Non-­saline Meadows and Grasslands in the Kola Peninsula

Meadows and grasslands attract attention of researchers for a long time. And their defini--‐ tions were changing in the course of history of the vegetation science. At this moment the most complete definition is from T. A. Rabotnov [1, 1984]. He determined meadows as biogeocoenoses with more or less closed cover of grasses, perennial mesophytes, with winter pause or sharp de--‐ crease in growth due to temperature drop, and without summer depression in growth due to lack of water supply. A. P Shennikov divides meadow to glykophytic, or non--‐saline (without halophytes) and halophytic (with halophytes adapted to life in salinity of soil > 5%). Coastal meadows of the White and Barents Seas in the mouth of rivers constitute a specific zone of "Arctic maritime and floodplain meadows" [2, 1941].

This article presents the data on history and present state of knowledge about glycophytic (non--‐saline) meadows on the Kola Peninsula, taking into account that these plant communities are less studied than coastal halophytic meadows and marshes ([3, 2002], [4, 2008], [5, 2011], [6, 2012] et al.)

Geographical and climatic features of the Murmansk region which influence meadows distribution

The Murmansk region occupies the northeastern part of the Baltic shield composed by Pre--‐ cambrian crystalline rocks. Frequent outcrops of acidic granites and gneisses of the Archean or Proterozoic era are covered by shallow glacier moraine. Morainic deposits represent main soil--‐ forming nutrient--‐poor parent rocks. Tectonic movements in the Proterozoic and Paleozoic era formed modern mountainous topography and determined characteristics of hydrology of the Murmansk region --‐ most rivers flow on the bottoms of tectonic faults and fractures, often their flow includes large lakes.

Network of rivers and lakes of the Kola Peninsula is young. Alluvial deposition occur in the mouths of large rivers, they are represented by layered clays, sands and pebbles [7, 1977]. Most of the major rivers in the north of the Peninsula (Teriberka, Voronya, Kharlovka, Vostochnaja Litca, Iokanga, Lumbovka) have a narrow floodplain. Parts with quiet and rush flow alternate each other, numerous falls and cascades occur, and due to this features area of floodplain meadows is small. The lower flow of large rivers flowing to the White Sea (Varzuga, Ponoy, Umba) sometimes have well--‐developed floodplain. Estuaries of the rivers are filled with salt Sea water during high tides, and this influences the floodplain vegetation.

Climate of the Murmansk region is determined by position in the Atlantic--‐Arctic Climatic zone, with frequent inflow of warm air masses from the Atlantic Ocean and intense cyclonic activi--‐ ty. Climate of area is humid with a predominance of precipitation over evaporation due to moder--‐ ate temperatures and high relative air humidity. Humid climate is an important factor in the for--‐ mation of meadows and grasslands.

Zonal vegetation types on the Kola Peninsula include northern taiga, subarctic birch forest and tundra. Zonal types were formed about 7500 years ago, after glaciers retreat [8, 2009], [9, 2007].

The composition of rocks, the hydrographic network and relative youth of plant cover de--‐ termine the small area covered by meadows and grassland and low proportion in plant cover of the Murmansk Province, compared with neighboring regions (such as Karelia and Arkhangelsk Re--‐ gion). But the diversity of meadows is high. Specific features of meadows in Murmansk region are resulted from their location at the northern limit of the riparian vegetation and in different natural zones in lowland and in mountains (from taiga to tundra). Although meadows represent a classic example of intrazonal communities but zonal position of non--‐saline meadows and grasslands de--‐ termine their composition and structure.

Non--‐saline meadows and grasslands on the Kola Peninsula include shortgrass early snow--‐ bed meadows in tundra zone in mountains, meadows of floodplain along rivers valleys, meadows on the banks of lakes, coastal meadows on the high shores, secondary (abandoned agricultural, including seeded) grasslands, and ornithogenic meadows nearby bird colonies (Fig. 1). Natural up--‐

Fig. 1. Non--‐saline meadow communities in the Varzuga river valley (flood--‐plain meadows) with dominance of Hedysarum arcticum (left) and Polemonium caeruleum (right)

land meadows in the region are absent [10, 1983].

