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БИОЛОГИЧЕСКИЕ НАУКИ
УДК 574.472
ON THE MOSS FLORA OF THE YUGYD VA NATIONAL PARK (MA-LYI PATOK RIVER BASIN, SUBPOLAR URALS)
G.V. ZHELEZNOVA, T.P. SHUBINA, S.V. DEGTEVA, Y.A. DUBROVSKY, T.N. PYSTINA
Institute of Biology, Komi Science Centre, UB RAS, Syktyvkar zheleznova@ib.komisc.ru, tshubina@ib.komisc.ru, degteva@ib.komisc.ru, dubrovsky@ib.komisc.ru, pystina@ib.komisc.ru
An annotated list of mosses from the basin of the Malyi Patok River (Yugyd Va National park, Subpolar Urals) is presented. It includes 80 species from 43 genera and 22 families. Data on habitats of the mosses and brief description of natural conditions and main types of vegetation are given. The following rare and protected in the Komi Republic species are registered: Ochyraea norvegica (Bruch et al.) Ignatov et Ignatova, Pohlia ludwigii (Spreng. ex Schwflgr.) Broth, and Polytrichastrum sexangulare (Floerke ex Brid.) G. L. Sm.
Keywords: mosses, the Malyi Patok River basin, Yugyd Va National Park, Subpolar Urals
Г.В. ЖЕЛЕЗНОВА, Т.П. ШУБИНА, С.В. ДЕГТЕВА, Ю.А. ДУБРОВСКИЙ, Т.Н. ПЫСТИНА. К ФЛОРЕ ЛИСТОСТЕБЕЛЬНЫХ МХОВ НАЦИОНАЛЬНОГО ПАРКА «ЮГЫД ВА» (БАССЕЙН Р. МАЛЫЙ ПАТОК, ПРИПОЛЯРНЫЙ УРАЛ)
Представлены сведения об основных типах растительности и видовом составе листостебельных мхов труднодоступной части национального парка «Югыд ва» в пределах верхнего, среднего и нижнего течения р. Малый Паток (Приполярный Урал). Впервые приведен аннотированный список видов мхов, насчитывающий 80 видов из 43 родов и 22 семейств, включающий редкие и охраняемые в Республике Коми виды: Ochyraea norvegica (Bruch et al.) Ignatov et Ignatova, Pohlia ludwigii (Spreng. ex Schwдgr.) Broth., Polytrichastrum sexangulare (Floerke ex Brid.) G. L. Sm. Показана роль мохообразных в сложении напочвенного покрова растительных сообществ. Полученные данные расширяют представление о бриофлоре национального парка «Югыд ва», изученность которой к настоящему моменту оценивается как недостаточная.
Ключевые слова: листостебельные мхи, река Малый Паток, национальный парк «Югыд ва», Приполярный Урал
Introduction
The «Yugyd Va» National Park with the total area of about 1894,1 thousand ha is situated in the Subpolar and Northern Urals [1] and has the status of protected natural territory of Federal level. Analysis of the currently available information about the biological diversity of the natural complexes of the park suggests that its flora is investigated insufficiently. First data on the bryophytes of the Park were contained in the works of R.R.Pole [2]. Then the Sablya Mountain Range attracted the greatest attention of the botanists [3, 4, 5]. Since 2005 scientists from the Institute of Biology of the Komi Science Centre, Ural Branch, RAS, carry out detailed study of the bryoflora of the northern part of the Park. At present the leafy Mosses of the «Yugyd Va» National Park includes 301 species of110 genera
and 37 families [6, 7]. This is 60 % of the entire Moss flora of the Komi Republic. With obtaining of new data the list of mosses is periodically reconsidered. This paper contains research results of the bryoflora of the Malyi Patok River basin - a part of the underinvesti-gated southern area ofthe National Park.
Studies were carried out in 2005-2006 in the remote areas of the Malyi Patok River basin at the key sites in the upper, middle and lower reaches of the river (Figure). The studied landscapes being a part of the largest national protected area (NPA) of the Komi Republic - the Yugyd Va National Park, are not impacted by anthropogenic pressure. Plant communities develop in the regime of spontaneous dynamics. Natural successions in forest communities are disturbed by such exogenous factors as the impact of wind and fire. There are no significant areas of windfalls and burnt areas.
Figure. Study area and collecting localities in the Malyi Patok River basin (Yugyd va national park, Subpolar Urals).
Locality 1 - Vicinities of the lakes Patok and Bolshoye (64°28' N, 59°33' E); Locality 2 - The Malyi Patok River, vicinities of environmental station «Uschelye» (64°18' N, 59°03' E); Locality 3 - Mouth of the Lorcempeya River (64°14' N, 58°51' E); Locality 4 - The Malyi Patok River, 7,3 km downstream the mouth of the Lorcempeya River (64°14' N, 58°42' E); Locality 5 - Island in the riverbed of the Schugor River, 2 km downstream the mouth of the Malyi Patok River (64°10' N, 58°31' E). ▲ - Places of collecting of mosses.
