency, and distribution patterns in the study area remains insufficient. Most of the former investigations of the Subpolar Urals directed at lowland plains and hilly foothills mainly in the southern and western parts of the territory, whereas data on mosses of the northeastern part of the Subpolar Urals in the Khul- ga River Basin are extremely poor. There was only one publication by B. N. Go- rodkov (1932) on the Polar Urals with a list of mosses, of which 59 species were collected in the upper reaches of the Khulga River at the northern boundary of the Subpolar Urals. In 2013–2015 we conducted studies of mosses and liverworts on the eastern slope of Subpolar Urals in the area of Mount Ner-Oika (1645 m) and 10 km to the south at the headwaters of the Puyva River (64°30'–64°33'N, 59°35'–59°38'E). The list of liverworts and partial data on mosses was published earlier (Konstantinova, Lapshi- na, 2014, 2017; Lapshina et al., 2015, 2016; Skuchas, Lapshina, 2018). The Khulga River Basin is located in the northeastern part of the Subpolar Urals, at a distance of about 120 km from the previously surveyed sites. The goal of this paper is to present a contribution on the moss flora of this nearly unexplored territory of the Polar and Subpolar Urals.

212 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 54(1): 211–241. 2020

Study area The Khulga River (the old name: Lyapin) starts to the south of the Polar Circle on the border between the Polar and Subpolar Urals and flows down along the eastern slope of the Ural Mountains to the south to its confluence with the Severnaya Sosva River (a left tributary of the Ob River) near the village Saranpaul. Administratively, the basin of the Khulga River occupies the extreme northwestern part of the Beloyarsk District and the entire Khanty-Mansi Autonomous Area. It is located between the mountain ranges of the northern part of the Subpolar Urals and the boggy areas of the West Siberian Plain within the Northern Taiga subzone. On mountain tops, tundra vegetation and forest-tundra open woodland extend far to the south. Thereby three main mountain vegetation belts are presented in the study area: forest belt (up to 350–400 m), forest-tundra belt (450–500 m), dominated by Sibe- rian pine (Pinus sibirica) and birch (Betula pubescens) open woodlands with an admixture of larch (Larix sibirica), alternating with thickets of alder (Duschekia fruticosa) and dwarf birch (Betula nana) communities — “yerniks”, and a mountain tundra belt, represented on the highest peaks up to 600–750 m (names of vascular plants follow Cherepanov, 1995). The study area is located at the southern boundary of the distribution of local permafrost with active current thermokarst processes. Most of the territory at the Middle and Lower Khulga River is plain or in some places gently undulating plain with absolute elevations of 40–90 m a. s. l. The Khulga River bed is stony, 60–80 m wide with 4–6 m deep pits and flat shallows. At a low water level, numerous islands and silty-stony shallows occur. The river valley is from several hundred meters to 3–4 km wide, the well-drained sites occupied by river valley forests are alternated with wet lowlands. Flat terraces are covered with transitional fens and raised bogs, as well as open woodland on shallow peat with the stunted tree layer of spruce (Picea obovata) and Siberian pine (Pinus sibirica). In places where river valley is getting narrow the slopes of hill and ridges covered by mountainous northern taiga forests and open woodlands approach the river. The proximity of mountain ranges, composed of metamorphic and intrusive rocks with close occurrence and frequent outcrops of carbonate rocks, results in a variety of rocky substrates and nutrient-rich groundwaters, which provide a wide variety of types of peatlands and mire vegetation. The vegetation of the Khulga River Basin is formed by flood plain and mountain forests, boggy open woodlands, and raised bogs, transitional and minerotrophic fens, thickets of subalpine shrubs (yerniks, willow shrubs), mountain tundras, plant communities on stony substrates, including carbonate rocks, vegetation of silty-gravelly riverbanks and coastal outcrops. Material and methods The moss specimens were collected during the geobotanical survey of the main vegetation types. Special attention was paid to rocky and stony outcrops, coasts of rivers

213 Lapshina et al. The mosses of the Khulga River Basin (Subpolar Urals) and lakes. Well recognizable species were collected from a few sites only, whereas in other cases they were only noted in the relevés. A total of 194 sites were surveyed, for which 178 geobotanical relevés were made, the full list of bryophyte species being identiﬁed. For the rest of the sites, such as the banks of rivers, streams, rocky outcrops, moss specimens were collected without vegetation relevés. On the map all collecting sites were combined into 30 localities within 4 key areas (Fig. 1).

I — Upper reaches of Khulga River near mouth of Tykotlova River (right tributary of Khulga River). Valley of Khulga River: 1 — right bank, 2 km below mouth of Tykotlova River, 65.252°N, 62.170°–62.177°E, 61 m above sea level; 2 — opposite the mouth of Tykotlova River, 65.253°– 65.269°N, 62.177°–62.184°E, 61 m a. s. l.; 3 — left bank, 2 km above mouth of Tykotlova River, 65.278°–65.286°N, 62.189°–62.207°E, 66 m a. s. l. Tykotlova peatland area between the valleys of the Khulga and Tykotlova Rivers: 4 — flat palsa-lake-hollow complex, 65.275°–65.285°N, 62.132°–62.147°E, 66 m a. s. l.; 5 — minerotrophic fen of nutrient-rich water supply in area of carbonate groundwater discharge, 65.253°–5.269°N, 62.177°–62.184°E, 68 m a. s. l.; 6 — frozen flat palsa bog, 65.291°–65.294°N, 62.146°–62.177°E, 68 m a. s. l.; 7 — groundwater discharge and swampy stream valley at base of mountain limestone spurs, 65.302°–65.303°N, 62.123°–62.131°E, 61 m a. s. l.; 8 — hanging bog at base of carbonate rock slope, 65.297°N, 62.128°E, 72 m a. s. l. Mid-mountain belt of the eastern slope of the Subpolar Urals: 9 — dry carbonate rocks of south-eastern exposure and carbonate outcrops in spruce-larch forest on north-eastern slope, 65.301°N, 62.119°E, 105 m a. s. l.; 10 — dry carbonate rocks on steep slope of southern exposure, 65.298°N, 62.085°E, 81 m a. s. l.; 11 — separate stone block on gentle north-eastern slope, 65.277°N, 62.111°E, 92 m a. s. l.; 12 — shrub-sedge tussock-moss fen in an intermontane basin 65.291°N, 62.109°–62.111°E, 71 m a. s. l.; 13 — minerotrophic fen of nutrient-rich groundwater supply at base of carbonate rock by lake, 65.298°N, 62.095°E, 75 m a. s. l.; 14 — shore of lake in intermountain basin, 65.302°N, 62.075°E, 75 m a. s. l.; 15 — valley of Tykotlova River at its lower reaches, 65.278°N, 62.101°E, 72 m a. s. l. II — Upper reaches of the Khulga River in the area of the Balbanty Lake. The valley of Khulga River: 16 — mesotrophic sedge-herb-Sphagnum fen on low terrace of Khulga River on right bank between Balbanty Lake and mouth of Tykotlova River 65.208°N, 62.140°–62.143°E, 54 m a. s. l.; 17 — left bank, 2 km east of Balbanty Lake, 65.150°–65.154°N, 62.108°–62.126°E, 52 m a. s. l. Mid-mountain belt of the eastern slope of the Subpolar Urals: 18 — lower part of western concave slope and swampy valley of Balbanshor Brook on eastern and northern shores of lake in forest belt, 65.161°–65.176°N, 62.026°–62.048°E, 81 m a. s. l.; 19 — rock outcrops on steep slope of deep intermountane small stream valley in upper part of forest belt, 65.201°N, 61.996°E, 487 m a. s. l.; 20 — gentle slopes and plateaus at top of mountain pass in tundra belt, 65.202°–65.210°N, 61.970°–61.986°E, 510 m a. s. l.; 21 — upper reaches of Balbanshor Brook under snowfield with rocky outcrops and small alpine mire in tundra belt, 65.216°–65.222°N, 61.922°–61.939°E, 608 m a. s. l.; 22 — valley of Khulga River, 1 km below mouth of Balban’yu River, 65.104°N, 62.216°–62.218°E, 46 m a. s. l. III — Valley of the Khulga River in its middle course: 23 — high palsa-hollow complex with frozen peat mounds and thermokarst subsidence of different ages on right bank of Khu- lga River 64.936°–64.937°N, 61.993°–61.999°E, 45 m a. s. l.; 24 — coastal silty-sand sediment

214 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 54(1): 211–241. 2020

Fig. 1. Collecting sites of mosses. 215 Lapshina et al. The mosses of the Khulga River Basin (Subpolar Urals) outcrop along left bank of river, 64.870°N, 62.034°E, 40 m a. s. l.; 25 — mesooligotrophic aapa mire1 complex with frozen palsa2 on left-bank terrace of Khulga River, 64.870°–64.871°N, 62.042°–62.046°E, 43 m a. s. l.; 26 — in bed of Khulga River on right bank, 64.659°N, 61.824°E, 31 m a. s. l.; 27 — left bank of Khulga River opposite the mouth of Khalmer’yu River, 64.558°N, 61.642°E, 34 m a. s. l. IV — Valley of the Khulga River at its lower reaches: 28 — left-bank terrace near the mouth of Man’ya River (right tributary of Khulga River) 64.461°N, 61.359°E, 24 m a. s. l.; 29 — mesooligotrophic fen with aapa complex in central part, 64.348° –64.349°N, 61.107°–61.150°E, 25 m a. s. l.; 30 — mesotrophic dwarf shrub-sedge-Sphagnum fen and aapa complex, 64.337°– 64.339°N, 61.066°–61.067°E, 24 m a. s. l.

In total about 1200 moss specimens were collected from July 7 to 18, 2018 and over 2500 identifications were made. All specimens are kept in the Biological Collection of the Yugra State University, Khanty-Mansiysk (YSU), selected samples of duplicates are in MW and MHA. To characterize the ecological conditions of phytocoenotic confinement of moss species, all the variety of habitats is combined in the following groups: Mountain and plain North taiga forests and open woodlands are common on drained, well-developed soils. Mixed dwarf shrub-green moss and dwarf shrub-lichen-green moss forests predominate. The tree layer is formed in different ra- tios by Picea obovata, Pinus sibirica, Betula pubescens and Larix sibirica. Large areas are also occupied by secondary, mainly young birch forests at different stages of post- fire restoration. In the ground cover Hylocomium splendens, Pleurozium schreberi, Di- cranum species dominate, often with some admixture of Polytrichum commune, Sphag- num girgensohnii, S. wulfianum. In mountain forests Sciuro-hypnum reflexum, S. curtum, Stereodon plicatulus sporadically occur. River valley forests develop on well-drained ridges along Khulga River and its large tributaries. These are the most productive tall dark coniferous and mixed herb and herb-green moss forests. The tree layer is formed by the same species as in described above zonal forests, often with well-defined alder undergrowth (Duschekia fruticosa) less often with Salix spp. and Sorbus sibirica. Moss cover varies from 10 to 80%, depending on the duration of flooding. Along with typical boreal forest species Climacium dendroides, Eurhynchiastrum pulchellum, Mnium lycopodioides, Plagiothe- cium spp., Pohlia cruda are very common in the moss layer. At the bases of trees Pylai- sia polyantha and Myrinia pulvinata were recorded. Dwarf shrub-sphagnum open spruce woodlands are common in flat ar - eas of river terraces and in the lower parts of slopes on a peat layer up to 50 cm thick. Picea obovata forms a light low tree layer 5–8 (10) m high, often with admixture of Pinus sibirica. In the lower layers Betula nana, Ledum palustre, Rubus chamaemorus, Oxycoccus microcarpus prevail. In the moss cover Sphagnum girgensohnii associated

1 Aapa mire is a conventional term used to designate mesooligotrophic ribbed fen (also called string-ﬂark mire complex or string-ﬂark mire). 2 Palsa is a mound of peat with a permafrost core, surrounded by seasonally melting fens.