Modern abandoned agricultural grasslands are widespread near settlements, where they were established after forests cuttings and bog melioration followed by sowing of grasses. Low--‐ land abandoned agricultural grasslands are situated in drained depressions and river valleys. Up--‐ land meadows are situated on watersheds and near settlements. Small patches of shortgrass early snowbed meadows are widespread in tundra zone of mountains. They are characterized by high portion of tundra dwarf shrubs. Meadows of flood plain look like narrow strips along rivers or on river islands both in the forest and tundra zones [11, 2011]. They were used by villagers as pas--‐ tures and hayfields. Now the area of this meadows and grasslands decreases and they persist only near the large settlements (Luvenga, Varzuga, Tchapoma, etc.). Meadows on the banks of the lakes have a limited distribution. This type of meadows must be differentiated from the grasses--‐ dominated fens (mires). Coastal meadows on the high seashores occur on rocky, sandy, loamy and clayey shores of the White and Barents Seas. The most extensive they are in the mouths of large rivers on river alluvia and on the coastal terraces. Ornithogenic nitrophilous meadows occur on the rocks near birds’ nests or colonies and are similar in composition with halophytic coastal meadows.

The typology and classification of meadows and grasslands

Meadows and grasslands is such a kind of vegetation where fundamental problems of geo--‐ botany and phytocoenology were tested. Science about meadows and grasslands (‘Lugovedenie’)

arose as a science to solve specific problems of these plant communities, for example, the spatial structure of community, long--‐term and short--‐term dynamics, relationships between plants and layers of community and interactions with other components of the biogeocoenose (such as mi--‐ crobes and fauna). One of the fundamental issues of this science is typology and classification of meadows and grasslands. These plant communities don’t have specific methods of classification. But each classification method applied to meadows and grassland vegetation had to solve special problems. The contradiction between the main purpose of classification and typology (to distin--‐ guish types of plant communities) and continual nature of spatial (horizontal) structure of flood--‐ plain meadows even resulted in denial of the validity of classification as a method of vegetation study. Leontij Ramenskyi (1884--‐1953) author of a theory of environmental coordination of mead--‐ ows [12, 1956] thought that classification method isn’t of use in continuously varying meadow plant cover.

At the beginning of 20 century Russian science about grasslands and meadows (lugovedenie) traditionally practiced a dominant approach to classification, with such basic units as formation and association. Soviet botanist Alexandr Shennikov (1888--‐1962) [2, p. 285--‐286] dis--‐ tinguished association basing on similar flora, structure and habitat (e.g., soil condition). For ex--‐ ample, associations Festucetum rubrae rhinanthosum , F. r leucanthemosum , F. r troliosum and etc. The group of associations with similar dominants (edificators) were united in formation, i.e. --‐ Alo--‐ pecureta pratensis , Parvoherbeta mixta and etc.

Marianna Ramenskaya (1915--‐1991) [13, 1958] practiced ecological and phytocenotic classi--‐ fication of meadows and grasslands of Murmansk Region and Karelia. She considered meadow as--‐ sociation as some group of phytocenoses with identical morphology, phytocenose structure and flora, and with common dominants and co--‐dominants (usually in number from 2 to 4). This species determine the structure of plant community and form a large part of biomass. Associations with dominance of one or several species form a formation (e.g., formations Nardeta strictae , Ag--‐ rosteta vulgaris and Humidiherbeta . This classification was used some scientists who practiced dominant method.

Method of dominants together with composition of layers and synusiae and temporary edificators was in early--‐ and mid last century the most common approach in the study of grassland in Russia [14, 1910], [15, 1928], [16, 1964]. But in meadows with several dominants, with season--‐ ally and permanently changing composition of dominants, this approach led to the establishment of several different classification schemes which do not correspond to the real diversity of mead--‐ ows. This approach was useful only for classification of mono--‐dominant communities with con--‐ stant composition --‐ such as i.e. seashore marches.