The studied part of the Malyi Patok River basin is located in mountains and foothills of the Subpolar Urals. The river mainly flows in a narrow valley where the floodplain is often poorly expressed. The bottom of the river is covered by pebbles and rocks. Large rounded boulders brought by ancient glacier are often found in the riverbed. Water has low mineralization, clear and cold. River flow is fast. Riverbed meanders strongly and is characterized by alternating reaches and sand spits.
The region has severe climatic conditions. Average annual temperature - 3.5°C. The temperatures of the coldest month (January) often fall to - 40°C. During the warmest month (July) air temperatures may reach +25-30°C, but mean temperature + 14°C. The vegetation period lasts from 60 to 120 days with late spring early autumn frosts. On the highest mountain tops the stable snow cover forms since September and completely melts in May. Meridional Ural ridges redistribute precipitations brought by moist northwest winds. Annual precipitation is up to 1500 mm [8].
Mountain-taiga landscapes of the Subpolar Urals and northern taiga landscapes of its western foothills located in the middle and lower reaches of the Malyi Patok River basin belong to the Novozemelsky-Ural mountain country [9]. In mountain landscape area the absolute heights vary from 230 m (in the valley of the river Malyi Patok) to 600-1035 m (on the tops of ridges) above sea level. Altitudinal belts are presented by
mountain stony belt, mountain tundra belt and mountain forest belt. Within rather small territory fragments of landscapes of stony deserts, mountain tundra, birch crooked forests of Betula tortuosa (nomenclature according to S.K. Cherepanov [10]), mountain dark coniferous forests are combined with complexes of mountain valleys drained by rivers and mountain streams. Mountain forest belt is formed by spruce (Picea obovata), fir-spruce (Abies sibirica) and, to a lesser extent, fir spruce-birch vegetation mainly of green moss type. High-grass and high-grass-moss dark coniferous forests developed along the streambed. Larix sibirica often occurs in the stands. Oligotrophic and mesotrophic mires occur at the foothills and on the terraces of the Malyi Patok River valley and its large tributaries, around mountain lakes marsh areas with peat deposits of the transitional and upland types are spread. For the river valleys small areas of high-grass meadows, birch and willow stands along the stream bed and interspersed with arrays of dark coniferous herb-green moss forests, are usual.
Foothills are covered by northern taiga landscape complexes of elevated ridges and large undulating depressions. Depressions are cut by valleys of semi-mountain rivers often swampy. Plateau-like tops of the highest ridges, pre-top terraces, extensive depressions within the plateau are occupied with stony gravels alternating with areas of mountain tundra, and in the depressions of the relief - small upland dwarf-
shrub grass-sphagnum mires with Betula nana. More low on the slopes the strip of foothill forests is located. Dark coniferous forests dominate here as in the mountain landscape zone. Stands are formed mainly by Siberian tree species - Picea obovata and Abies sibirica, Pinus sibirica and Betula pubescens often occur in the stands. At the watershed, forests alternate with hummock-ridge mires. Valleys of the rivers are covered by coniferous and birch forests and small meadows. Mesotrophic mires and spring bogs occur in the flood-plains and depressed terraces.
Results and discussion
Vegetation of the river valley has high coenotical and species diversity. Communities from perennial herbs, spruce, birch and willow stands often of herbal type and mires establish here.
Thickets of Petasites radiatus changing by meadows are usual for shallow waters and towpath habitats of the Malyi Patok River and its largest tributary - the Lorcempeya River. Fontinalis antipyretica (nomenclature follows «Check-list of mosses of East Europe and North Asia»[11]) is the most common species in the Malyi Patok River. It often forms thickets on the bottom of shallow parts of the riverbed. This species is typical for the rivers of most regions of the Northern hemisphere and was registered in all the studied rivers of the Timan and the Urals. Ochyraea duriusculaand and O. norvegica were found on the flooded gravels. Warnstorfia exannulata was always found in the meander lakes. The following species occur along the water edge: Philonotis fontana, Callier-gonella lindbergii, Calliergon cordifolium, Pseudobryum cinclidioides, Warnstorfia exannulata, Brachythecium mildeanum, Brachythecium rivulare, Rhizomnium pseudopunctatum and Sphagnum riparium. Ni-photrichum canescens was found many times at dry sandbanks. This species is actively involved in overgrowth of sloping sandbanks as well as Pleurozium schreberi, Hylocomium splendens and Rhytidiadelphus subpinnatus.
High grasses (Calamagrostis purpurea, Phalaroides arundinacea, and Alopecurus pratensis), herbs (Aconitum septentrionale, Angelica archangelica, Cirsium hetrophyllum, Crepis sibirica, Filipendula ul-maria and Thalictrum minis) and sedges (Carex atherodes) dominate in the meadow communities. Cli-macium dendroides, Calliergonella lindbergii, Brachy-thecium mildeanum, Plagiomnium ellipticum, Pohlia wahlenbergii, Philonotis fontana and species of genus Bryum are characteristic for floodplain meadows cenoses. Calliergonella lindbergii and Aulacomnium palustre grow in overwet places of the meadows. Meadows are not mowed down and so are gradually replaced by shrub and tree vegetation. We noted establishing of trees Picea obovata, Betula pubescens and shrubs (more often Salix phylicifolia and other Salix species) at many investigated meadows. But these trees and shrubs do not form a closed canopy. In a narrow valley of the Malyi Patok River woody plants are damaged or destroyed during the ice drift.