216 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 54(1): 211–241. 2020 with S. angustifolium and Polytrichum commune dominate. In small abundance Aula- comnium palustre, Polytrichum strictum, Pleurozium schreberi and, less often, Sphagnum russowii were found. In conditions of a nutrient-richer groundwater supply at the base of mountain slopes, spruce woodlands are characterized by the presence of such moss species as Tomentypnum nitens, Sphagnum warnstorfii and Rhizomnium pseudopunctatum. In some cases, Sphagnum fuscum dominates in the moss layer. Mires occupy up to 30% of the Khulga River Basin in its lower and middle cours- es. In the upper reaches, where the river valley narrows and comes close to the eastern spurs of the Urals, the area of mires decreases, but their typological diversity increases. Raised bogs of atmospheric nutrition represented by low pine (Pinus sylvestris, P. sibirica)-dwarf shrub-Sphagnum communities, as well as flat palsa and flat palsa-lake complexes are widespread. In the moss cover with a dominant role of Sphagnum fuscum, participation of Pleurozium schreberi, Sphagnum angustifolium, S. capillifolium and fruticose lichens is notable. On low flat palsa S. lenense occurs along with Sphag- num fuscum. In thermokarst hollows and waterlogged areas of flat palsa complexes cotton grass (Eriophorum russeolum)-Sphagnum and sedge (Carex rotundata)-Sphagnum oligotrophic communities with Sphagnum balticum, S. lindbergii, S. jensenii, S. majus are developed with minor admixture of Polytrichum jensenii in moss layer. Transitional fens. In the early stages of paludification in thermokarst subsidence within high palsa-hollow complexes and on the low river terraces, if the hydro- logical connection to the river is still retained, sedge (Carex rostrata)-herb (Comarum palustre, Equisetum fluviatile)-Sphagnum and cotton grass (Eriophorum angustifolium, E. gracile)-sedge (Carex limosa)-Sphagnum communities dominated by Sphagnum riparium, S. obtusum are developed. Vast poor fens and aapa complexes in extended sec- tions of river terraces are the most typical of the study area. In aapa complexes, dwarf shrub (Betula nana, Chamaedaphne calyculata)-sedge (Carex lasiocarpa)-Sphagnum (S. divinum) ridges alternate with waterlogged hollows, where the moss cover is represented by single shoots and small patches of Sphagnum platyphyllum, S. subsecundum, Warnstorfia exannulata, W. fluitans. In relatively homogeneous peripheral areas, dwarf shrub-sedge (Carex rostrata, C. lasiocarpa, C chordorrhiza)-Sphagnum (S. angustifolium, S. girgensohnii, S. centrale) communities are rather common. Minerotrophic fens. On the right bank of the Khulga river, in its upper reaches, specific horsetail (Equisetum fluviatile)-sedge (Carex diandra, C. juncella)-brown moss (Cinclidium stygium, Bryum pseudotriquetrum, Scorpidium cossonii), shrubby (Salix phylicifolia, Betula nana)-sedge (Carex juncella)-brown moss (Scorpidium scorpioides) and dwarf birch (Betula nana)-Sphagnum (S. warnstorfii)-brown moss (Scorpidium revolvens) fens of rich-nutrient groundwater supply occur. Beside the abovementioned dominants in the moss cover of these fens Calliergon giganteum, C. richardsonii, Campyli- um stellatum, Fissidens osmundoides, Hamatocaulis vernicosus, Meesia triquetra, Paludella squarrosa and Tomentypnum nitens were found. In addition in places, where groundwater emerges at the base of mountain slopes composed of carbonate rocks, Loeskypnum badium, Warnstorfia exannulata, W. sarmentosa and W. tundrae were recorded.

217 Lapshina et al. The mosses of the Khulga River Basin (Subpolar Urals)

On gentle slopes in the mountain-tundra belt, there are small spots of shrub-sedge (Carex arctisibirica)-Sphagnum alpine mires dominated by Sphagnum warnstorfii and with abundance of liverworts (Schljakovianthus quadrilobus, Tritomaria quinquidenta- ta, Ptilidium ciliare, Schljakovia kunzeana) with admixture of Hylocomium splendens, Polytrichastrum alpinum, Flexitrichum flexicaule, Scorpidium cossonii. The spring-water supply indicators Cyrtomnium hymenophylloides and Philonotis tomentella are also common. Tundras replace subalpine open woodlands at 450–470 m a. s. l. Debris-lichen-green moss, dwarf shrub (Betula nana, Vaccinium vitis-idaea, V. uliginosum, Em- petrum hermaphroditum, Diapensia lapponica) and dwarf birch (Betula nana)-green moss tundras are most common. In ground layer, along with numerous lichens and liverworts, Aulacomnium turgidum, Dicranum spadiceum, D. flexicaule, D. undulatum, Hylocomium splendens, Pohlia nutans, Polytrichastrum alpinum are the most common. On protruding stones Andreaea rupestris, Racomitrium lanuginosum and Hymenoloma crispulum are common. Carbonate rocky outcrops within the forest belt are the richest habitats in terms of moss species diversity. Schistidium spp., Orthotrichum spp., Drepanium recurvatum, Grimmia spp., Leucodon sciuroides, Stereodon vaucheri, S. bambergeri, Tortella tortuosa occur on their surface. On ledges and between stones on humified fine earth, Encalypta spp., Flexitrichum flexicaule, Myurella spp., Plagiopus oederianus, Platydictia jungerman nioides and Timmia austriaca are common. In shaded wet niches Cyrtomni- um hymenophylloides, Distichium capillaceum and Neckera oligocarpa were recorded. Only at carbonate rock outcrops Amphidium mougeotii, Anomodon attenuatus, Didimo- don perobtusus, Hedwigia mollis, Oxystegus tenuirostris, Trichostomum crispulum, Ulota curvifolia and Zygodon sibiricus were found. Water and river bank habitats. On silty-gravel banks of the Khulga River and streams Bryum pallens, Calliergon cordifolium, Calliergonella lindbergii and Pohlia filum are common. In the middle course of the river in such habitats Atrichum tenellum and Philonotis caespitosa occur. Along the coast and on the protruding stones in the channels of small streams and rivers, on fine earth in the forest belt in conditions of periodic flooding Bryum turbinatum, B. pseudotriquetrum, Hygrohypnella ochracea, H. polare, Ochyraea alpestris, Philonotis tomentella, Pohlia wahlenbergii, Schistidium agassizi, S. papillosum, S. platyphyllum and Warnstorfia exannulata were found. In the mountain tundra belt along the banks of streams flowing from under snowfields, the species composition of mosses changes. Blindia acuta, Ochyraea duriuscula, Hymenolo- ma crispulum, Oncophorus virens, Schistidium rivulare, S. sordidum and Sciuro-hypnum glaciale are the most common in such habitats. Noteworthy that in the water of rivers and streams, on stones and submerged trunks, only two species of Fontinalis, F. antipyretica, F. hypnoides were found. Bare soil and eroded substrates. Dicranella crispa, Ditrichum cylindricum, Pohlia lescuriana and P. proligera are pioneers growing over sandy and sandy loam

218 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 54(1): 211–241. 2020 outcrops along the Khulga River banks. In such habitats Psilopilum cavifolium grows abundantly and often dominates. Species list The nomenclature of mosses follows mainly the “List of Mosses of Eastern Europe and North Asia” (Ignatov et al., 2006), taking into account some later changes (Tu- banova, 2010; Hessel et al., 2018; Ignatov et al., 2017, 2018). The liverworts nomenclature follows mainly Söderström et al. (2016) with some changes (Bakalin, 2016). The species are listed in alphabetical order. For each species its frequency is indi- cated and evaluated in accordance with the following scale: common (Cm, the species is registered at more than 15 sites), frequent (Fr, 7–15 sites), sporadic (Sp, 4–6 sites), rarely (Rar, 2–3 sites), a single find (Un). The numbers of key areas (Roman numerals in boldface) and localities (Arabic numerals), where the species was recorded are given in accordance with the map (Fig. 1). For common, frequent and sporadic species the generalized habitat characteristics are given. For rare species collected from 1–2 localities the labels are cited with reference to catalog number in the Biological Collection of Yugra State University (YSU) in square brackets or field number site in italic. Collector name is not mentioned for most specimens gathered by the first author, E. D. Lapshina. Geographical reference: Khanty-Mansi Autonomous Area, Berezovo District, basin of the Khulga River. The presence of sporophytes is marked by S+. Extended data on all specimens and their labels in Russian are provided in the electronic supplement3 to this article arranged according to the specimen numbers in the Biological Collection of Yugra State University (YSU). Asterisks before species name mean: * — new record for the Khanty-Mansi Auton- omous Area — Yugra; ** — new record for the Subpolar Urals, *** — new records for the entire Urals. Abietinella abietina (Hedw.) M. Fleisch. — Sp. I: 9, 10; II: 19–21. In well lighted places along rocky outcrops and in ground cover in forest belt and mountain tundra. Amblystegium serpens (Hedw.) Bruch, Schimp. et W. Gümbel — Un. I: 1, birch-spruce valley forest, at the base of tree trunk, 65.25283°N, 62.17799°E, 11 VII 2018, Filippov, YSU- MH-00088. S+. Amphidium mougeotii (Bruch, Schimp. et W. Gümbel) Schimp. — Rar. I: 11, stone block in larch-birch forest on gentle mountain slope on stone surface, 65.27757°N, 62.11185°E, 12 VII 2018, YSU-MH-00356; II: 21, rocky outcrops in Betula nana-green moss tundra, on surface of stones, on fine earth in mixture with Hygrohypnum norvegicum, Oxystegus tenuirostris, Isoptery- giopsis pulchella, 65.22070°N, 61.92725°E, 14 VII 2018, YSU-MH-00601. The species is widespread in the mountain systems of the Holarctic, but in the Urals it is known from a few sites (Ignatov et al., 2006). Andreaea rupestris Hedw. — Rar. I: 11; II: 19–20. On separate stones and rocks in mountain tundra, rarely forest belts. S+.

3 Electronic supplement is available at the end of the article page on the journal website (https://doi.org/10.31111/nsnr/2020.54.1.211).