In 20 century there was arisen Braun--‐Blanquet approach to classification in Central and Southern Europe, and after 1980ths has been widely distributed in Russia. Its advantages is use of floristic combination and diagnostic species for combining communities into associations and then arrange associations in the hierarchy. “Old” syntaxa can change their position and rank in the hier--‐ archy, and new syntaxa can be introduced in existing scheme, etc. [17, 2012]. This method is wide--‐ ly used in meadows and grasslands. In Prodromuses (list of syntaxa) of European countries the part of communities with dominance of grasses and herbs is very significant. For example, in Pro--‐ dromus of Russia [17, p. 377] unions of meadows, steppes, forest edges take approximately 20% of the total list. In Northern Europe associations of meadows, tundra meadows and clearings communities constitute 24% of the entire list of syntaxa [18, 1996], in the Prodromus of Czech Re--‐ public associations of steppe meadows, wet and dry meadows and forest edges communities ac--‐ count for about 25% of the list [19, 2003].

In the typology of vegetation In Fennoscandia (Finland, Norway) there is practiced ecoligi--‐ cal--‐and--‐dominant approach in classification of meadows. This method is based on the grouping of communities according to the environmental conditions of habitat, dominants and characteristic species [20, 1994], [21, 2003]. To characterize the community--‐type there is used diagnostic group of species as in method of Braun--‐Blanquet.

Modern classification of meadows and grasslands are based on mathematical methods (such as coordination and clustering) with use of special computer programs (TWINSPAN, Coctail, JUICE, Graphs, etc.). The results of such classifications are interpreted involving methodology of Braun--‐Blanquet or typology basing on habitats and species composition [17, P. 211].

History of the study meadow and grassland communities of the Murmansk region

Study of meadows and grasslands in the Murmansk region started in late XIX --‐ early XX cen--‐ turies. It was the time of first active botanical expeditions (including study of meadow and grass--‐ lands together with other plant communities) and first lists of vascular plants of Russian Lapland [22, 1864], [23, 1831], [24, 1882]. There were published some general reviews of the investigation of Murmansk Province vegetation, including seashore meadows [25, 1953], [26, 1981], [27, 2012].

• K.    Regel [28, 1922] presented the first descriptions of plant cover of Kola Peninsula includ--‐ ing descriptions of coastal meadows, meadows of flood plain, early snowbed meadows in tundra zone in mountains. Descriptions were performed during field routes and contain only the most

dominant and characteristic species. It is difficult to compare them with present complete descrip--‐ tion of meadows and grasslands. For example, the association of Regel Anthoxantho--‐Nardetum stricti described in the alluvial terrace on the left bank of Varzuga includes only 7 species.

Yu. D. Zinserling [29, 1935] explored the eastern part of the Kola Peninsula (river valleys of Iokanga, Ponoy, Strelna, Pyalitsa, Lumbovka) from 1927 to 1928 and classified meadows and grass--‐ lands based on dominants and position of communities in landscape. Zinserling divided all non--‐ saline meadows and grasslands on the Kola Peninsula to floodplain meadows, early snowbed meadows in tundra zone in mountains, upland meadows and paludified meadows. According Zin--‐ zerling, diversity of floodplain meadows is determined by geographic position of the river (wheth--‐ er the meadow lies on the area of the floodplain riverbed, in the middle part of a river terrace or on the slope of the eroded by water bank), by differences in the hydrology and the nature of allu--‐ vial processes, by differences of floodplain topography and human activities. So, running streams of the Kola Peninsula with narrow valleys and short--‐term flood have only terrace and riverbed zones. Such a meadows Zinserling has described along rivers Strelna, Iokanga and Ponoy.

Zinserling classified continental meadows of Northwestern Russia basing on the classifica--‐ tion approach of V. N. Sukachev and distinguished three classes (meadows on poor soils, meadow on soils of average fertility and meadows on rich soils). These three classes were subdivided ac--‐ cording to the level (gradient) of the soil moisture.

