Abstract
The paper presents general characteristics of climate, geomorphology, parent rocks and soils in birch forests area in Murmansk Province. Numerical classification of birch forests resulted in following groups: heath birch forests are presented by Arctostaphylos uva-ursi type, Empetrum - Flavocetraria type, Empetrum - Cladina type, and Empetrum - Hylocomium type. Empetrum - Cornus type contains the transitional characteristics of the heath to meadow birch forest. Equisetum sylvaticum type and Cicerbita alpina type belong to the meadow birch forests.
Antropogenic impact on the birch forests ecosystems is analysed in following aspects: industrial pollutions, fires, felling, reindeer grazing and recreation.
1. Introduction
Birch forests occupy about 20% of the Murmansk Province covering huge area of the plains and low mountain areas and forming a well-defined narrow band between coniferous (pine and spruce) forests and tundra in high mountains. The narrow zone of birch forest fringes the eastern and south-eastern maritime part of the Kola Peninsula, mainly on the coastal slopes, and bordering with seashore heathlands, and coniferous forests and peat bogs - from the inland countryside. The aims of this study are:
- to survey mountain birch forest ecosystems in the Murmansk area, being the most north-eastern area of Fennoskandia;
- to present results of classification of plant communities, and
- to assess the human impact.
2. Area of Investigations.
2a. Geology, geomorphology and soils.
The area of study covers most of Province’s mountains, including Khibiny, Sal’nye tundry, Keyvy, Chuna- ans Monche-tundra, and north shore of the White Sea. As far as geomorphology is concerned, the study area is subdivided into two remarkably different sections - the west and the east. The border between them can be shown as a line following Voronya river valley - Lovozero - Umbozero - to the Umba river valley. The western part is sharply rugged, with mountains attaining 800 - 1200 m, most prominent are the Sal’nye Tundry, Chuna-tundra, Monche-tundra, Khibinskie and Lovozerskie mountains. Owing to influence of recent glaciation, the mountains have flat surfaces and steep slopes, with well-developed glacial morainic deposits. The eastern part has less complicated image and consists of a range of low of ‘up-hill’ mountains (the collective name is Keyvy) situated in the central part of peninsula, which decline gradually towards the Southeast. This range formed a major division during the last glaciation, and it is the reason for its rounded and gently sloping surface.
Fig. 1. Map of Murmansk Region showing the boundaries between tundra zone, mountain birch forest zone and northern taiga, and following study areas:
1. Surroundings of Pechenga,
2. Chuna-tundra, Monche-tundra, and Salnye Tundry mountains,
3. Khibiny mountains,
4. Central Keyvy mountains,
5. Surroundings of Sosnovka,
6. Tersky Bereg.
The parent rocks in the birch forest area are of various structures and composition (Geologicheskoe..., 1958). Sub-maritime birch forests are located on maritime sediments. Mountain birches grow on multiform bedrocks. The major part of mountains and mountain systems of western region (Salnye Tundry, Chil’tald, Chuna-tundra, Monche-tundra) consist mainly of crystalline rock (granite, gneiss, granulites, and shist) which has been subject to basic and ultrabasic random intrusion. The Khibiny and Lovozersky alkaline intrusion represent the richest type of bedrocks. The Eastern mountain range - Keyvy - is composed mainly of crystalline schists. As a result of glaciation, quaternary deposits almost entirely cover the bedrock, and bare bouder beds are rather sparse in the birch forests zone. Soils derived from morainic deposits are characterised by sandy texture with a lot of stone and gravel in the soil profile.
Podzolis soils prevail in all the birch forests of Murmansk Province. But owing to rich parent bedrocks in Khibiny and Lovozersky mountains and particular climatic conditions (more rainfall and snow cover, and a longer growing season) the podzol layer formation is reduced. This soil is called Al-Fe humic podzol. The most outstanding feature of this soil is that the humus content in its mineral layers is relatively high: it amounts to as much as 5,8% in illuviale horizon, whereas the podzol horizon contains 4.1% of humus (Ushakova, 1997).
2b. Climate.