Birch forests from Betula tortuosa and B. pubescens are usual for the valley of the Malyi Patok River. Moss cover is underdeveloped, fragmented and has, as a rule, low total projective cover (1-5 %) under the canopy of herb birch forests. Inhibition of bryophytes in these plant communities is due to the dense herb cover and abundant annual leaves falling off trees and shrubs. In these ecosystems, mosses inhabit mainly such substrates as rotting wood, butts and trunks of living trees and small areas of soil. Pleurozium schre-beri, Hylocomium splendens, Ptilium crista-castrensis and Rhytidiadelphus triquetrus grow on soil in small clumps, Plagiomnium ellipticum, Climacium dendroides and Brachythecium mildeanum often grow in wet depressions (on the bottom of windfall hollows). Butts of trees are typical habitats for Dicranum fuscescens, D. scoparium, Rhytidiadelphus subpinnatus, Pleurozium schreberi, Hylocomium splendens, Ptilium crista-castrensis and Polytrichastrum alpinum. Decaying fallen trunks, stumps and various wood residues in birch grass are entirely covered with mosses including species occurring only on floodplain ecotopes - Dicranum montanum, Myrinia pulvinata and Pylaisia polyantha. The last two are common epiphytes in the southern and middle taiga.
Herb-sphagnum birch forests associated with depressions on the terraces have similar moss diversity comparing with birch herb forests. We noted the increase and strengthening of the role of hygrophytes -Brachythecium mildeanum, Sphagnum girgensohnii and S. majus. Projective cover of mosses reached 80 %.
Moss layer in birch forests of green moss type emerging on the above the floodplain terraces in ecotopes with poor soils is formed by very few species -Pleurozium schreberi, Hylocomium splendens, Poly-trichum commune, Dicranum scoparium. Trunks of fallen trees and stumps are often covered by Brachythecium spp., Dicranum scoparium, D. fuscescens, Sanionia uncinata, Polytrichum juniperinum, and Ptilium crista-castrensis. Trunks of birch are covered by mosses only in the lower parts.
Moss cover of willows is underdeveloped and is mainly presented by narrow strips along the banks of the Malyi Patok River, streams and lakes and on the edges of mires. Projective cover of mosses varied from 5 to 30-40 %. Dominants are not clear, bryophytes grow fragmentarily and form clumps. The following Sphagnum mosses are abundant in willow communities in stream valleys on the swampy banks of lakes besides the typical taiga green mosses: Sphagnum squarrosum, S. riparium, S. girgensohnii, S. capilli-folium, S. magellanicum, S. angustifolium, as well as the species tolerant to temporary flooding - Calliergon cordifolium, C. giganteum, Straminergon stramineum, Calliergonella lindbergii, Aulacomnium palustre.
Willow communities growing in conditions of flowing humidification show another species diversity of bryophites. Projective cover of mosses there increased and sometimes reached 60 %. Soil surface is covered by Climacium dendroides, Plagiomnium ellipticum, Brachythecium rivulare, B. mildeanum and Philonotis fontana. Rotting wood is covered by clumps from San-
ionia uncinata, Brachythecium mildeanum, Plagiomnium ellipticum, Rhythidiadelphus triquetrus, R. subpinnatus, Calliergonella lindbergii, Ptilium crista-cast-rensis, Pleu-rozium schreberi, Hylocomium splendens, Dicranum fuscescens, D. scoparium and some other moss species. Moss cover is less developed in more drained sites (total projective cover 5-10 %). Such oligomesotrophic and mesotrophic species as Pleurozium schreberi, Dicranum scoparium, D. fuscescens and Hylocomium splendens were registered in these communities as well as the species preferring richer soils - Rhodobryum roseum and Rhythidiadelphus triquetrus growing on rotting wood, on the butts of trees and more rare on soil.
The core of floristic complex of vascular plants in dark coniferous forests in the valley of the Malyi Patok River is formed by the species typical for the taiga picea forests. Gymnocarpium dryopteris is the most constant and abundant among them. Vaccinium myrtillus, Trien-talis europaea, Maianthemum bifolium, Linnaea borealis, Dryopteris expansa, Rubus arcticus, Equisetum sylvati-cum and Carex globularis also have high constancy. Sufficiently high illuminance under the canopy of northern open spruce forests contributes to the growth of light-demanding marginal and meadow species. Many of them meet suitable conditions in valley spruce forests. Calamagrostis purpurea, Chamae-nerion angustifolium, Cirsium heterophyllum, Geranium albiflorum, Solidago virgaurea, Veratrum lobelianum are most constant species of these groups. Moss cover is dominated by green mosses. Hylocomium splendens and Pleurozium schre-beri are most abundant among them. Ptilium crista-castrensis, Sanionia uncinata, Dicranum fuscescens and species from genus Brachythecium usually grow on fallen stems and butts of trees. The role of sphagnum mosses, especially Sphagnum girgensohnii, increases with the increase in stagnant humidification.