Early snowbed meadows occur only in the tundra of the Kola Peninsula, on well moistened slopes on the fine earth, often influenced by melting water from late--‐melting snow patches and permanent snow field, with good supply of water. The growing season starts here later compared to the surrounding tundra. Some grasses, like Anthoxanthum alpinum, Nardus stricta , herbs Vera--‐ trum lobelianum , Trollius europaeus , Geranium silvaticum , Alchemilla spp., Viola biflora etc. pre--‐ vail here.

• N.    M. Savitch studied meadows and grasslands in the central and southern parts of the peninsula (floodplain meadows of rivers Tuloma and Kola, coastal meadows of Kandalaksha Bay and secondary grasslands along the railway from Kola to Kandalaksha) [30, 1926]. Savitch based of the dominant approach and described three formations and 11 associations on the base of species composition. So, grass--‐and--‐forbs formation included 4 associations: Agrosticetum albae , Festucetum rubrae on sandy soil, Agroryretum repens and Elymusetum arenarius . Formation of paludified meadows comprised associations Caricetum norvegicae , Heleocharicetum palustris , Juncetum gerardi , Festucetum rubrae on muddy substrate, Calamagrosticetum neglectae, Carice--‐ tum sainae and Caricetum imosae . The latter two associations were described on the shoreline of

Kandalaksha Bay and in the central parts of the Kola Peninsula. In the taiga zone Savitch described formation of forest meadows.

Some descriptions of the floodplain meadows were presented in the survey of reindeer pastures of Kola Peninsula [31, 1936]. There are following types: shortgrass early snowbed mead--‐ ows in tundra zone, riverine meadows of "low level" and meadows on the middle terrace above the floodplain of rivers Nota, Lotta, Tuloma and other rivers of north--‐west part), antropogenous “continental meadows" located nearby fishers’ and Sami huts, secondary grasslands in areas after birch cutting and coastal meadows.

The monography of R. Kalella [32, 1939] contains the tables of descriptions and character--‐ istics of communities with the dominance of herbaceous perennials on the Rybachiy (Fisher) pen--‐ insula, former Finnish territory. Herbal community on the Rybachiy were divided into following types: slope meadows, tall--‐grass meadows, alluvial meadows, paludified and riverbank meadows and grasslands, meadows on sandy substrates, marches, snowbed meadows with dwarf shrubs. Kalela noticed the narrow floodplains and thin alluvial deposits in valleys of small rivers of the Rybachiy Peninsula. Kalela divided alluvial meadows to following types: Equisetum arvense--‐Wiese, Calamarostis neglecta--‐Wiese, Equisetum fluviatile--‐Wiese, Carex aquatilis--‐Wiese, Juncus filiformis--‐ Wiese, Caltha palustris--‐Wiese, Ranunculus repens--‐Wiese, Petasites frigidus--‐Wiese, Filipendula ul--‐ maria--‐Wiese, Ranunculus reptans--‐Soziation, Alopecurus aquatilis--‐Soziation, Subularia aquatica--‐ Soziation, Carex vesicaria--‐ and Carex lasiocarpa--‐Überschwemmungsmoor, Carex caespitosa--‐ Zsombeck--‐Moor.

• M.    L. Ramenskaya [13, p. 339] presented survey of main types of meadows of the Mur--‐ mansk region comparing with meadows of Karelia. In the Murmansk region Ramenskaja described negligible small paludified meadows, small alluvial meadows (due to special hydrology and sand drifts on the banks of running rivers), grass--‐sedges and grass--‐forbs meadows and grasslands with Deschampsia cespitosa , Phleum alpinum , Poa alpina and P. alpigena , Festuca rubra and consider--‐ able part of arctic species. Ramenskaja considered shortgrass early snowbed meadows in tundra zone as a distinctive characteristic of the Kola Peninsula. Meadows of Murmansk region and Kare--‐ lia are similar only in the coastal area.