In general, the climate of the central part of Province is more continental, than the climate of the eastern and coastal parts. Meteorological data are available from the Handbook of climate of USSR (Anon. 1965, 1968) for the stations, situated in the birch forest zone.The western part has a higher precipitation, than the central and coastal area of the White Sea. Maximum annual temperatures in the western and coastal regions are less than in centre. The average annual temperature is below 0o C in the whole area, but inland and at the White Sea it is clearly colder than in the western region. The western and coastal parts of the region show evidence of maritime climate by their reduced summer rainfall (less than 35 % of the annual distribution for June, July, and August)
3. Methods.
3a. Data collection.
Field work was carried out in 1990-1999 in the Khibiny, Keyvy, Chuna-Tundra, Monche-tundra and Salnye Tundry Mts, and along the shore of the White Sea (Fig.1). 82 sample plots (10 x 10 m) were made in birch stands above the coniferous forest zone. The stands were considered uniform in floristic composition and structure. Percentage cover of each taxon was estimated using the following variant of the Braun-Blanquet scale: r: << 1% cover; + : < 1%; 1: 1-5%; 2: 6-25%; 3: 26-50%; 4: 51-75%; 5: 76-100%.
Altitude, exposition and inclination were estimated with the help of a map and compass. Height and diameter of tree species were measured, the density of canopy and cover of understorey species were estmated.
The botanical nomenclature follows Lid and Lid (1994) for vascular plants, Ignatov and Afonina (1992) and Konstantinova et al. (1992) for mosses and Santesson (1993) for lichens.
In addition, birch forest vegetation samples taken by Avrorin et al. (1935), Nekrasova (1938), Neshatayev & Neshatayeva (1993) were included. These samples were comparable to Braun-Blanquet releves, albeit the Drude and percentage scale used by these authors had to be converted to the above scale as follows: sol - very small, cover< 1% = +; sp1, cover 1-5% = 1; sp2, cover 6-20% = 2; cop1, cover 21-40% = 3; cop2, cover 41-60% = 3; cop3, cover 61-80% = 4; soc, cover 81-100% = 5.
3b. Data analyses
Numerical technique TWINSPAN, which gives both a grouping of species and classification of sample plots (Hill, 1979), had been used to analyse composition of vegetation. The analyses contains more than 110 releves (stands). Three levels of division were taken into account in order to produce the final community units. The TWINSPAN sequence of species was rearranged in order to characterize the groups of communities obtained in terms of their floristic composition and to reveal clearly at a glance the similarity and dissimilarity of samples (Tab. 1)
4. Results.
The two large groups marked on the first TWINSPAN division correspond to heath and meadow birch forests (Fig.2)..
The first unit is Arctostaphylos uva-ursi type of heath birch forests, widely distributed at low elevations, mainly on southerly plains. The tree layer is open, and policormic birches grow far apart, and the field-layer consists mainly of Arctostaphylos uva-ursi, Empetrum hermaphroditum. Lichens of genera Cladina- and Cladonia- form the ground layer.
The second unit is an Empetrum - Flavocetraria type and comprises the driest and the most infertile birch stands. They are common in the mountains of the eastern part of Province. The birches here grow in bush islands, standing far apart. Dwarf shrubs and chionophobous lichens of genera Cladina, Cladonia and Flavocetraria form scattered cover.
The third group of TWINSPAN division for birch forests is where mosses make up essential part in the ground-layer. Here birches are often in the form of more individual trees, and the stands occupy more sheltered positions and differ in appearance because of dense layer of vascular plants, great portion of green mosses and liverworts, and the constant presence of mesophilous foliose lichens in ground layer. This group is divided into sub-units, which differ in accordance with the abundance of lichen and moss in the ground layer, which are of Empetrum - Cladina and Empetrum - Hylocomium types.
A fourth group - Empetrum - Cornus type - contains the transitional characteristics of the heath to meadow birch forest. These stands occupy rather steep warm slopes with good water supplies served by numerous brooks. Polycormic oblique birches form dense tree layer, field layer consisis mainly of herbs, although dwarf shrubs are a significant component too. The ground layer is weakly developed (almost absent). Owing to their particular position and ecological conditions these stands provide shelter to some interesting rare Red Book species such as Epipactis atrorubens, Veronica fruticans, Castilleja lapponica.