Fir forests are located in the valley of the river and prefer sites with well drained soils. Moss cover is well developed and its total projective cover reaches 90 %. Pleurozium schreberi and Hylocomium splendens dominate here. Such common moss species as Sanionia uncinata, Pleurozium schreberi, Hylocomium splendens, Dicranum fuscescens, Ptilium crista-cast-rensis and species from genus Brachythecium cover lower parts of tree stems, fallen trees and stumps. Roots and butts of trees in fir forests are covered by mosses growing on soil surface and rotting wood. Mosses-indicators of stagnant humidification are not abundant. We found Polytrichum commune, P. strictum, Sphagnum girgensohnii and Calliergon cordifolium.
Wetland complexes have high diversity in the valley of the Maliy Patok River. This is largely due to their different origin. A lot of lakes were formed in the valley during post-glacial time. Their overgrowth during the primary succession promoted the formation of mires. Most of mires are mesotrophic, sedge-sphag-num or cotton grass-sphagnum with stagnant humidification.
Cold springs are common in the valley of the river. Such ecotopes are also waterlogged, but there is fundamentally different structure of wetlands - herb-sphagnum-hypnum communities prevail here. Paludel-la squarrosa, Helodium blandowii and Tomentypnum nitens that grow on small elevations within the wetter
sites or in moderately humid sites within the evtrophic mires are indicators of rich mineral nutrition of mires often associated with spring humidification. Plagiom-nium ellipticum, Pseudobryum cinclidioides, Climacium dendroides, Rhodobryum roseum, Warnstorfia exannu-lata, species from genus Calliergon, and sphagnum mosses - Sphagnum girgensohnii, S. squarrosum and S. riparium are abundant at the edges of lowland bogs. Species of the mosses less selective to the soil richness are found near the central part of the bogs - Aulacom-nium palustre, Sphagnum angustifolium. Such forest species as Hylocomium splendens, Rhythidiadelphus triquetrus and Ptilium crista-castrensis grow on rotting wood and butts of trees at the edges of the bogs.
Lower part of mountain slopes and slopes of moraine hills and river terraces at altitudes 200-250 m above sea level are covered mainly by spruce-fir, fir and spruce-birch forests of green moss type. Herb-dwarf shrub layer is well developed and dense. Its species composition widely varies depending on conditions of the habitats. The most abundant species are Vaccinium myrtillus and Gymnocarpium dryopteris, in moss cover - Pleurozium schreberi and Hylocomium splen-dens as well as Aulacomnium palustre, Polytrichum commune, Sphagnum girgensohnii, Sanionia uncinata, Dicranum fuscescens, Ptilium crista-castrensis and species from genus Brachythecium grow on rotting wood and butts of trees.
Spruce and fir-spruce forests of herb type occur in water flow depressions. Dryopteris expansa dominates in herb-dwarf shrub layer of such communities. Other species that demand to richness and humidification of soil increase their abundance comparing with species typical for spruce green moss forests: Angelica archangelica, Athyrium filix-femina, Cirsium heterophyl-lum, Crepis paludosa, Geranium albiflorum, Milium effusum, Phegopteris connectilis, Rubus chamaemorus and Stellaria bungeana. Mosses cover up to 30 % of soil surface. Hylocomium splendens, Pleurozium schreberi and Plagiomnium ellipticum are most abundant on soil, on fallen trees - species from genus Brachythecium. Calliergon richardsonii, Pohlia wahlen-bergii, Plagiomnium ellipticum, Rhizomnium magni-folium, Brachythecium mildeanum and Sphagnum rus-sowii occur on soil in the overlogged sites.
Brachythecium rivulare, Calliergon cordifolium, Rhizomnium pseudopunctatum, Bryum weigellii, Cli-macium dendroides and Warnstorfia exannulata are often registered in overlogged depressions in spruce herb-sphagnum forests in the valleys of the streams. Sphagnum mosses can also form closed cover on the soil surface: Sphagnum capillifolium, S. squarrosum and S. russowii. Total projective cover of the mosses reached 75-80 %. Butts of stems of spruce and birch are often covered by Climacium dendroides, Pleurozium schreberi, Hylocomium splendens, Sanionia uncinata, Rhythidiadelphus subpinnatus, Aulacomnium palustre, Sphagnum capillifolium and some other species. Pleurozium schreberi, Sanionia uncinata, Di-cranum fuscescens and Hylocomium splendens cover stumps and fallen trees.