219 Lapshina et al. The mosses of the Khulga River Basin (Subpolar Urals)

*Anomodon attenuatus (Hedw.) Hübener — Un. I: 10, rocky outcrop of limestone, on shaded surface of stones, small admixture among Drepanocladus aduncus, Barbula convoluta, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00319. A rare nemoral species outside the broad-leaved forest zone, known in the Northern Urals from isolated finds (Ignatova et al., 1996). Atrichum tenellum (Röhl.) Bruch, Schimp. et W. Gümbel — Un. III: 27, silty-gravel river sandbank, in willow-grass-green moss (Calliergonella lindbergii, Calliergon cordifolium) community, 64.55854°N, 61.64242°E, 8 VII 2018, YSU-MH-00009. Aulacomnium palustre (Hedw.) Schwägr. — Cm. I: 1–3, 5–8, 12–14; II: 16–18, 20–22; III: 23, 25, 27; IV: 28–30. In spruce dwarf shrub-cloudberry-Sphagnum bogs and woodlands with Picea obovata and Pinus sibirica, on ridges of transitional fens, in raised bogs, and in swampy forests. A. turgidum (Wahlenb.) Schwägr. — Un. II: 20, dwarf birch-stony tundra, in niches between stones, 65.21719°N, 61.93727°E, 14 VII 2018, YSU-MH-00512. *Barbula convoluta Hedw. — Un. I: 10, on outcrops of carbonate rocks, in niches between stones, on fine earth, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00310. Bartramia ithyphylla Brid. — Un. II: 21, on grassy slope along stream bank under snowfield, 65.22291°N, 61.92264°E, 14.VII.2018, YSU-MH-0054. S+. Blindia acuta (Hedw.) Bruch, Schimp. et W. Gümbel — Un. II: 21, on shore and stones in stream bed under snowfield, on concave stones, 65.22291°N, 61.92264°E, 14 VII 2018, YSU- MH-00581. S+, immature. Brachythecium capillaceum (F. Weber et D. Mohr) Giacom. [= B. rotaeanum De Not] — Un. I: 3, birch-spruce-shrub (Duscheckia fruticosa, Salix spp.)-herb river valley forest, at base of tree, in pure mats and with Pylaisia polyantha, 65.28656°N, 62.18911°E, 11 VII 2018, YSU-MH-00185, YSU-MH-00186. B. mildeanum (Schimp.) Schimp. — Sp. I: 1, 3, 8; II: 18. In wet birch-herb-Calamagrostis river valley forests, in dwarf birch-cloudberry-Sphagnum open spruce woodlands, and minerotrophic fens. B. salebrosum (F. Weber et D. Mohr) Bruch, Schimp. et W. Gümbel — Un. II: 21, shrubby tundra, in niches between stones, single shoots in mat of Pseudoleskeella rupestris, 65.21958°N, 61.93000°E, 14 VII 2018, YSU-MH-00515. *B. turgidum (Hartm.) Kindb. — Rar. I: 10, carbonate rocks, between stones on organic and fine earth material, in mixture with Campyliadelphus chrysophyllus, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00285; II: 21, shrub (Salix, Betula nana)-sedge (Carex arctisibirica)-Sphagnum alpine mire, single shoots in pure mat of Orthocaulis quadrilobus, 65.22070°N, 61.92725°E, 14 VII 2018, 75/7-18. Bryoerythrophyllum recurvirostrum (Hedw.) P. C. Chen — Sp: I: 9–11, outcrops of carbonate and basic rocks in forest belt, at ledges and in niches between stones on fine earth. Bryum capillare Hedw. — Un. II: 21, rocky outcrops in yernik tundra, in niches between stones, on fine earth in mixture with Isopterygiopsis pulchella, Distichum capillaceum, 65.22070°N, 61.92725°E, 14 VII 2018, YSU-MH-00600. S+. B. creberrimum Taylor — Un. I: 10, outcrops of carbonate rocks, in niches, between stones on fine earth, in pure mats, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00307. S+. *B. intermedium (Brid.) Blandow — Rar. I: 10, outcrops of carbonate rocks, in niches between stones, in pure mats and in mixture with Encalypta rhaptocarpa, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00316. S+; I: 14, on peaty shore of lake at the base of carbonate mountains, single shoot in pure mat of Distichium capillaceum, 65.30276°N, 62.07579°E, 12 VII 2018, YSU-MH-00324. A rare arcto-boreal species, known in the Urals from single finds. 220 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 54(1): 211–241. 2020

B. moravicum Podp. — Un. I: 10, carbonate rock outcrop, in shaded niches between stones, in pure mat, in mixture with Bryoerythrophyllum recurvirostrum, and as single admixture between Myurella sibirica, Pohlia cruda, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00303. B. pallens Sw. ex anon. — Rar. I: 1, along bank of stream, 65.25149°N, 62.17349°E, 11 VII 2018, Lapshina 39/7-18; III: 27, silty-gravel sandbank in willow-grass-green moss community, noticeable admixture among Pohlia filum, Calliergonella lindbergii, Scapania irrigua, 64.55854°N, 61.64242°E, 8 VII 2018, YSU-MH-00006. B. pseudotriquetrum (Hedw.) P. Gärtn., B. Mey. et Schreb. — Cm: I: 1, 3, 5, 7, 12, 15; II: 21; III: 27, in minerotrophic sedge-brown moss fens, in swampy and wet floodplain forests, along banks of rivers and streams. S+. B. turbinatum (Hedw.) Turner — Rar. I: 1, along bank of stream, in pure mats, 65.25149°N, 62.17349°E, 11 VII 2018, YSU-MH-00252. S+; II: 22, on flooded gravelly river bank, 65.10409°N, 62.21820°E, 18 VII 2018, YSU-MH-00032. S+. B. weigelii Spreng. — Un. II: 21, along bank of stream under snowfield, in pure mats, 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00571. Callicladium haldanianum (Grev.) H. A. Crum — Rar. I: 3, flooded birch-spruce-shrub (Dus- checkia fruticosa, Salix spp.)-herb river valley forest, at base of trees, 65.28656°N, 62.18911°E, 11 VII 2018, YSU-MH-00652. Calliergon cordifolium (Hedw.) Kindb. — Sp. II: 16, 18; III: 23, 27, along banks of rivers and streams, in minerotrophic sedge-hypnum fens, in willow thickets. C. giganteum (Schimp.) Kindb. — Fr. I: 1, 5, 7, 12, on minerotrophic horsetail-sedge-brown moss and shrub (Betula nana)-tussock-sedge-brown moss rich fens. C. richardsonii (Mitt.) Kindb. — Rar. I: 5, in sedge (Carex rostrata)-horsetail-brown moss fen, small admixture among Hamatocaulis vernicosus, 65.29494°N, 62.14051°E, 10 VII 2018, Filippov, YSU-MH-00037; I: 12, shrub (Salix philicifolia, Betula nana)-tussock sedge (Carex juncella) fen in intermountain depression, singlе shoots in pure mat of Scorpidium cossonii, 65.29348°N, 62.14253°E, 10 VII 2018, YSU-MH-00650. Calliergonella lindbergii (Mitt.) Hedenäs — Fr. I: 1, 10, 14–15; II: 17, 22; III: 27. At river and stream banks, in willow-grass-green moss communities on silty-gravel soil, in river valley forests. Campyliadelphus chrysophyllus (Brid.) R. S. Chopra — Un. I: 9, outcrops of carbonate rocks, in niches between stones, on fine earth, in mixture with Brachythecium turgidum, Drepa- nium recurvatum, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00286. *Campylidium calcareum (Crundw. et Nyholm) Ochyra — Un. I: 14, on dry peat shore of lake at the base of carbonate mountains, in mixture with Lophocolea minor, Mesoptychia colla- ris and single plants of Bryum cf. intermedium, Barbilophozia barbata, 65.30276°N, 62.07579°E, 12 VII 2018, YSU-MH-00325. S+. A rare species outside Europe, known in the Urals from single finds. Campylidium sommerfeltii (Myrin) Ochyra — Rar. I: 5, in horsetail (Equisetum fluviatile) fen with rare oppressed trees, at the base of spruce and on decayed stumps, admixture in pure mat of Mesoptychia heterocolpos. 65.29546°N, 62.13940°E, 10 VII 2018, YSU-MH-00092. S+; II: 18, in birch-grass (Calamagrostis, Equisetum sylvaticum) river valley forest, on peaty soil, in pure mat. 65.16155°N, 62.04456°E, 14 VII 2018, YSU-MH-00388. S+. Campylium stellatum (Hedw.) C. E. O. Jensen — Fr. I: 5, 7, 12–13. In shrub (Betula nana, Salix)-tussock sedge-brown moss and horsetail-brown moss rich fens . S+. Ceratodon purpureus (Hedw.) Brid. — Sp. I: 2, 5; II: 19, 21, III: 24, 27. Occurs in wide variety of habitats; most abundant along dry edges of river bank outcrops. S+.

221 Lapshina et al. The mosses of the Khulga River Basin (Subpolar Urals)

*Cinclidium stygium Sw. — Sp: I: 5, 7, 12. In tussock-sedge-brown moss and horsetail-brown moss fens with a mineal-rich groundwater supply. S+. Climacium dendroides (Hedw.) F. Weber et D. Mohr — Sp. I: 3; II: 17. In birch-spruce-grass floodplain forests and along river banks. Cynodontium cf. strumiferum (Hedw.) Lindb. — Un. II: 19, rock outcrops in birch forest, on fine earth in nishes between stones, with single admixture of Hedwigia mollis, Ceratodon purpureus, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00472. S+. Cyrtomnium hymenophylloides (Hübener) T. J. Kop. — Sp. I: 9, 10, 11; II: 21. In moist shaded niches at outcrops of carbonate rocks in mountain forest and tundra belts. Once collected in alpine shrub (Salix, Betula nana)-sedge (Carex arctisibirica)-Sphagnum mire. Dicranella crispa (Hedw.) Schimp. — Un. III: 24, on coastal exposure, on sandy substrate, single shoot among Pohlia lescuriana, Dicranella varia, Psilopilum cavifolium, 64.87081°N, 62.03472°E, 16 VII 2018, YSU-MH-00618. S+. D. varia (Hedw.) Schimp. — Un. III: 24, coastal outcrop, on sandy soil, in mixture with Psilopilum cavifolium, Ditrichum cylindricum, 64.87081°N, 62.03472°E, 16 VII 2018, YSU-MH-00619. S+. **Dicranum bardunovii Tubanova et Ignatova — Un. II: 17, in mixed larch-spruce- birch-Pinus sibirica-dwarf shrub-green-moss river valley forest, at base of larch, 65.15105°N, 67.11533°E, 13 VII 2018, YSU-MH-00373. Until recently, the species was known only in the Asian part of Russia east of the Yenisei River (Tubanova, Ignatova, 2011) and from Vishersky Nature Reserve on the western slope of the North Urals (Ignatova et al., 2019). As a result of the revision of herbarium specimens, this species was identified at two localities in the Nadym and Taz regions of the Yamalo-Nenets Autonomous Area in the north of Western Siberia (Pisa- renko et al., 2017). Therefore, its find in the Subpolar Urals is not surprising. D. bonjeanii De Not. — Rar. I: 5, in Betula nana-Sphagnum open spruce woodland, single shoots among Sphagnum fuscum, Tomentypnum nitens, Rhizomnium pseudopunctatum, 65.29751°N, 62.12825°E, 10 VII 2018, Filippov, YSU-MH-00022. D. flexicaule Brid. — Rar. II: 21, in dwarf shrub tundra, with single shoots of Polytrichas- trum alpinum, 65.21958°N, 61.93000°E, 14 VII 2018, YSU-MH-00517. S+; ibidem, among stones in Betula nana-green moss tundra, in pure mats, YSU-MH-00509. D. fragilifolium Lindb. — Fr. I: 1, 3, II: 17–18, 20. In river valley shrub-herb-green moss forests, in boggy dwarf shrub-Sphagnum open woodlands with Pinus sibirica, less often in mountain dwarf shrub tundras. D. fuscescens Turner — Rar. II: 17–18; III: 23. In mountain and valley forests, in boggy woodlands. **D. laevidens R. S. Williams — Rar. I: 5, in boggy spruce-Betula nana-Rubus chamaemorus-Sphagnum (S. fuscum) open woodlands at the base of mountain slope, with Tritomaria quinquedentata, 65.29749°N, 62.12861°E, 10 VII 2018, YSU-MH-00102; IV: 29, in meso-oligotrophic aapa mire complex, on dwarf shrub (Andromeda polifolia)-Sphagnum (S. divinum) ridge with rare low pine trees, in mixture with Sphagnum russowii, 64.34894°N, 61.10882°E, 17 VII 2018, YSU-MH-00651. A hypoarctic species widespread in the Arctic and Subarctic of Eurasia. It was previously found in the northern flat part of the Khanty-Mansi Autonomous Area (Lap- shina et al., 2018), and in Vishersky Nature Reserve on the western slope of the North Urals (Ignatova et al., 2019). D. majus Turner — Un. II: 18, in birch-grass (Calamagrostis purpurea, Equisetum sylvaticum) mixed forest, pure mats on peaty soil, 65.16155°N, 62.04456°E, 14 VII 2018, YSU-MH-00395. D. montanum Hedw. — Sp. I: 1; II: 17–19. In the upland and river valley mixed forests, in green moss-lichen Pinus sibirica-birch open forests, at base of trees and on decayed wood.