Two last groups of TWINSPAN division represent meadow birch forests, which are mainly situated close to springs and rivers as well as in mountains, as in submaritime zone, where meadow forests are most prevalent. The birches here are tall, straight or oblique. Mountain ash and alder are common in tree-layer. Luxurious tall herbs, grasses and ferns form field layer and the ground layer is poorly developed. The Equisetum sylvaticum type comprises of fresh meadow birch forests situated in ravines, on well-moistered slopes in mountains, and on seashore terrains. Cicerbita alpina type comprises of more humid birch forests, situated mainly in mountains, near brooks and springs.
Whereas birch forests of Fennoscandia form oroarctic, arctic and maritime timberline (Hamet-Ahti, 1963, Ahti et al., 1968), in Murmansk Province most of birch forest form just only oroarctic timber-line. They are common on extensive plains in central and western part of Province at altitudes of 150-350 m a.s.l., where prevail lichenous, mossy, heath forests. Fresh heath and meadow birch forests are common in high mountains of Province, and are associated with springs and brooks. The submaritime birch forest are practically absent from the Barents Sea shore, owing to particular topography of this area. Meadow and heath submaritime birch forests occur only on the White Sea shore, as rather narrow zone, where they are found at elevations from 20 to 50 m a.s.l.
Tab.1. Synoptical table of mountain and seashore birch forests of the Murmansk Province. Romanic numbers mean classes of constancy (presense of species), arabic numbers mean cover degree.
Arctostaphylos uva-ursi type |
Empetrum - Flavocetraria type |
Empetrum - Cladina type |
Empetrum- Hylocomium type |
Empetrum - Cornus type |
Equisetum sylvaticum type |
Cicerbita alpina type |
|
unit |
1 |
2 |
3 |
4 |
6 |
5 |
7 |
Arctostaphylos uva-ursi |
IV+-3 |
II+-1 |
I+ |
- |
- |
- |
- |
Cladonia deformis |
V+ |
II+ |
II+ |
- |
I+ |
- |
II+ |
Cladonia pleurota |
IV+ |
I+ |
- |
I+ |
- |
- |
- |
Diphasiastrum complanatum |
I+ |
III+-1 |
I+-1 |
- |
- |
- |
- |
Flavocetraria nivalis |
II+-2 |
IV+-3 |
I+ |
- |
- |
- |
- |
F. cucullata |
I+ |
III+-3 |
I1 |
- |
- |
- |
- |
Cladina rangiferina |
V1-4 |
V+-2 |
IV+-2 |
I+-1 |
I+ |
- |
- |
Cladina stellaris |
IV+-3 |
III+-3 |
III+-3 |
- |
- |
- |
- |
Cladonia gracilis |
V+-1 |
I+-1 |
IV+-2 |
- |
I1 |
- |
- |
Cladonia crispata |
V+ |
- |
IV+-2 |
III+ |
- |
- |
- |
Nephroma arcticum |
I+ |
I+ |
III+-1 |
IV+-2 |
- |
- |
- |
Peltigera aphthosa |
I1-2 |
- |
II+-1 |
IV1-2 |
- |
- |
- |
Ledum palustre |
I+ |
I+ |
III+-1 |
IV1-3 |
I+ |
- |
- |
Dicranum majus |
II+-1 |
II+-1 |
I+-1 |
IV1 |
- |
- |
- |
Cladonia uncialis |
IV+-2 |
III+-2 |
II+ |
II+ |
I+ |
- |
- |