Birch-spruce and spruce forests of green moss-sphagnum type are common on the watersheds in
List of mosses species of the Malyi Patok River basin (Yugyd Va National Park, Subpolar Urals)
№/№ Species Collecting localities* Habitats Geographic al element Ecological element
1. Andreaea rupestris Hedw. 1 on stones aa mx
2. Aulacomnium palustre (Hedw.) Schwagr 1, 2, 3, 4 sedge-herb-sphagnum mires and spring bogs; spruce-birch sedge-sphagnum forest, on tussocks; floodplain willow community b hg
3. Bartramia ithyphylla Brid. 1 vicinities of the lake, along the water's edge aa m
4. Brachythecium mildeanum (Schimp.) Schimp. 2, 3 floodplain birch forest, herb-moss spruce forest, on soil b mhg
5. Brachythecium rivulare Bruch et al. 1, 2, 3, 4 birch-spruce herb-grass forest near the stream, on stones; large herb-moss spruce forest; stony herb-sedge-moss towpath community with streams; on bank of the stream b hg
6. Bryum weigelii Spreng. 1, 2 birch-spruce herb-grass forest near the stream, on stones; on the bank of the stream b hg-hd
7. Bucklandiella heterosticha (Hedw.) Bednarek-Ochyra et Ochyra 1 near the stream, open willow stand of grassherb green moss type; vicinities of the lake, on stones m mhg
8. Bucklandiella sudetica (Funck) Bednarek-Ochyra et Ochyra 1 near the stream, open willow stand of grassherb green moss type; vicinities of the lake, along the water's edge aa mhg
9. Calliergon cordifolium (Hedw.) Kindb. 1 on the bank of the stream b hg
10. Calliergon richardsonii (Mitt.) Kindb. 3 spruce herb-moss forest, on soil ham hd
11. Calliergonella cuspidata (Hedw.) Loeske 4 stony herb-sedge-moss towpath community with streams b hd
12. Calliergonella lindbergii (Mitt.) Hedenas 3, 4, 5 stony towpath communities b hg
13. Climacium dendroides (Hedw.) F. Weber et D. Mohr 2, 4, 5 floodplain birch forest, stony herb-sedge-moss towpath community with streams; island in the watercourse, upstream the tow-path b hgm
14. Codriophorus aquaticus (Brid. ex Schrad.) Bednarek-Ochyra et Ochyra 1 vicinities of the lake, along the water's edge; near the river, swampy site, dwarf shrub community dominated by willow and Betula nana m hd-hg
15. Dicranella grevilleana (Brid.) Schimp. 3 towpath communities ham m
16. Dicranum acutifolium (Lindb. et Arnell) C. E. O. Jensen 1 vicinities of the lake, along the water's edge aa m
17. Dicranum scoparium Hedw. 3 bilberry-fern green moss spruce forest, on soil; bilberry-fern green moss spruce-birch forest, on soil ham m
18. Dicranum flexicaule Brid. 1 near the river, swampy site, dwarf shrub community dominated by willow and Betula nana b m
19. Dicranum fuscescens Turner 1, 2 bedrock bank of the lake, on the butt; birch-spruce herb-grass forest near the stream, on stones; floodplain willow stand b m
20. Dicranum montanum Hedw. 2 floodplain birch forest b m
21. Dicranum polysetum Sw. 3 bilberry-fern green moss spruce forest, on soil b m
22. Dicranum spadiceum J. E. Zetterst. 1 near the river, swampy site and on the bank of the stream, dwarf shrub community dominated by willow and Betula nana aa m
23. Fontinalis antipyretica Hedw. 4 stony herb-sedge-moss towpath community with streams m hd
24. Helodium blandowii (F. Weber et D. Mohr) Warnst. 2, 3, 4 sedge-herb-sphagnum mires with streams; valley spruce forest ham hg
25. Hygrohypnella ochracea (Turner ex Wilson) Ignatov et Ignatova 1 along the water's edge, flooded sites ham hd
26. Hylocomiastrum pyrenaicum (Spruce) M. Fleisch 1 vicinities of the lake m hgm
27. Hylocomium splendens (Hedw.) Bruch et al. 1, 2 birch-spruce herb-grass forest near the stream, on stones; on the bank of the lake b m
28. Hymenoloma crispulum (Hedw.) Ochyra 1 vicinities of the lake, along the water's edge; near the waterfalls aa hgm
29. Kiaeria starkei (F. Weber et D. Mohr) I. Hagen 1 vicinities of the lake, near the stream, open willow stand of grass-herb green moss type; on the bank of the stream and along the water's edge; near the waterfalls аа hgm
30. Myrinia pulvinata (Wahlenb.) Schimp. 2 floodplain birch forest nm hgm
31. Niphotrichum canescens (Hedw.) Bednarek-Ochyra & Ochyra 1 on the bank of the stream m mx
32. Ochyraea duriuscula (De Not.) Ignatov & Ignatova 1 in the water and on stones аа hd
33. Ochyraea norvegica (Bruch et al.) Ignatov & Ignatova 1, 2 birch-spruce herb-grass forest near the stream, on stones; in the water, on stones аа hd-hg
34. Paludella squarrosa (Hedw.) Brid. 2, 3 sedge-sphagnum mire with streams ham hd-hg
35. Palustriella decipiens (De Not.) Ochyra 4 stony herb-sedge-moss towpath community with streams m hd-hg
36. Philonotis caespitosa Jur. 1 on the bank of the stream; flooded sites on the bank of the lake; near the waterfalls b hg
37. Philonotis fontana (Hedw.) Brid. 4 stony herb-sedge-moss towpath community with streams b hg