222 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 54(1): 211–241. 2020

**D. schljakovii Ignatova et Tubanova — Un. II: 21, at rock outcrops in yernik tundra, without admixture of other species, 65.22070°N, 61.92725°E, 14 VII 2018, YSU-MH-00599. Apparently, not a rare species in the north of Western Siberia (Pisarenko et al., 2017), was previously noted in the highlands of the Southern Urals, in the Caucasus, Khibiny and Altai Mts. For the Subpolar Urals, the species is identified for the first time. D. spadiceum J. E. Zetterst. — Sp. I: 11; II: 20–21. In rocky dwarf-shrub tundra, in Betula nana-sedge (Carex arctisibirica)-Sphagnum mountain mire, on stony outcrops in forest belt. D. undulatum Schrad. ex Brid. — Fr. I: 2–3, 8–10; II: 18, 20, 22; III: 23. In Pinus sibirica-spruce dwarf shrub-Sphagnum boggy woodlands, less often in larch-Pinus sibirica-spruce dwarf shrub-green moss forests and rock outcrops. ***Didymodon perobtusus Broth. — Un. I: 9, on dry southeast-faced slope at outcrops of carbonate rocks, in dense pure mat. 65.30118°N, 62.11997°E, 10 VII 2018, YSU-MH-00135. This is the first and up to now the only find of the species in the Urals. Earlier, the species has been observed only in the mountains of Southern Siberia and Kamchatka (Ignatov et al., 2006). Distichium capillaceum (Hedw.) Bruch, Schimp. et W. Gümbel — Sp. I: 9–10, 14; II: 21. Along outcrops of carbonate and basic rocks in forest belt and mountain tundra, in niches on fine earth, in pure mats and mixed with other species. S+. Ditrichum cylindricum (Hedw.) Grout — Un. III: 24, shore exposure on sandy soil, single shoots among Dicranella varia, Psilopilum cavifolium, Ceratodon purpureus. 64.87081°N, 62.03472°E, 16 VII 2018, YSU-MH-00619. S+. Drepanium recurvatum (Lindb. et Arnell) G. Roth — Rar. I: 9, on dry southeast slope, at outcrops of carbonate rocks, in pure mats and in mixture with Stereodon vaucherii, Encalypta cf. streptocarpa; I: 10, at outcrops of carbonate rocks, in pure mats on the surface of stones. Drepanocladus aduncus (Hedw.) Warnst. — Rar. I: 10, at outcrops of carbonate rocks, in moist shaded niches, on stones, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00319; III: 23, in high palsa-mire complex, in sedge community of thermokarst depression, with admixture Bryum pseudotriquetrum, 64.93691°N, 61.99780°E, 16 VII 2018, Filippov, YSU-MH-00204. D. polygamus (Bruch, Schimp. et W. Gümbel) Hedenäs — Rar. I: 10, at outcrops of carbonate rocks, in moist shaded niches, on stones, in admixture with Calliergonella lindbergii, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00317; III: 23, in high palsa-mire complex, in sedge community of thermokarst depression, in ground cover with admixture of Sanionia uncinata, 65.17568°N, 62.02801°E, 14 VII 2018, Filippov, YSU-MH-00167. **Encalypta pilifera Funck — Un. I: 10, at stone block on gentle mountain slope, on fine earth on surface of stone, without admixtures, 65.27757°N, 62.11185°E, 12 VII 2018, YSU-MH-00336. S+. Previously, the species was known from isolated places in the Southern and Middle Urals. For the northern part of the Urals is identified for the first time (Ignatov et al., 2017). S+. E. rhaptocarpa Schwägr. — Rar. I: 9–11. At outcrops of carbonate and basic rocks in forest belt, in small pure mats, often together with or among other species. S+. *Encalypta cf. streptocarpa Hedw. — Rar. I: 9–11. At outcrops of carbonate and basic rocks in forest belt, in pure mats and in mixture with Encalypta rhaptocarpa, Drepanium recurvatum, Bryoerythrophyllum recurvirostrum, Cyrtomnium hymenophylloides, Platydictya jungermannioides. All specimens are collected in a sterile condition. We presumably attribute them to this species, since we found E. streptocarpa with capsules on the adjacent territory of the Subpolar Urals at the upper reaches of the Puyva River in 2015. This species was also reported for the adjacent Komi region at the western border of the Khanth-Mansi Autonomous Area at the upper reaches of the Levma River (Kildyushevskiy, 1956). Eurhynchiastrum pulchellum (Hedw.) Ignatov et Huttunen — Sp. I: 2–3; II: 17. In birch- spruce- and mixed alder-grass-green moss river valley forests, on bare soil and at bases of tree stems.

223 Lapshina et al. The mosses of the Khulga River Basin (Subpolar Urals)

Fissidens bryoides Hedw. — Rar. I: 3, 10. In birch-spruce-shrub (Duschekia fruticosa, Salix spp.)-grass river valley forests, at outcrops of carbonate rocks, in moist niches between stones, everywhere as minor admixture. F. osmundoides Hedw. — Rar. I: 13, in Betula nana-sedge-moss fen at the base of carbonate cliff near lake, in mixture with Mesoptychia rutheana, Campylium stellatum, Scorpidium revolvens, 65.29878°N, 62.09535°E, 12 VII 2018, YSU-MH-00269; I: 12, in Betula nana-tussock- sedge (Carex juncella) mire in the intermontane basin, small admixture between Campylium stellatum, 65.29137°N, 62.11055°E, 12 VII 2018, YSU-MH-00259. Flexitrichum flexicaule (Schwägr.) Ignatov et Fedosov [Ditrichum flexicaule (Schwägr.) Hampe] — Sp. I: 9; II: 20–21. In dwarf shrub tundras, at outcrops of rocks of different composition in forest belt and on shrub-sedge-Sphagnum (S. warnstorfii) alpine mire. S+. **F. gracile (Mitt.) Ignatov et Fedosov — Un. I: 9, on southeast slope, at outcrops of carbonate rocks, in pure mats in crevices in fine earth, 65.30118°N, 62.11997°E, 10 VII 2018, YSU- MH-00113. Since this species was not previously distinguished from F. flexicaule its distribution is poorly understood. Occurs sporadically in the Arctic, Subarctic and in mountains of more southern regions (Ignatov et al., 2017; Ignatova et al., 2019). For the Subpolar Urals F. gracile is recorded for the first time. Fontinalis antipyretica Hedw. — Un. I: 7, in bed of small river flowing from lake, on dead wood, 65.30334°N, 62.13155°E, 10 VII 2018, YSU-MH-00168. F. hypnoides Hartm. — Un. III: 26, at middle course of the Khulga River, on stones in water, 65.30334°N, 61.82424°E, 17 VII 2018, YSU-MH-00628. S+. ** Grimmia elatior Bruch ex Bals.-Criv. et De Not. — Un. II: 21, at rock outcrops in dwarf birch tundra, on stones in pure mats, 65.22070°N, 61.92725°E, 14 VII 2018, YSU-MH-00594. The species is widespread in mountainous regions, including the Southern, Middle and North- ern Urals. The find in the Subpolar Urals is quite predictable. * G. longirostris Hook. — Rar. I: 11; II: 19–20. At rock outcrops in the upper part of forest and mountain tundra belts, on rocks, in pure mats and with Leucodon sciuroides. **G. tergestina Tomm. ex Bruch, Schimp. et W. Gümbel — Rar. I: 9, on southeast slope, at outcrops of carbonate rocks, in small pure mats, 65.30118°N, 62.11997°E, 10 VII 2018, YSU- MH-00131; I: 10, at rocky outcrop of carbonate, on surface of stones, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00308. Previously, it was listed for the Southern and Northern Urals (Ignatova, Muñoz, 2004). Hamatocaulis vernicosus (Mitt.) Hedenäs — Sp. I: 5; IV: 29–30. On sedge-brown moss and Betula nana-tussock-sedge-brown moss rich fens, less often in hollows of aapa mires. Hedwigia mollis (Hedw.) P. Beauv. — Un. II: 19, rocky outcrops on steep mountain slope in young birch forest, on stones in pure mats, 65.18130°N, 62.99642°E, 14 VII 2018, YSU- MH-00460. According to the recent revision of the Hedwigia ciliata (Hedw.) P. Beauv complex the specimens collected on the eastern slope of the Subpolar Urals were attributed to Hedwigia mollis (Ignatova et al., 2016). S+. Helodium blandowii (F. Weber et D. Mohr) Warnst. — Rar. II: 18; III: 23. In swampy thickets of willow, in sedge community, in boggy dwarf shrub-Sphagnum larch-spruce open forest. Hygrohypnella ochracea (Turner ex Wilson) Ignatov et Ignatova — Rar. I: 1; II: 19, 21. In streams and rivers, on stones. Mostly mountainous species, also known from single locations in the lowlands of the Khanty-Mansi Autonomous Area (Lapshina, Pisarenko, 2013). *H. polaris (Lindb.) Ignatov et Ignatova — Un. I: 1, in the channel of stream, on stones, 65.25149°N, 62.17349°E, 11 VII 2018, YSU-MH-00234. This species was previously known from single locations for the Northern and Western slopes of the Subpolar Urals (D’yachenko, 1997; Zheleznova, 1994).