Arctostaphylos alpinus |
III+-5 |
III+-1 |
II+-2 |
- |
- |
- |
- |
Calluna vulgaris |
II+-5 |
I1-2 |
I+ |
I2 |
- |
- |
- |
Cladonia amaurocraea |
I1 |
II+-1 |
II+-1 |
I+ |
I+ |
- |
- |
Cladonia bellidiflora |
I+ |
I+ |
II+-1 |
I+ |
- |
- |
- |
Cladonia cornuta |
II+ |
I+ |
I+ |
III+ |
- |
- |
- |
Cladina mitis |
III+-3 |
III+-4 |
III+-2 |
III+-1 |
II+ |
- |
- |
Stereocaulon spp. |
III+-3 |
II+-2 |
II+-1 |
I1 |
I+-1 |
- |
- |
Cetraria islandica |
III+-2 |
III+-3 |
III+-2 |
III+-1 |
II+-1 |
- |
I+ |
Pleurozium schreberi |
II+-5 |
III+-5 |
IV+-5 |
V1-3 |
II2-4 |
II1 |
- |
Hylocomium splendens |
I1 |
I1 |
III+-3 |
V2-5 |
I+-1 |
III1 |
- |
Equisetum sylvaticum |
- |
- |
- |
- |
- |
IV+-1 |
- |
Bistorta vivipara |
- |
- |
- |
- |
- |
III+-1 |
- |
Veratrum album |
- |
- |
- |
- |
- |
III+-1 |
- |
Athyrium distentifolium |
- |
- |
- |
- |
I3 |
- |
IV+-5 |
Cicerbita alpina |
- |
- |
- |
- |
- |
- |
IV+-1 |
Salix glauca |
- |
I+ |
II+-2 |
- |
I+ |
- |
V+-3 |
Salix phylicifolia |
- |
I+ |
I1 |
I1 |
- |
- |
III+-1 |
Sanionia uncinata |
- |
I+ |
- |
- |
I+ |
- |
III+ |
Milium effusum |
- |
- |
I+ |
I+ |
II+ |
- |
III+-1 |
Calamagrostis lansdorfii |
- |
- |
I+ |
- |
I+ |
III+-1 |
IV+-3 |
Geranium sylvaticum |
- |
- |
II+-3 |
- |
III+-1 |
IV+-4 |
V+-3 |
Trollius europaeus |
- |
- |
I1 |
- |
I1 |
III1 |
IV+-1 |
Gymnocarpium dryopteris |
- |
- |
II1 |
- |
III+-1 |
III+-3 |
V+-3 |
Brachythecium salebrosum |
- |
- |
- |
- |
II+ |
II+ |
III+-2 |
Rhizomnium pseudopunctatum |
- |
I+ |
- |
- |
- |
II+ |
II+ |
Melampyrum pratense |
II+ |
I+ |
I1 |
I+ |
IV+ |
V+-2 |
III+ |
Trientalis europaea |
- |
I+ |
III+-2 |
IV+ |
V+-1 |
III+-1 |
IV+ |
Cornus suecica |
- |
I+-1 |
II1 |
II1 |
IV+-5 |
V1-4 |
IV+-1 |
Anthoxanthum odoratum |
- |
I+ |
II1-2 |
- |
III+-3 |
II+ |
IV+-1 |
Sorbus aucuparia |
- |
I1 |
I+-1 |
III+-1 |
II+-1 |
III1 |
V+-1 |
Betula nana |
II2-3 |
IV+-3 |
II1-2 |
III+-4 |
I+-3 |
I2 |
I+ |
Empetrum nigrum spp. hermaphroditum |
V2-5 |
V+-5 |
IV+-5 |
V1-3 |
IV+-2 |
II+-1 |
II+ |
Vaccinium vitis-idaea |
V+-3 |
IV+-2 |
IV1-3 |
V1-2 |
V+-1 |
III+-1 |
II+ |
Vaccinium uliginosum |
V1-4 |
III+-3 |
IV+-3 |
IV1-2 |
III+-5 |
III2-3 |
II+ |
Juniperus commune spp. alpina |
II+-1 |
IV+-2 |
IV+-1 |
III1-3 |
IV+-3 |
V+-2 |
III+-1 |
Betula pubescens |
V2-5 |
V1-4 |
V2-5 |
V2-5 |
V2-5 |
V2-5 |
V2-5 |
Vaccinium myrtillus |
I+ |
IV+-3 |
IV+-4 |
IV1-5 |
V+-3 |
V+-1 |
V+-1 |
Solidago virgaurea spp. minuta |
I+ |
II+ |
V+-2 |
I1 |
V+-3 |
V+-2 |
V1-2 |
Luzula pilosa |
I1 |
- |
I1 |
I1 |
II+-1 |
I1 |
III+ |
Linnaea borealis |
- |
I+ |
V+-1 |
V+-1 |
II+ |
II+ |
III+ |
Pyrola minor |
- |
- |
I+-1 |
III+ |
II+ |
- |
III+-3 |
Barbylophozia lycopodioides |
- |
II+-2 |
IV+-3 |
V1-2 |
II+-3 |
I+ |
I1 |
Lycopodium annotinum |
- |
I+ |
III+-1 |
II+ |
II+ |
II+ |