38. Philonotis tomentella Molendo 1 near the stream, open willow stand of grassherb green moss type; on the bank of the lake аа hg
39. Plagiomnium ellipticum (Brid.) T. J. Kop. 1, 2, 3, 4 floodplain birch forests, birch-spruce and spruce herb-moss forests, on soil, on stones; stony towpath with streams; floodplain willow stand b hgm
40. Pleurozium schreberi (Brid.) Mitt. 2 birch-spruce herb-grass forest near the stream, on stones; bedrock bank of the lake, on butts of trees b m
41. Pohlia cruda (Hedw.) Lindb. 1 on the bank of the lake b m
42. Pohlia ludwigii (Spreng. ex Schwägr.) Broth. 1 near the stream, open willow stand of grassherb green moss type; along the water's edge m m
43. Pohlia nutans (Hedw.) Lindb. 1 on the bank of the lake b m
44. Pohlia wahlenbergii (F. Weber et D. Mohr) A. L. Andrews 1, 3, 4 stony herb-sedge-moss towpath community with streams; spruce herb-moss forest, on soil; open willow grass-herb green moss community near the stream; on the bank of the stream, along the water's edge; in the water, on stones, in flooded sites ham hg-hd
45. Polytrichastrum alpinum (Hedw.) G. L. Sm. 1, 2 floodplain birch forest; open willow grassherb green moss community near the stream аа m
46. Polytrichastrum sexangulare (Floerke ex Brid.) G. L. Sm. 1 on the bank of the stream and along the water's edge, on stones; near the waterfalls аа m
47. Polytrichum commune Hedw. 1, 2 floodplain birch forest; on the bank of the stream b hgm
48. Polytrichum juniperinum Hedw. 1 swampy site, dwarf shrub community dominated by willow and Betula nana b xm
49. Polytrichum piliferum Hedw. 1 on the bank of the stream and along the water's edge b mx
50. Polytrichum strictum Brid. 2 spruce-birch sedge-sphagnum forest, on hummocks b hgm
51. Pseudobryum cinclidioides (Huebener) T. J. Kop. 3, 4 floodplain spruce and birch-spruce herb-moss forests; stony herb-sedge-moss tow-path community with streams ham hd-hg
52. Ptilium crista-castrensis (Hedw.) De Not. 2 bedrock bank of the lake, on butts of trees b m
53. Pylaisia polyantha (Hedw.) Bruch et al. 2 floodplain forest, on stems of trees nm m
54. Racomitrium lanuginosum (Hedw.) Brid. 1 along the water's edge аа xm
55. Rhizomnium magnifolium (Horik.) T. J. Kop. 3 spruce herb-moss forest ham hgm
56. Rhizomnium pseudopuncta-tum (Bruch et Schimp.) T. J. Kop. 2 birch-spruce herb-grass forest near the stream, on stones ham hg
57. Rhodobryum roseum (Hedw.) Limpr. 2 birch-spruce herb-grass forest, herb-moss forest, on soil b m
58. Rhytidiadelphus subpinnatus (Lindb.) T. J. Kop. 2 floodplain birch and spruce-birch herb-grass forests, on soil, on stones b m
59. Rhytidiadelphus triquetrus (Hedw.) Warnst. 5 island in the watercourse, on soil b m
60. Rhytidium rugosum (Hedw.) Kindb. 5 island in the watercourse, on stones ham xm
61. Sanionia uncinata (Hedw.) Loeske 2 floodplain birch forest; bedrock bank of the lake, on butts of trees b m
62. Sphagnum angustifolium (C. E. O. Jensen ex Russow) C. E. O. Jensen 2, 3 sphagnum mires, on hummocks, in swampy hollows; spruce-birch swampy and valley spruce forests, on hummocks b hg
63. Sphagnum balticum (Russow) C. E.O. Jensen 2 spruce-birch sedge-sphagnum forest, on hummocks b hg
64. Sphagnum capillifolium (Ehrh.) Hedw. 2, 3, 4 sphagnum mire, spring bog, on hummocks, in swampy hollows; birch-spruce herb-grass forest, herb-moss forest, floodplain spruce forest, on stones, on hillocks b hg
65. Sphagnum centrale C. E. O. Jensen 3 watershed mire dominated by Eriophorum, Carex and mosses, in swampy hollows b mhg
66. Sphagnum flexuosum Dozy et Molk. 2 hummock-ridge sphagnum mire, on hillocks b hg
67. Sphagnum fuscum (Schimp.) H. Klinggr. 2 spruce-birch sedge-sphagnum forest, on hillocks; hummock-ridge sphagnum mire, on hillocks b hg
68. Sphagnum girgensohnii Russow. 1, 2, 3, 4, 5 spruce-birch herb-grass forest, on soil; spruce dwarf shrub green moss forest, on soil; birch-spruce dwarf shrub sphagnum forest near the mire, on soil; birch-spruce fen green moss forest, on soil; edge of lowland bog; on the bank of the stream; near the waterfalls b hg
69. Sphagnum jensenii H. Lindb. 3 floodplain spruce forest; watershed mire dominated by Eriophorum, Carex and mosses, in swampy hollows ha hd
70. Sphagnum lindbergii Schimp. 1 on the bank of the lake ha hg
71. Sphagnum magellanicum Brid. 2, 3 sedge-sphagnum mire, spruce-birch sedge-sphagnum forest, valley spruce forest, on soil, on hillocks b hg
72. Sphagnum majus (Russow) C. E. O. Jensen 2 hummock-ridge sphagnum mire, on hillocks b hd
73. Sphagnum riparium Angstr. 2 hummock-ridge sphagnum mire, on hillocks b hg-hd
74. Sphagnum russowii Warnst. 2, 3, 4 sphagnum mires; birch-spruce herb-grass forest near the stream; spruce herb-moss forest, on stones, on hillocks b hg
75. Sphagnum squarrosum Crome 4 birch-spruce herb-moss b hd
76. Sphagnum teres (Schimp.) Angstr. ex Hartm. 4 herb-sphagnum mire b hg