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Hylocomiadelphus triquetrus (Hedw.) Ochyra et Stebel [≡ Rhytidiadelphus triquetrus (Hedw.) Warnst.] — Rar. I: 3 II: 17. In birch-spruce shrub-grass-green moss valley forests, in forest floor. Hylocomium splendens (Hedw.) Bruch, Schimp. et W. Gümbel — Fr. I: 1–3; II: 17–19; III: 23; IV: 28. In river valley and mountain mixed and dark coniferous dwarf shrub-green moss and Vaccinium vitis-idaea-green moss forests. Hylocomiadelphus triquetrus (Hedw.) Ochyra & Stebel — Rar. I: 3 II: 17. In birch-spruce shrub-grass-green moss valley forests, in forest floor. *Hymenoloma crispulum (Hedw.) Ochyra — Sp. II: 19, 21. At rock outcrops, on protruding stones along banks of streams in forest and tundra belts. S+. Hypnum cupressiforme Hedw. — Rar. I: 11, stone block on gentle mountain slope, on fine earth on surface stones, in admixture among Radula complanata, 65.27757°N, 62.11185°E, 12 VII 2018, YSU-MH-00347; II: 19, rock outcrops on steep slope in young birch forest, in pure mats and with Barbilophozia barbata, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00444. *Isopterygiopsis muelleriana (Schimp.) Z. Iwats. — Un. II: 19, rock outcrops on steep mountain slope in upper part of forest belt, in niches on fine earth, without admixture of other species, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00477. A rare hypoarctic species, previously cited on the basis of unspecified data for the Polar Urals (Afonina, Czernyadjeva, 1995). Our finds confirmed the presence of this species in the Urals. I. pulchella (Hedw.) Iwats. — Fr. I: 3, 11; II: 19, 21. In birch and birch-dark coniferous river valley forests on bare soil, on rock outcrops in dwarf birch tundra, in niches on fine earth. S+. *Kiaeria blyttii (Bruch, Schimp. et W. Gümbel) Broth. — Un. II: 20, in dwarf shrub (Em- petrum-Vaccinium-Diapenisia) tundra, in niches between stones, without admixture of other species, 65.20243°N, 61.98634°E, 14 VII 2018, YSU-MH-00486. *Kiaeria starkei (F. Weber et D. Mohr) I. Hagen — Un. II: 21, on slope along bank of stream, under snowfield, with minor admixture of Sanionia uncinata, 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00529. Leptobryum pyriforme (Hedw.) Wils. — Un. I: 5, in spruce-horsetail fen, on decayed wood of stump, single shoots in pure mat of Mesoptychia heterocolpos, 65.29546°N, 62.13940°E, 10 VII 2018, YSU-MH-00093. S+. *Lescuraea radicosa (Mitt.) Moenk. — Un. II: 21, on grassy slope along bank of stream under snowfield, in mixture with Hymenoloma crispulum, 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00562. *L. saxicola (Bruch, Schimp. et W. Gümbel) Molendo — Rar. II: 19, rock outcrops on steep slope in young birch forest, on ledges of stones on fine earth, in mixture with Barbilophozia hatcheri, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00417; II: 21, on grassy slope along bank of stream under snowfield, in pure mats and as admixture among Sciuro-hypnum glaciale, Mnium blyttii, Saccobasis polita, 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00545. *Leucodon sciuroides (Hedw.) Schwägr. — Rar. I: 9, 11; II: 19. At outcrops of carbonate and basic rocks, mainly in pure mats. ***Lewinskya iwatsukii (Ignatov) F. Lara, Garilleti et Goffinet [≡ Orthotrichum iwatsukii Ig- natov] — Un. II: 19, rock outcrops on steep slope in young birch forest, on stones, single shoots in pure mat of Schistidium pulchrum, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00421. Ear- lier, the species was observed in Russia only in the Far East, in the mountains of Siberia and the Caucasus. The find in the Urals significantly expands the knowledge about its distribution. S+. *Loeskypnum badium (Hartm.) H. K. G. Paul — Un. II: 18, groundwater spread within larch-spruce-shrub-Sphagnum open woodland, in mixture with Paludella squarrosa, 65.17609°N, 62.02684°E, 14 VII 2018, YSU-MH-00414. 225 Lapshina et al. The mosses of the Khulga River Basin (Subpolar Urals)

Meesia triquetra (Jolycl.) Ångstr. — Sp. I: 5, 12, 13. In Betula nana-tussock sedge-brown moss and sedge-brown moss rich fens. M. uliginosa Hedw. — Un. II: 21, on grassy slope along bank of stream under snowfield, noticeable admixture among Pseudolophozia sudetica, Mesoptychia heterocolpos, Anthelia ju- ratzkana, Brepharostoma trichophyllum, Schljakovianthus quadrilobus, 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00535. *Mnium blyttii Bruch, Schimp. et W. Gümbel — Un. II: 21: on grassy slope along bank of stream under snowfields, noticeable admixture between Sciuro-hypnum glaciale, 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00553. Previously, the species was known from single finds on the northern and western slopes of the Subpolar Urals (Zheleznova, 1994; D’yachenko, 1997). M. lycopodioides Schwägr. — Sp. I: 1, 3. In periodically flooded birch-spruce and mixed shrub-grass and herb-green-moss river valley forests, on bare soil. Predominantly mountainous species, it is very common in floodplain forests. M. marginatum (Dicks.) P. Beauv. — Rar. I: 1, 9, 11. At outcrops of carbonate and basic rocks, in niches between stones, on fine earth, as admixture among Orthothecium strictum, Dis- tichium capillaceum, Myurella tenerrima, M. julacea, Platydictya jungermannioides, Bryoerythro- phyllum recurvirostrum. Found also on bare soil in birch-spruce river valley forest, mixed with Lophocolea minor, Eurhynchiastrum pulchellum, Barbilophozia barbata. M. spinulosum Bruch, Schimp. et W. Gümbel — Un. II: 19, rock outcrops on steep mountain slope in young birch forest, in shaded niche on fine earth, in pure mats and with Isopterygi- opsis pulchella, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00434. M. stellare Hedw. — Rar. I: 3, 5. In birch-spruce herb-green moss river valley forests, on soil. Once met in spruce-Sphagnum fen, on decayed wood of spruce stump, single shoots in pure mat of Mesoptichia heterocolpos. M. thomsonii Schimp. — Rar. I: 10, 14; II: 21. At outcrops of carbonate rocks in the forest and mountain tundra belts, in moist shaded niches between stones; along shore of mountain lake, on peaty soil, single shoots in pure mat of Distichium capillaceum. Myrinia pulvinata (Wahlenb.) Schimp. — Rar. I: 3, in small herb-green moss flooded birch- spruce forest, at base of trunks, insignificant admixture among Pylaisia polyantha, 65.27844°N, 62.20731°E, 11 VII 2018, YSU-MH-00215; ibidem, in Pinus sibirica-birch-spruce valley forest, single admixture in pure mat of Sciuro-hypnum curtum, 65.27852°N, 62.20757°E, 11 VII 2018, Filippov 12F-12. * Myurella julacea (Schwägr.) Bruch, Schimp. et W. Gümbel — Ra. I: 9–10. At outcrops of carbonate rocks in forest belt, by fine earth, sometimes forms pure mats, often grows in mixture with other species. M. sibirica (Müll. Hal.) Reimers — Un. I: 10, on outcrops of carbonate rocks, in moist shaded niches, on fine earth, single shoots among Bryoerythrophyllum recurvirostrum, Pohlia cruda, Bryum moravicum, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00303. *M. tenerrima (Brid.) Lindb. — Rar: I: 9–10. Along outcrops of carbonate rocks in forest belt, in niches on fine earth, always in ± significant admixture among other species of mosses. S+. *Neckera oligocarpa Bruch. — Rar: I: 11; II: 21. Along rock outcrops in forest and mountain tundra belt, on surface of stones in pure mats and in mixture with Radula complanata, Pseudoleskeella rupestris, Hypnum cupressiforme. S+. Specimens of Neckera pennata (Hedw.) Bruch, Schimp. et W. Gümbel previously cited for the upper reaches of the Pol’ya River and the basin of the Man’ya River (Kildyushevskiy, 1956), apparently also belong to this species. *Niphotrichum canescens (Hedw.) Bednarek – Ochyra et Ochyra [≡ Racomitrium canescens Bridel] — Rar. I: 1, on protruding stones in stream bed, 65.25149°N, 62.17349°E, 11 VII 2018,

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YSU-MH-00240; ibidem, in Festuca ovina-lichen-moss community, on sandy soil in stream valley, prevails in moss cover cover, 65.25136°N, 62.17379°E, 11 VII 2018, YSU-MH-00254. S+. *Ochyraea alpestris (Hedw.) Ignatov et Ignatova — Un. I: 1, in channel of stream, on stones, in pure mats, 65.25149°N, 62.17349°E, 11 VII 2018, YSU-MH-00236. *O. duriuscula (De Not.) Ignatov et Ignatova — Un. II: 21, in channel of stream under snowfield, on stones on fine earth, 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00573. *O. norvegica (Bruch, Schimp. et W. Gümbel) Ignatov et Ignatova — Un. II: 21, rock outcrops in dwarf birch tundra, in mixture with Amphidium mougeotii and admixture of Oxystegus tenuirostris, Isopterygiopsis pulchella, 65.22070°N, 61.92725°E, 14 VII 2018, YSU-MH-00603. Previously, this species was known only from two locations in the northern part of the Urals (Zheleznova, 1994; D’yachenko, 1997). Oncophorus virens (Hedw.) Brid. — Un. II: 21, on grassy slope along bank of stream under snowfield, mainly in pure mats, once in mixture with Scorpidium revolvens, 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00548. S+. O. wahlenbergii Brid. — Sp. I: 1–3. In larch-birch-spruce Vacccinium vitis-idaea-green moss river valley forests, on decayed wood in mixture with other species of mosses and liverworts, less common in shrubby mountain tundra. S+. *Orthothecium strictum Lorentz — Rar. I: 9, at outcrops of carbonate rocks, on southeast-facing slope, in niches between stones, in pure mats and in mixture with Cyrtomnium hymenophylloides, 65.30118°N, 62.11997°E, 10 VII 2018, YSU-MH-00124; I: 11, stone block on gentle slope of mountain, on ledges of stones with minor admixture of Eurhynchiastrum pulchellum, Mnium marginatum, Cyrtomnium hymenophylloides, 65.27757°N, 62.11185°E, 12 VII 2018, YSU-MH-00361. *Orthotrichum anomalum Hedw. — S. I: 9, 10. At outcrops of carbonate rocks in forest belt, on surface of stones, mainly in pure mats. Once noticed in mixture with O. cupulatum and Porella platyphylla. S+. *O. cupulatum Brid. — Un. I: 10, outcrops of carbonate rocks on stones, in mixture with O. anomalum and Porella platyphylla, 65.29862°N, 62.08511°E, 12 VII 2018, YSU- MH-00306. S+. **O. hyperboreum Fedosov et Ignatova — Un. II: 19, rock outcrops on steep slope in young birch forest, on stones, without admixture of other species, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00443. The species was recently separated from O. pallens based on genetic analysis (Fedosov et al., 2017). It is found in the mountains of the East Siberian Arctic and Sub- arctic and in the Polar Urals. For the Subpolar Urals, the view is reported for the first time. S+. *Oxystegus tenuirostris (Hook. et Tayl.) A. J. E. Sm. — Rar. II: 19, rock outcrops on steep slope in young birch forest, in shaded niches on fine earth, in small mats and as a minor admixture among Schistidium pulchrum and Zygodon sibiricus, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00430; II: 21, rocky outcrops in dwarf birch tundra, single shoots among Ochyraea norvegica, Amphidium mougeotii, 65.22070°N, 61.92725°E, 14 VII 2018, YSU-MH-00603. Paludella squarrosa (Hedw.) Brid. — Fr. I: 5, 7; II: 18. In sedge-brown moss rich fens, at groundwater outlets. *Philonotis caespitosa Jur. — Un. III: 27, on silty-gravel shallows of river bank and in willow-grass-green moss community, noticeable admixture among Bryum pseudotriquetrum, Pohlia filum, Calliergonella lindbergii, 64.55854°N, 61.64242°E, 8 VII 2018, YSU-MH-00002. P. tomentella Molendo — Rar. I: 1; II: 21. Along banks of rivers and streams in forest and mountain tundra belts, in shrub (Betula nana, Salix)-sedge (Carex arctisibirica)-Sphagnum mire in mountain tundra, mainly in pure mats. S+ (immature).