III+ |
Festuca ovina |
IV+-2 |
II+ |
V+-1 |
III+-3 |
II+ |
III+-2 |
- |
Deschampia flexuosa |
III+ |
III+-1 |
IV+-3 |
I+ |
V+-4 |
- |
V+-1 |
Picea abies spp. obovata |
- |
I+-1 |
I+ |
I1 |
II+-2 |
II+ |
I1 |
Polytrichum juniperinum |
II+-1 |
I+ |
II+-3 |
I+ |
II+-3 |
- |
II+-1 |
Polytrichum commune |
II+-1 |
I+ |
IV+-3 |
I+ |
II+-3 |
III+-2 |
- |
Brachythecium reflexum |
- |
- |
I+ |
II+ |
I1 |
II+ |
I+ |
Carex bigelowii |
- |
III+-1 |
III+-2 |
- |
- |
- |
- |
Dicranum fuscescens |
- |
I+-2 |
II+-1 |
I+ |
- |
- |
- |
Cladonia furcata |
I+ |
- |
I+ |
I+ |
- |
- |
- |
Lophozia groenlandica |
- |
I+ |
I+ |
II+ |
- |
- |
- |
Dicranum scoparium |
I1 |
II+-1 |
II+-1 |
- |
- |
- |
I1 |
Dicranum congestum |
- |
I+-2 |
III+-3 |
I1 |
II+ |
- |
I+ |
Cladonia coccifera |
III+-1 |
I+-1 |
II1 |
- |
II+ |
- |
- |
Rubus saxatilis |
- |
- |
I+ |
I+ |
I+ |
- |
- |
Orthilia secunda |
- |
- |
II+-1 |
I+ |
I1 |
- |
- |
Peltigera canina |
- |
- |
II+ |
- |
I+ |
- |
- |
Omalotheca norvegica |
- |
- |
I+-1 |
- |
- |
- |
I+ |
Phyllodoce caerulea |
- |
II+-1 |
II+-3 |
- |
II+-3 |
- |
I1 |
Cladonia fimbriata |
I+ |
- |
II+ |
- |
I1 |
- |
- |
Bartsia alpina |
- |
I+ |
I+ |
- |
- |
- |
I+ |
Hieracium sp. |
- |
II+-1 |
II+-1 |
- |
I+ |
- |
II+ |
Populus tremula |
- |
II+-1 |
II+-1 |
- |
I+ |
- |
- |
Plagiothecium laetum |
- |
I+ |
I+ |
- |
I+ |
- |
- |
Orthocaulis floerkei |
- |
I+ |
I+ |
- |
- |
I+ |
- |
Antennaria dioica |
I+ |
- |
I+ |
- |
I+ |
- |
- |
Cladonia ecmocyna |
I+ |
I+ |
I+ |
- |
I+ |
- |
- |
Cirsium helenioides |
- |
- |
- |
I1 |
I+ |
- |
- |
Pedicularis lapponica |
- |
I+ |
II+-1 |
II+ |
I+ |
- |
II+ |
Salix caprea |
- |
I+ |
II+-1 |
- |
I+ |
- |
II+ |
Crepis paludosa |
- |
- |
I+ |
- |
- |
- |
I1 |
Pyrola rotundifolia |
- |
- |
I+ |
- |
II+ |
- |
- |
Salix lanata |
- |
- |
I1 |
- |
I1 |
I1 |
I1 |
Cladonia chlorophaea |
II+ |
- |
I+ |
- |
- |
- |
- |
Peltigera scabrosa |
I1 |
- |
I+ |
- |
- |
- |
- |
Bryocaulon divergens |
- |
II+ |
I1 |
- |
- |
- |
- |
Alectoria nigricans |
- |
I+ |
I2 |
- |
- |
- |
- |
Alectoria ochroleuca |
- |
I+ |
I+ |
- |
- |
- |
- |
Pinus sylvestris |
I4 |
I+-1 |
I1 |
- |
- |
- |
- |
Juncus trifidus |
- |
II+-1 |
- |
- |
- |
- |
- |
Alnus incana |
- |
- |
- |
- |
- |
- |
I1 |
5. Human impact.
In Murmansk Province the birch forests cover a large area which is very different with regard to local population density and level of industrial development, that determines the character and pressure of antropogenic impact. The most populated, most trafficked, most industrial sectors are the central part of Province, situated along the road Petersburg-Murmansk, and north-western part. The eastern part of Murmansk Province is almost uninhabited, and there is situated the main pasture area of reindeer husbandry.