77. Sphagnum warnstorfii Russow 1 on the bank of the lake b hg
78. Straminergon stramineum (Dicks. ex Brid.) Hedenäs 1, 2, 3, 4 spruce-birch sedge-sphagnum forest, on hillocks, watershed swampy mire dominated by Eriophorum, Carex and mosses, in swampy hollows; stony herb-sedge-moss towpath community with streams; on the bank of the stream, on the bank of the lake, along the water's edge, in flooded sites b hg-hd
79. Tomentypnum nitens (Hedw.) Loeske 2, 3 sedge-sphagnum mire, spring bog ham hg
80. Warnstorfia exannulata (Bruch et al.) Loeske 1, 3, 4 watershed swampy mire dominated by Erio-phorum, Carex and mosses, swampy hollows; birch-spruce herb-moss forest, valley spruce forest; stony herb-sedge-moss tow-path community with streams; open willow grass-herb green moss community near the stream; on the bank of the lake; near the waterfalls b hg-hd
Geographical element: aa - arctic-alpine, b - boreal, m - mountain, ha - hypoarctic, ham - hypoarcto-montain, nm - nemoral.
Ecological element: hd - hydrophyte (гидрофит), hd-hg - hydro-hygrophyte (гидрогигрофит), hg - hygrophyte (гигрофит), hg-hd - hygro-hydrophyte (гигрогидрофит), hgm - hygromesophyte (гигромезофит), xm -xeromesophyte (ксеромезофит), m - mesophyte (мезофит), mhg - mesohygrophyte (мезогигрофит), mx -mesoxerophyte (мезоксерофит). *Collecting localities are given in figure
habitats with stagnant humidification for example at the edges of mires. These communities have well developed herb-dwarf shrub layer, but not as diverse as in herbal or herb-sphagnum forests. The main dominants of such communities are Vaccinium myrtillus and Chamaepericlimenum suecicum as well as Equisetum sylvaticum and Gymnocarpium dryopteris. Other species mostly belonging to the "spruce" eco-coenotical group are found with low abundance. Mosses cover 40-50 % of soil surface. Large clumps of Sphagnum girgensohnii interchange with clumps of green mosses dominated by Pleurozium schreberi.
Flat and depressed areas of the watersheds are characterized by overlogging that result in inundating of different intensity. In such sites, spruce forests of green moss-sphagnum and herb-hypnum types are mixed with arrays of hummock-ridge mesotrophic mires. Dwarf shrubs in lower layers are displaced by hydrophilous herbs (Calamagrostis purpurea, Equisetum sylvaticum, Carex globularis, C. cespitosa, Filipendula ulmaria, Bistorta major and Ranunculus repens). Poly-trichum commune, Plagiomnium ellipticum and sphagnum mosses increase their coenotical role.
Mires can be open or afforested. Single stag-headed trees Betula pubescens, Picea obovata u Pinus sibirica were noted at the edges of mires. Central parts of mires are often treeless. Shrub layer is well developed in the elevated sites and dominated by Betula nana with mixture of Salix lapponum and S. myrtilloides. Dwarf shrub-herb layer is often dominated by Carex rostrata or C. lasiocarpa. Oxycoccus palus-tris, Andromeda polifolia, Vaccinium uliginosum and Empetrum hermaphroditum are also abundant. Moss layer is dominated by sphagnum mosses. The most common dominant is Sphagnum angustifolium. The following species can be met in sites with average humidification as well as on the elevated sites (often near stems of Betula nana): Sphagnum angustifolium, S. fuscum, S. magellanicum, S. capillifolium, S. russowii, Aulacomnium palustre, Straminergon stramineum, Polytrichum commune and P. strictum, rarely - Sphagnum. flexuosum. All the species also grow in watered depressions where they form closed moss cover with Sphagnum jensenii, S. majus, S. flexuosum, Straminergon stramineum, Warnstorfia exannulata and Polytrichum commune. Swampy hollows are filled with water. Herb layer is not as dense as on the ridges. Carex limosa, C. paupercula, Eriophorum russeolum, Scheuchzeria palustris and Menyanthes trifoliata are the most abundant species here. Such species as Carex chordoriza, C. elongata, Cicuta virosa, Drosera anglica and Pedicularis palustris are common in the depressions of relief. Hummocks up to 60 cm in height are formed mostly by Sphagnum fuscum with a mixture of S. angustifolium, S. magellanicum, S. capillifolium, S. russowii, Polytrichum strictum, Aulacomnium palustre and Straminergon stramineum. Species diversity of mosses growing on rotting wood is lower in mesotro-phic and oligotrophic mires comparing with lowland bogs. This substrate is mainly covered by Pleurozium schreberi, Polytrichum strictum and Pohlia nutans. The Lake Patok is located in the upper course of the Malyi Patok River at 770 m above sea level where