227 Lapshina et al. The mosses of the Khulga River Basin (Subpolar Urals)

Plagiomnium ellipticum (Brid.) T. J. Kop. — Fr. I: 3, 5; II: 18; IV: 29. In sedge-brown moss and horsetail fens and spruce-willow-tussock sedge swampy low open forests, less often in flooded birch and birch-spruce shrubbery (alder, willow)-grass forests. Plagiopus oederianus (Sw.) H. A. Crum et L. E. Anderson — Un. I: 10, at outcrop of carbonate rocks, on ledges, on fine earth, in pure mats and together with Myurella tenerrima, Bryo- erythrophyllum recurvirostrum, Campyliadelphus chrysophyllus, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00289. S+. Plagiothecium cavifolium (Brid.) Z. Iwats. — Sp. I: 1, 3, 15. In birch-spruce and mixed shrub-herb and herb-green moss floodplain forests, in the forest floor. P. denticulatum (Hedw.) Bruch, Schimp. et W. Gümbel — Sp. I: 3; II: 17–18, 21. In shrub- grass and mountain green moss flooded river valley forests, less often on rocky outcrops in dwarf birch tundra, on fine earth between stones. P. svalbardense Frisvoll — Fr. I: 13; II: 17, 19. In river valley and mountain birch-spruce, birch-Pinus sibirica-spruce green moss forests, on decayed wood. S+. Earlier the Plagiothecium piliferum-like plants from the northern part of Asian Russia according to identification keys came out to P. laetum. Recently, based on molecular phylogenetic analysys, it was proved that they belong to Plagiothecium svalbardense (Ignatova et al., 2019), described from Svalbard (Frisvoll, 1981; Frisvoll, Elvebakk, 1996). Platydictya jungermannioides (Brid.) H. A. Crum — Sp. I: 9–11; II: 19. At outcrps of carbonate and basic rocks in forest belt, mainly as admixture to other species of mosses and liverworts. Pleurozium schreberi (Brid.) Mitt. — Cm. I: 1–3, 5–9, 14–15; II: 17–20; III: 23, 25; IV: 28–30. In different types of river valley and mountain forests, in dwarf shrub-Rubus chamaemorus-Sphagnum boggy open spruce forests and mountain tundra. Pohlia cruda (Hedw.) Lindb. — Cm: I: 1–3, 9–11, 14; II; 17, 19, 21. In river valley and mountain forests, along outcrops of carbonate rocks and rocky ledges, on soil and in niches between stones. S+. P. drummondii (Müll. Hal.) A. L. Andrews — Un. II: 21, along bank of stream under snowfield, on fine earth, single admixture to Diplophyllum taxifolium, Fuscocephaloziopsis albescens, Lophozia murmanica. 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00559. P. elongata Hedw. — Rar. II: 19, rock outcrops on steep slope in young birch forest, in niches between stones, on fine earth, with admixture of Rhabdoweisia crispata, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00471. S+; II: 22, on gravelly periodically flooded river bank, with Rhabdoweisia crispate, 65.10409°N, 62.21820°E, 18 VII 2018, YSU-MH-00036. S+. P. filum (Schimp.) Mårtensson — Rar. I: 1, on sandy soil in stream valley, in mixture with Bryum pseudotriquetrum, Calliergonella lindbergii, 65.25136°N, 62.17379°E, 11 VII 2018, YSU- MH-00653; III: 27, on silty-gravel shallows and in willow-grass-hypnum community, in mixture with Pellia neesiana, Scapania irrigua, Calliergonella lindbergii, 64.55854°N, 61.64242°E, 08 VII 2018, YSU-MH-00011. *P. lescuriana (Sull.) Grout — Un. III: 24 on river-bank sediment exposure, on silty sandy soil in mixture with Dicranella crispa, D. varia, Psilopilum cavifolium. No nodules were found in the studied specimens, 64.87081°N, 62.03472°E, 16 VII 2018, YSU-MH-00617. S+. P. nutans (Hedw.) Lindb. — Cm. I: 1–3, 5, 7–9, 14–15; II: 17–20; III: 23–25; IV: 27–30. In all types of river valley and mountain forests, boggy open forests, and mires, shrubby tundra, on stony outcrops on fine earth between stones. S+. Along with typical forms in one habitat, samples of dioecious plants were collected at the base of rock outcrops in the upper part of the forest belt. In this review, the species is considered in broad sense, including P. sphagnicola (Bruch, Schimp. et W. Gümbel) Broth.

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P. proligera (Kindb.) Lindb. ex Broth. — Sp. I: 3; III: 24. In birch-spruce and Pinus sibirica-birch-spruce shrub-grass river valley forests, along coastal cliffs and outcrops. P. wahlenbergii (F. Weber et D. Mohr) A. L. Andrews — Un. I: 1, along bank of stream, on sandy alluvium, in pure mats, 65.25149°N, 62.17349°E, 11 VII 2018, YSU-MH-00242. Polytrichastrum alpinum (Hedw.) G. L. Sm. — Fr. I: 3, 7, 11; II: 19–21. In shrubby tundras, by outcrops of rocks in forest belt, less often in spruce-shrub (alder, willow)-grass valley forests, along banks of streams. S+. P. longisetum (Sw. ex Brid.) G. L. Sm. — Un. I: 1, birch-willow-sedge-Sphagnum community in swampy valley, 65.25291°N, 62.17188°E, 11 VII 2018, YSU-MH-00232. Polytrichum commune Hedw. — Fr. I: 1–3; II: 17, 22; III: 27–28. In spruce-Betula nana-Sphagnum bogs and Pinus sibirica-birch-spruce shrub-lichen-green moss open forests, in river valley green-moss forests. S+. P. jensenii I. Hagen — Sp. I: 14; II: 16; III: 23. In swampy willow-sedge-Sphagnum birch low forests, Menyanthes trifoliata-sedge-Sphagnum thermokarst subsidences, in hollows of oligotrophic mire complexes. S+. P. juniperinum Hedw. — Rar. I: 2, in birch-spruce-Pinus sibirica-shrub-grass forest, single shoots among Ceratodon purpureus, 65.27042°N, 62.18309°E, 08 VII 2018, YSU-MH-00654; I: 5, in spruce-Betula nana-sedge (Carex juncella) open low forests, on sedge tussock, 65.29345°N, 62.14450°E, 10 VII 2018, 11E-18. P. piliferum Hedw. — Rar. I: 1–2; II: 19. Along edges of coastal cliffs, on sand and gravel deposits in valley of stream, on rock outcrops on fine earth. P. strictum Brid. — Fr. I: 1–8, 14; II: 17–18; III: 23, 25: IV: 28–30. In raised bogs and transitional fens on dwarf shrub-Sphagnum ridges and frozen palsas, in spruce dwarf shrub-Rubus chamaemorus-Sphagnum boggy open forests. S+. P. swartzii Hartm. — Un. II: 17, in the floodplain sedge communities (Carex rostrata), with minor admixture of Straminergon stramineum, 65.15069°N, 62.12681°E, VII VII 2018, YSU- MH-00366. Pseudobryum cinclidioides (Hübener) T. J. Kop. — Fr. I: 13; II: 16, 18; III: 23. In swampy and regularly flooded river valley forests, in swampy open low woodlands, in willow thickets. *Pseudoleskeella catenulata (Brid. ex Schrad.) Kindb. — Rar. I: 10–11; II: 19. At rock outcrops in forest and tundra belts, in pure mats and in admixture among Zygodon sibiricus, Neckera oligocarpa. *P. rupestris (Berggr.) Hedenäs et L. Söderstr. — Sp. I: 9, 14; II: 19–21. At rock outcrops of different composition in forest belt and in dwarf birch tundras, mainly in admixture among Neckera oligocarpa, Tortella tortuosa, Radula complanata, Porella platyphylla, Encalypta cf. streptocarpa. Psilopilum cavifolium (Wilson) I. Hagen — Un. III: 24, on shore exposure on sandy soil, forms pure carpets, and also in mixture with Dicranella varia, D. crispa, Pohlia proligera, 64.87081°N, 62.03472°E, 16 VII 2018, YSU-MH-00614. Pterigynandrum filiforme Hedw. — Rar. II: 19, 21. At rock outcrops in upper part of forest and tundra belts, on stones, in pure mats and mixed with Grimmia elatior and Pohlia nutans. Ptilium crista-castrensis (Hedw.) De Not. — Un. I: 1, in larch-birch-spruce dwarf shrub- green moss old-growth forest, on decayed wood, with Stereodon plicatulus and Sanionia uncinata, 65.27277°N, 62.17880°E, 11 VII 2018, YSU-MH-00655. Pylaisia polyantha (Hedw.) Bruch, Schimp. et W. Gümbel — Rar. I: 3. In Pinus sibirica-birch-spruce-alder (Duschekia fruticosa)-grass and Equisetum sylvaticum- herb river valley forests, at bases of tree trunks. S+.