Antropogenic impact on the birch forests ecosystems can be analysed in following aspects: Industrial pollutions, Fires, Felling, Reindeer Grazing and Recreation.
5a. Industrial pollutions.
Industrial development in Murmansk Province started from 1930’s, and nowadays numerous plants and mines ores cause essential damage to biota. Concern ‘Apatity’ produce fertilizers, etc. from apatit ores in Khibiny mountains. Copper-nickel smelters of the concern ‘Severonikel’ and ‘Pechenganikel’ operate with recovering of copper, nickel, cobalt as the metals. Vaste gases containing sulphur dioxide and dust are considered to be the major damaging factors, caused deforestation of surrounding of Monchegorsk, Nikel and Zapoljarnyi. Vastes of aluminium plant in Kandalaksha, ore-developing factories in Kovdor and Olenegorsk are lower, but contribute much to the tree-line ecosystem degradation in neighboring mountains.
5b. Fires and Felling usually accompany the industrial impact overall in zone of forest degradation. Although the felling is of minor occurrence in the timberline area, the creeping fires almost entirely (and sometimes repeatedly) cover the area adjoining industrial centers. They have dramatic effect, when ground layer and litter burn, bare soil slim downhill, and reveal bare mineral moraine horizon. This leads to complete ecosystem degradation, and fires even are reported to be a leading factor affecting the forest degradation in the industrial zone (Selikhovkin, 1993).
5c. The keeping of reindeer herds provides a traditionally form of subsistence for the Saami peoples. The wild reindeer population amounts to around 16 thousand whereas the tame reindeer population is around 70 thousand. Although these populations cover various areas, tame reindeer are tended under more or less the same natural conditions as wild reindeer (as far as the Saami people are concerned). Their horizontal migration is dictated by the availability of fresh pastures. During the summer, the reindeer disperse over a wide range of forest territory, because at this time they favour the leaves and twigs of the mountain birch as well as the various grasses and herbs that are available for grazing. Cladina-lichens, for example, are only eaten when wet. The damage to the vegetation caused by reindeer tramping around seems to be minimal, because they only use their migration route paths. These paths cover an extensive area and provide relief from insects for the reindeer. The winter pastures are situated mainly on the plains and have a predominance of Cladina vegetation. Tame reindeer are concentrate near the villages during winter and, as a result, winter pastures around the villages become overgrazed. This has resulted in the establishment of grasses, such as Nardus stricta, Deschampsia cespitosa and D. flexuosa.
The ‘Nenets’ way of tending to herds, when the summer flock is kept within close confines on limited pastures - protected by dogs - leads to important overgrazing of pastures near settlements and villages. However, in comparison to the situation in Finish-Lapland, where desertification of the pastures due to overgrazing is a major problem, the reindeer business in Kola is much less intensive and damaging.
5d. Recreation and reconnaissance. Surroundings of industrial centers and distant sites are similarly popular destination of some number of people - either for a rest, or for a exploring activity. Skiing, camping, trampling have locally detrimental effect on vegetation and soil. Under natural circumstances after decreasing of the pressure goes rehabilitation in plant cover. As the most successive invaders were reported graminoids, apocarpic mosses, and fruticose and foliose lichens. Primary succession on this small-scale disturbances results in some increasing of species diversity comparingly with sample plots from adjoining plant cover. The damage caused by off road vehicular and tracked traffic, local drilling and local fires, especially caused by pouring of dizel fuel, are potential for seious degradation of landscapes.
Conclusion
Although a lot of studies have addressed the investigations of birch forest as a part of plant cover of in Murmansk Province, numerous unsolved problems relating to birch forest ecosystems still exist. On the one hand, Betula pubescens were reported as a species, highly resistant to aerial pollution, whereas on the other hand, birch timberline ecosystems are referred to as the most sensitive in relation to damage caused from environmental pollution. To make comprehensive study of the birch forests ability to resist and regenerate after disturbance caused by pollution, we need:
- firstly, to use the available background information on their structure and composition relative to the habitat;
- secondy, to examine all possible birch forest ecosystem parameters, such as the coenopopulation structure and properties, the litter and soil composition and processes, the microbiota characteristics, and so on;
- lastly, to organize the investigations along gradients of the main ecological factors.
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