mountain stony belt and mountain tundra belt occur. Stony gravels are common here. Among them, dwarf shrub-moss-lichen tundra communities can be met. The most common mosses of these tundra communities are Hylocomium splendens, Pleurozium schreberi, Racomitrium lanuginosum, Polytrichum piliferum and Dicranum fuscescens. Shrub tundra communities from Betula nana and Salix are spread in lower parts of the slopes, saddles and valleys. Pleurozium schreberi, Sanionia uncinata, Sphagnum russowii, S. squarro-sum, Aulacomnium palustre, Warnstorfia exannulata dominate in moss cover. Dicranum spadiceum, Kiaeria starkei, Rhizomnium pseudopunctatum, Pohlia wahlen-bergii and Polytrichastrum alpinum are also often. Grass-herb meadow communities cover banks of the rivers and streams in the valleys and on the slopes. These sites are often inhabited by Brachythecium rivu-lare, Bryum weigelii, Calliergonella lindbergii, Clima-cium dendroides and Kiaeria starkei. Species composition of mosses of spring bogs and towpaths is similar and presented by Climacium dendroides, He-lodium blandowii, Tomentypnum nitens, Paludella squarrosa, Palustriella decipiens, Brachythecium rivu-lare, Calliergonella cuspidata, Pseudobryum cinclidioi-des, Pohlia wahlenbergii and Warnstorfia exannulata. Small arrays of mesotrophic and oligotrophic mires are located at foothills and on the terraces of the river Nyamga. Here Sphagnum russowii, Sphagnum capillifolium, S. angustifolium and Aulacomnium palustre, on the hummocks - S. fuscum, in hollows - S. jensenii, S. majus are abundant.
The annotated list of mosses from the basin of the Malyi Patok River includes 80 species from 43 genera and 22 families (Table).
Studies of eco-coenotical structure of mosses showed the highest taxonomic diversity in forest communities - 55 species. The largest number of the mosses was revealed in spruce forests - 49, in communities of green moss type 8 species were found, in forests of herb type - 13 species. 42 species were registered in willow communities, 21 species - in birch forests and 16 species - in fir forests. Epigeos of ground mosses are common in all the studied forests. 42 moss species were found in mires, 13 species - in meadows.
The discovered moss species are divided into nine groups relative to the humidity of the substrate. Most of the species belong to two groups - meso-phytes (22 species or 27,5 %) and hygrophytes (21 species or 26,2 %).
Boreal moss species are the most abundant in the Malyi Patok River basin (44 species or 55 %). They are dominants of soil surface cover in many vegetation communities. Soil surface cover in spruce and deciduous forests of green moss and herb types are dominated by Pleurozium schreberi and Hylocomium splendens, green moss-sphagnum - Sphagnum gir-gensohnii and Pleurozium schreberi, herb-sphagnum -Plagiomnium ellipticum, Sphagnum girgensohnii and S. squarrosum, meadows - Climacium dendroides and Plagiomnium ellipticum, towpath communities -Calliergonella lindbergii and Calliergon cordifolium, spring bogs and mesotrophic mires - Sphagnum capil-lifolium and Sphagnum russowii.
Arcto-alpine species present 16,3 % of the list (13 species), hypoarctic-mountain species - 15 % or 12 species. Mountain species are not numerous, amount 8,7 % (7 species) and prefer stony substrates in the riverside habitats. Hypoarctic and nemoral elements are not abundant and are presented by 2 species in both cases.
Three species of the mosses found in the basin of the Malyi Patok River are included in the Red Data Book of the Komi Republic [12]. The finding of these species adds new information about distribution of these species in the Komi Republic. For example, Ochyraea norvegica was found earlier only in the basins of the Ilych river, the Elima river (Northern Urals). The species is included in the Red Data Book of European bryophytes [13]. Pohlia ludwigii is known from the literature on the Telpos-is ridge [2] and mountain Narod-naya [5]; it also was found in the basins of the rivers Ilych (Northern Urals) and Voyvozh Synya (Subpolar Urals). Polytrichastrum sexangulare was earlier collected on the mountain Otorten (Northern Ural) and the Maldynyrd ridge (Subpolar Urals).
Moreover, in riverside and mountain habitats of the basin of the Malyi Patok River, several rare for the Komi Republic species were collected - Bartramia ithyphylla, Calliergon richardsonii, Codriophorus aqua-ticus, Dicranella grevilleana, Hymenoloma crispulum, Ochyraea norvegica, Polytrichastrum alpinum.
Bryoflora of the basin of the Malyi Patok River shows low species diversity of the mosses that is connected mainly with the lack of data due to the remoteness of the territory. The data obtained extend our understanding of the flora and vegetation cover of the "Yugyd Va" National Park which is currently being assessed as insufficient and makes a significant contribution to the study of the bryoflora of the Komi Republic and Russia.
Acknowledgements
We are grateful to Dr. I. Chadin for preparing the
map.
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Статья поступила в редакцию 10.06.2015.