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*Racomitrium lanuginosum (Hedw.) Brid. — Rar. II: 20, in dwarf shrub (Empetrum, Va c- cinium)-Diapenisia tundra, on stones, 65.20243°N, 61.98634°E, 14 VII 2018, YSU-MH-00489; ibidem, in rocky dwarf shrub-green moss tundra, 65.20436°N, 61.98496°E, 14 VII 2018, YSU-MH-00497. *Rhabdoweisia crispata (Dicks. ex With.) Lindb. — Rar. II: 19, rock outcrops on steep slope in young birch forest, on fine earth in niche between stones, in admixture to Pohlia elongata, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00471; II: 22, on gravelly flooded bank of river, with Pohlia elongata, 65.10409°N, 62.21820°E, 18 VII 2018, YSU-MH-00036. S+. Rhizomnium pseudopunctatum (Bruch et Schimp.) T. J. Kop. — Sp. I: 5, 8; II: 18. In Betula nana-sedge-moss fens, in dwarf birch-Sphagnum boggy woodlands, less often in valley forests. Rhytidium rugosum (Hedw.) Kindb. — Un. II: 21, under large boulder among dwarf birch tundra, in ground cover in pure mats and with Abietinella abietina, 65.21678°N, 61.93953°E, 14 VII 2018, YSU-MH-00508. Saelania glaucescens (Hedw.) Broth. — Rar. I: 13. In birch-spruce shrub-grass flooded river valley forests, on bare soil, in small pure mats and as admixture among Isopterigiopsis pulchella, Pohlia cruda, Mnium lycopodioides, Lophocolea minor. S+. Sanionia uncinata (Hedw.) Loeske — Cm. I: 1–3, 8, 14–15; II: 17–19; III: 23; IV: 28. In all types of river valley and mountain forests, less often in Pinus sibirica-spruce dwarf shrub-Sphag- num boggy woodlands. Schistidium agassizii Sull. et Lesq. — Sp. I: 1; II: 19, 22; III: 27. On the protruding stones in channels of streams and along river banks. S+. *Schistidium boreale Poelt — Rar. I: 9, at outcrops of carbonate rocks, 65.30118°N, 62.11997°E, 10 VII 2018, YSU-MH-00126; I: 7, on the top of small mountain, in pure mats, 65.30458°N, 62.14071°E, 10 VII 2018, YSU-MH-00170. ***S. frisvollianum H. H. Blom — Rar. I: 9, carbonate rock outcrops on southeast-fecing slope , in pur mats on stones, 65.30118°N, 62.11997°E, 10 VII 2018, YSU-MH-00127; on slope of north-eastern exposure with protruding carbonate boulders in Pinus sibirica-larch-spruce- shrub-green-moss forest, in niches between stones, 65.30118°N, 62.11997°E, 10 VII 2018, YSU- MH-00147. Earlier, the species was known only in Karelia from single finds, the Nenets Auton- omous District and in the Arctic of the Asian part of Russia, expanding southwards to 61°N (Ignatov et al., 2017). A find in the Subpolar Urals is quite predictable. *S. papillosum Culm. — Un. II: 19, in channel of mountain stream valley, on protruding stones, in pure mats, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00481. *S. platyphyllum (Mitt.) Perss. — Un. II: 19, in channel of creek in mountain valley, in pure mats, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00482. S+. *S. pulchrum H. H. Blom — Rar. I: 10–11; II: 19. At rock outcrops in upper part of forest belt, on stones, mainly in pure mats. S+. *S. rivulare (Brid.) Podp. — Un. II: 21, in stream bed under snowfield, in pure mats, 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00544. S+. *S. sordidum I.Hagen — Un: II: 21, in stream bed under snowfield, in pure mats, 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00578. S+. Sciuro-hypnum curtum (Lindb.) Ignatov — Sp. I: 1, 3; II: 18–19. In river valley and mountain spruce and Pinus sibirica-birch-spruce herb-green moss forests. *S. glaciale (Bruch, Schimp. et W. Gümbel) Ignatov et Huttunen — Un. II: 21, on grassy slope along bank of stream under snowfield, in mixture with Mnium blyttii, Lescuraea saxicola, 65.22291°N, 61.92264°E, 14 VII 2018, YSU-MH-00554. A rare arctic alpine species, known in the Urals from single finds (D’yachenko, 1997).

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*S. latifolium (Kindb.) Ignatov et Huttunen — Un. II: 21, on grassy slope along bank of stream under snowfield, in pure mats, 65.22291°N, 61.92264°E, 14 VII 2018, YSU- MH-00568. S. reflexum (Starke) Ignatov et Huttunen — Rar. II: 17, in larch-spruce-birch-Pinus sibirica-shrub-green moss river valley forest, 65.15105°N, 67.11533°E, 13 VII 2018, YSU- MH-00369; II: 19, in young birch forest in upper part of forest belt, in pure mats and in mixture with Sciuro-hypnum curtum, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00479. Scorpidium cossonii (Schimp.) Hedenas — Fr. I: 5, 712–13; II: 21. On Betula nana-sedge- brown moss mires and sedge-brown moss rich fens. *S. revolvens (Sw. ex anon.) Rubers — Sp: I: 1, 13; II: 18, 21. On Betula nana-sedge-brown moss mires and sedge-brown moss fens with nutrient-rich groundwater supply, in pure mats and with admixture of Scorpidium cossonii; Meesia triquetra, Campylium stellatum, Mesoptychia rutheana. Scorpidium scorpioides (Hedw.) Limpr. — Sp. I: 5, 12. In Betula nana-tussock sedge-bown moss mires and waterlogged sedge-brown moss fens. Sphagnum angustifolium (C. E. O. Jensen ex Russow) C. E. O. Jensen — Cm. I: 1–8, 14; II: 16–18; III: 23, 25; IV: 28–30. In raised bogs and transitional fens, in spruce- and birch-Pinus sibirica-spruce dwarf shrub-Sphagnum mires and boggy open woodlands, often as a dominant and co-dominant. **S. annulatum H. Lindb. ex Warnst. — Rar. IV: 29, in mesooligotrophic ridge-hollow complex, in Menyanthes trifoliata-Sphagnum lawns, 64.33912°N, 61.06640°E, 18 VII 2018, Filippov, YSU-MH-00265. Sphagnum annulatum is rare in Siberia, close to S. jensenii, which until recently was not separated from the latter. In the flat part of the autonomous area, it is found at three sites very remote from each other (Lapshina et al., 2019). For the Subpolar Urals is recorded for the first time. S. balticum (Russow) C. E. O. Jensen — Fr. I: 4, 6; II: 17–18, 22; III: 23, 25; IV: 29–30. In oligotrophic wet hollows and lawns of flat palsa mires, and in thermokarst subsidence of high palsa-hollow complexes, often as a dominant. S. capillifolium (Ehrh.) Hedw. — Un. I: 6, in the dwarf shrub-S. (S. fuscum) community on frozen palsa, 65.30171°N, 62.14087°E, 10 VII 2018, YSU-MH-00645. S. centrale C. E. O. Jensen — Cm. I: 3; II: 16, 18; IV: 29–30. In minerotrophic and transitional dwarf shrub-sedge-Sphagnum fens and sedge-herb-Sphagnum mires. S. compactum Lam. et DC. — Rar. I: 3, in horsetail-sedge (Carex rostrata)-Sphagnum thermokarst subsidence among the Betula nana-Sphagnum open spruce woodlands, 65.27863°N, 62.19419°E, 12 VII 2018, YSU-MH-00217; ibidem, in sedge-Sphagnum community, 65.27875°N, 62.19427°E, 11 VII 2018, Filippov 13F-18. S. fimbriatum Wilson — Fr. I: 3; II: 16–17; III: 23, 25; IV: 29–30. In floodplain sedge (Carex rostrata) community, on transitional Menyanthes-sedge-Sphagnum fens. S. fuscum (Schimp.) H. Klinggr. — Cm. I: 2–-8; II: 17–18, 22; III: 23, 25; IV: 29–30. In raised pine-dwarf shrub-Sphagnum bogs, on frozen palsas of flat palsa complexes, in boggy dwarf shrub-Rubus chamaemorus-Sphagnum spruce and pine-spruce open woodlands, often as a dominant. S. girgensohnii Russow — Cm. I: 1–3, 5–8; II: 17–18, 21–22; III: 23, 25; IV: 28–30. In spruce and Pinus sibirica-spruce shrubby cloudberry-Sphagnum open woodlands, often as a dominant, less common in transitional fens. S. jensenii H. Lindb. — Sp: I: 3–4, 6; IV: 29–30. In oligotrophic wet hollows and watelogged lawnsof flat palsa complexes, often as a dominant.

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*S. lenense H. Lindb. ex L. I. Savicz — Rar. I: 6, in dwarf shrub (Ledum palustre, Betula nana)-Rubus chamaemorus-Sphagnum community of frozen palsa, 65.30051°N, 62.14294°E, 10 VII 2018, YSU-MH-00177; ibidem, on dwarf shrub-Sphagnum palsa in the lake-flat palsa complex, Glazunov, YSU-MH-00050. S. lindbergii Schimp. ex Lindb. — Sp. I: 4, 6; III: 25. In oligotrophic waterlogged hollows of flat palsa complex, in thermokarst dessions, often as a dominant or co-dominant. S+. S. divinum Flatberg et Hassel [= S. magellanicum auct.] — Cm. I: 6, III: 23, IV: 25–26. In ridges of transitional aapa mires, less often at the base of frozen hummocks in flat-palsa complexes. S. majus (Russow) C. E. O. Jensen — Rar. I: 4, in sedge (Carex rotundata)-Sphagnum hollows of flat palsa complex, 65.29921°N, 62.14483°E, 10 VII 2018, YSU-MH-00183; I: 6, in sedge (Carex rotundata)-Sphagnum wet depression in flat palsa complex, 65.27956°N, 62.14740°E, 12 VII 2018, YSU-MH-00656. ***S. mirum Flatberg et Thingsgaard — Un. III: 23, thermokarst depression in high palsa-mire complex, with minor admixture of S. obtusum, 64.93691°N, 61.99780°E, 16 VII 2018, Filippov, YSU-MH-00202. A rare arctoboreal species recently identified in Russia (Flatberg et al., 2016). Until present it was known from three localities in the autonomous area (Lapshina et al., 2018; Ellis et al., 2018). A new find of S. mirum in the Khulga River Basin was expected due to confinement to the subarctic regions of North Asia and Alaska. S. obtusum Warnst. — Cm. I: 13; II: 16–17, 22; III: 23; IV: 28–30. On sedge-herb-Sphagnum poor fens and Betula nana-Sphagnum transitional mires. S. platyphyllum (Lindb. ex Braithw.) Warnst. — Fr. I: 3; III: 25, IV: 29–30. In waterlogged sedge hollows and low ridges of transitional aapa mires. S. riparium Ångstr. — Cm. I: 3; II: 16–17; III: 23; IV: 29–30. In sedge-herb-Sphagnum poor fens and Betula nana-sedge-Sphagnum mesotrophic mires, often as a dominant and co-dominant. S. russowii Warnst. — Sp. I: 3; II: 17; IV: 30. In boggy Pinus sibirica-spruce dwarf shrub-Sphagnum open woodlands, on ridges of transitional aapa mires. S. squarrosum Crome — Fr. I: 13; II: 16; IV: 29–30. In transitional sedge-herb-Sphagnum mires and swampy open woodlands. S. subsecundum Nees — Fr. I: 3; II: 16; III: 25; IV: 29–30. In waterlogged sedge hollows and low ridges of transitional aapa mires. S. teres (Schimp.) Ǻngstr. — Sp. I: 5; II: 18; IV: 29–30. In boggy larch-spruce dwarf shrub-Sphagnum open woodlands, in sedge-moss fens, at groundwater outlets. S. warnstorfii Russow — Cm. I: 5, 13; II: 16, 18, 21. On dwarf shrub-sedge-Sphagnum mires of nutrient-rich groundwater supply in forest and mountain tundra zones. S. wulfianum Girg. — Rar. IV: 28–29. In wet spruce-birch-larch shrub-Sphagnum-green moss forest, twice on ridges in sedge-Sphagnum aapa complex, with S. angustifolium. Splachnum luteum Hedw. — Un. II: 17, in boggy spruce-Betula nana-Sphagnum (S. girgensohnii) open woodland, in a mixture with S. rubrum, 65.15403°N, 62.10863°E, 14 VII 2018, YSU-MH-00375. S+. S. rubrum Hedw. — Un. II: 17, in boggy spruce-Betula nana-Sphagnum (S. girgensohnii) open woodland, in a mixture with S. luteum, 65.15403°N, 62.10863°E, 14 VII 2018, YSU- MH-00376. S+. *Stereodon bambergeri (Schimp.) Lindb. — Un. I: 11, stone block on gentle slope of mountain, on stone on fine earth, single shoots among Sphenolobus minutus, Pohlia cruda, Isoptery-

232 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 54(1): 211–241. 2020 giopsis pulchella, Tritomaria quinquedentata, Barbilophozia hatcheri, 65.27757°N, 62.11185°E, 12 VII 2018, YSU-MH-00352. S. plicatulus Lindb. — Fr. I: 3, 11, II: 17–20. In the river valley and mountain forests, at outcrops of basic and silicate rocks in forest and mountain tundra belts, in pure mats and mixed with other species. *S. vaucheri (Lesq.) Lindb. ex Broth. — Rar. I: 9, at outcrops of carbonate rocks on southeast-facing slope, in pure mats, less often in mixture with Drepanium recurvatum, 65.30118°N, 62.11997°E, 10 VII 2018, YSU-MH-00107; I: 10, on rocky outcrop of carbonate rocks, on stones, single shoots between Tortella tortuosa, 65.29862°N, 62.08511°E, 12 VII 2018, YSU- MH-00297. Straminergon stramineum (Dicks. ex Brid.) Hedenäs — Cm. I: 5; II: 16–18, IV: 29–30. In transitional and minerotrophic fens, at groundwater outlets. S+. *Syntrichia ruralis (Hedw.) F. Weber et D. Mohr — Rar. I: 9, at outcrops of carbonate rocks, on slope of the eastern exposure, at stone ledges in fine earth, in pure mats, 65.30118°N, 62.11997°E, 10 VII 2018, YSU-MH-00133; I: 10, on rocky outcrop of carbonate rocks, in niches on fine earth, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00305. Tetraphis pellucida Hedw. — Rar. I: 3; II: 19; III: 23. In mountain and river valley forests on decayed wood, on birch-Pinus sibirica-shrub-Sphagnum high palsa, on stump. Timmia austriaca Hedw. — Rar. I: 9, at outcrops of carbonate rocks, on southeast slope, on ledges of stones in pure mats, 65.30118°N, 62.11997°E, 10 VII 2018, YSU-MH-00137; I: 10, on rocky outcrop of carbonate rocks, in niches on fine earth, mixed with Encalypta rhaptocarpa, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00293. Tomentypnum nitens (Hedw.) Loeske — F.: I: 5, 7–8, 12–13; II: 18. In minerotrophic sedge- brown moss rich fens, in spruce-dwarf shrub-tussock sedge and in boggy Betula nana-Rubus chamaemorus-Sphagnum open woodlands, at groundwater outlets. *Tortella fragilis (Hook. et Wils.) Limpr. — Un. II: 21, in Betula nana-sedge (Carex arctisibirica)-Sphagnum (S. warnstorfii) alpine mire, in pure mat, 65.22070°N, 61.92725°E, 14 VII 2018, YSU-MH-00526. T. tortuosa (Hedw.) Limpr. — Sp. I: 9-10; II: 20. At outcrops of rocks of different composition in forest and tundra belts. *Trichostomum crispulum Bruch — Rar. I: 9, at outcrops of carbonate rocks, on southeast slope, without admixture of other species, 65.30118°N, 62.11997°E, 10 VII 2018, YSU- MH-00134; I: 10, on outcrop of carbonate rocks, on stones, 65.29862°N, 62.08511°E, 12 VII 2018, YSU-MH-00309. A widely spread calcephilous species, for the Urals known from isolated finds from limestone rocks. Ulota curvifolia (Wahlenb.) Lilj. — Rar. I: 11, stone block on gentle mountain slope, on surface of stone, in pure mats, 65.27757°N, 62.11185°E, 12 VII 2018, YSU-MH-00334; II: 19, rocky outcrops on steep slope in young birch forest, 65.18130°N, 62.99642°E, 14 VII 2018, YSU-MH-00426. S+. Warnstorfia exannulata (Bruch, Schimp. et W. Gümbel) Loeske — Cm. I: 1, 5, 12; II: 16, 18; III: 23; IV: 29–30. In transitional and minerothophic fens, at groundwater discharge. S+. Warnstorfia fluitans (Hedw.) Loeske — Fr. I: 4, 6; IV: 29–30. In wet hollows, low Sphagnum hummocks and ridges of raised bogs and transitional aapa mires. S+. Warnstorfia pseudostraminea (Müll. Hal.) Tuom. et T. J. Kop. — Un. II: 16, vast mesotrophic open fen, in cotton grass (Eriophorim gracile)-sedge-herb community, minor admixture among Pseudobryum cinclidioides, Warnstorfia exannulata, Sphagnum obtusum, 65.20831°N, 62.14333°E, 16 VII 2018, YSU-MH-00657.

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Warnstorfia sarmentosa (Wahlenb.) Hedenäs — Un. II: 18, groundwater outflows among larch-spruce-horsetail-dwarf shrub-Sphagnum open forest, in pure mats and in admixture to Cinclidium stygium, 65.17609°N, 62.02684°E, 14 VII 2018, YSU-MH-00410. ***Warnstorfia tundrae (Arnell) Loeske — Un. I: 7, at places of groundwater discharge at base of mountain, forms lush carpet with Calliergon giganteum, 65.30458°N, 62.14071°E, 10 VII 2018, YSU-MH-00174. Previously W. tundrae was known only in the Northwest of Russia and in the Arctic of the Asian part of Russia, in Central Siberia, Yakutia and the Far East. Finds of the species in the Subpolar Urals fills the gap between in the Northwest of Russia and in the Arctic of the Asian part of Russia. Zygodon sibiricus Ignatov, Ignatova, Z. Iwats. et B. C. Tan — Un. II: 19, on cliff outcrops of rocks in young birch forest, on surface of stones, abundantly in pure mats and in admixture to Neckera oligocarpa, Schistidium pulchrum, YSU-MH-00423. S+.

In total, in the Khulga River Basin 210 species of mosses, including 24 Sphagnum species were revealed. The main part of the bryoflora of the Khulga River Basin is represented by widespread boreal holarctic species — Aulacomium palustre, Bryum pseudotriquetrum, Hy- locomium splendens, Pleurozium schreberi, Pohlia nutans, P. cruda, Sanionia uncinata, Sphagnum angustifolium, S. balticum, S. fuscum, S. girgensohnii, S. obtusum, S. riparium, S. warnstorfii, Straminergon stramineum, Warnstorfia exannulata, etc. In the foothills of the Khulga River valley 54 species were recorded. The species diversity of mosses increases noticeably along the right bank of the river at its upper reaches (key areas I and II), where the river basin embraces mid-mountain areas (up to 750 m high) along the eastern slope of the Urals. There 201 species were recorded, i. e., 95.3% of the recorded species in the study area. A high diversity of mosses in the Upper Khulga River basin in comparison with the foothill area and the adjacent part of the plain, can be explained by the presence of rocks with different composition, including carbonate rock outcrops and boulders, as well as the development of peat mires with nutrient-rich groundwater supply. The wide distribution of carbonate rocks resulted in a high number of calciphilous species in the flora. These include Anomodon attenuatus, Bryoerythrophyllum recurvirostrum, Cyrtomnium hymenophylloides, Distichium capillaceum, Drepanium recurvatum, Encalypta rhaptocarpa, E. cf. streptocarpa, Leucodon sciuroides, Myurella julacea, M. sibirica, M. tenerrima, Orthothecium strictum, Orthotrichum anomalum, Plagiopus oederianus, Platydictya jungermannioides, Pseudoleskeella catenulata, Schistidium frisvollianum, Syntrichia ruralis, Trichostomum crispulum. A significant part of species recorded in the Khulga River basin (92 species or 43.4%) were previously known from single finds of I. D. Kildyushevsky (1956) in the southern part of the Subpolar Urals on the territory of the Khanty-Mansi Autono- mous Area — Yugra (Man’ya, Pol’ya, Shchekur’ya river basins). Among 210 identified species, 56 species (28.9%) are new to the territory of the Khanty-Mansi Autonomous Area — Yugra. Of these, about a half were previously registered by us for the Subpolar Urals in Mount Ner-Oika area (in 2013) and (or) at the

234 Новости систематики низших растений — Novosti sistematiki nizshikh rastenii 54(1): 211–241. 2020 upper reaches of Puiva River (in 2015). However, the materials of these studies have not yet been published. There were only very brief records of the most characteristic and widespread species without indicating their exact collection sites (Lapshina et al., 2015, 2016; Skuchas, Lapshina, 2018). A number of species, which were recorded for the first time for the Khanty–Mansi Autonomous Area — Yugra are widespread in the Urals. The lack of information about their location on the eastern slope of the Subpolar Urals is resulted from an extremely poor knowledge of its moss flora. These are such widespread species as Grimmia elatior, Hymenoloma crispulum, Kiaeria starkei, Myurella julacea, M. tenerrima, Niphotrichum canescens, Orthotrichum anomalum, Racomitrium lanuginosum, Tortella fragilis. However, many species that are new to the autonomous area are certainly rare and are known in the Urals by only a few records. These are Amphidium mougeotii, Ano- modon attenuatus, Campylidium calcareum, Encalypta pilifera, Hedwigia mollis, Hygro- hypnella polare, Pseudoleskeella catenulata, Orthotrichum cupulatum, Oxystegus tenuirostris, Ochyraea norvegica, Sciuro-hypnum glaciale, Trichostomum crispulum. In the Khulga River Basin, five species new to the whole Urals were collected: Didymodon perobtusus, Lewinskya iwatsukii, Orthotrichum hyperboreum, Schistidium frisvollianum, S. mirum, Warnstorfia tundrae. Moreover, the last two species were noted earlier also in the plain part of the autonomous area (Lapshina et al. 2018; Ellis et al., 2018). The rest of the abovementioned species represent the first records for the whole autonomous area. Dicranum schljakovii, Encalypta pilifera, Grimmia elatior, G. tergestina, Orthotrichum hyperboreum were collected in the Subpolar Urals for the first time. In the upper and middle reaches of the Khulga River five moss species were found, which are included in the Red Book of the Khanty-Mansi Autonomous Area – Yugra. These are Meesia triquetra, Paludella squarrosa, Psilopilum cavifolium, Scorpi dium scorpioides and Zygodon sibiricus. Two more species — Calliergon richardsonii and Meesia uliginosa — are included into the additional list of species of special concern. Based on the research and new information on occurrence and data on the distribution of the species, a number of rare mountain species of mosses can be recommended for inclusion in the next edition of the Red Book of the Khanty-Mansi Autonomous Area – Yugra. The obtained data will be used in further investigation of the biota of the region and the history of its formation, in planning environmental protection measures and expert assessments. Acknowledgments The authors are grateful to the administration and staff of the Direction of Pro- tected Areas in the Khanty-Mansi Autonomous District — Yugra: I. V. Pavlovskikh, A. V. Korovin, O. S. Fedosov for organization of field work and logistic. We also very grateful to Ilya Filippov from Yugra State University (Khanty-Mansiysk), as well as Valeriy Glazunov and Svetlana Nikolayenko from the Institute for the Development of the North of the SB RAS (Tyumen) for their help during the field investigations.

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