There are not so much modern researches of non--‐saline meadows and grasslands of the Murmansk region. In the survey of secondary upland meadows on Varlamov Island in the Pasvik Reserve there were presented 7 formations of meadows and grasslands, together with species composition and dynamics [33, 2011]. Formations differ in the soil fertility, the number of vascular plants and bryophytes and the coverage. So, nitrophilous tall grass community with the lowest number of species (8--‐15) and coverage 95--‐100% was described on the soils with high fertility.

Some data about successions and the impact of colonial birds on the coastal non--‐saline meadows (including brackish areas and floodplains) were presented in some modern studies [34, 1969], [35, 1981], [36, 1988]. Coastal meadows on the high seashores include some forest species ( Trientalis europaea and Chamaepericlymenum suecicum) . Some halophytes are presented on high level meadows, which are influenced by salty sea water only during strong storms or high spring tides. I.P. Breslina [37, 1980] studied seashore grasslands and marshes and described asso--‐ ciation of Festuca ovina for sandy dry soils, and association of Phalaroides arundinacea with spe--‐ cies of forbs and grasses in the lower layer on sandy alluvia on some islands and on the mainland coast.

Ornithogenic meadows in the White sea and Barents sea significantly different in species and general appearance from zonal vegetation. Primary

Fig.2. Coastal meadows on the high shores during flow (Kandalaksha Bay)

ornithogenic meadows appear in areas with destroyed plant cover due to activity of colonies of birds (usually on coastal rocks). Ornithogenous vegetation on the White Sea islands [38, 1987], [39, 1965], [40, 1997] divided into three zones: the lower belt of coastal cliffs with a few halophytes; zone of peat accumulation and the formation of ribbon--‐like tufts with dominance of Puccinella pulvina--‐ ta; ribbon--‐like turfs of middle zone with a predomi--‐ nance of clumps of Tripleurospermum hookeri, Cochlearia arctica, Rumex pseudonatronatus,

Sonchus humilis , grasses Festuca rubra , Poa alpigena , Agrostis straminea , Calamagrostis lapponi--‐ ca , etc. Vegetation on cliffs of Eastern Murman hasn’t clear zonal subdivision, there took domi--‐ nance Cochlearia officinalis , Tripleurospermum hookeri and Festuca rubra spp. arenaria .

Secondary ornithogenic vegetation substitutes dry peatlands with dominance of Empetrum hermaphroditum or any other tundra vegetation under influence of nesting seabirds or follows to the primary ornithogenic vegetation. Groups of Cochlearia officinalis together with some other ‘specialists’ grow on the islands of the Eastern Murman and gradually substitute Empetrum her--‐ maphroditum and other tundra dwarf shrubs. Junperus sibirica , Vaccinium ulinginosum , Sedum acre, Festuca rubra, F. ovina, Poa alpine, Leymus arenarius , etc. dominated on the islands of the Kandalaksha bay on these secondary ornithogenic meadows [38, 1987].

• V. B. Golub and D. D. Sokolov [41, 1998] analyzed data on coastal plant communities in Western Europe and the European part of the former USSR using the method of Braun--‐Blanquet. Non--‐saline meadows with prevailing herbaceous perennials on the shores of the White and Bar--‐ ents seas were attributed to 5 classes: Phragmiti--‐Magnocaricetea Klika in Klika et Novak in 1941 (which combines hygrophytic communities on pond banks with fresh, brackish or salt water), Mol--‐ inio--‐Arrhenatheretea R.Tx. 1937 (non--‐saline communities of perennial herbaceous plants with mainly anthropogenic origin), Filipendulo--‐Convolvuletea Gehu et Gehu--‐Frank 1987 (tall--‐grass ni--‐ trophytc and hygroophytic communities with a high proportion of broad--‐leaf herbs), Crithmo--‐ Staticetea Br.--‐Bl. 1947 (hazmophytic vegetation on rocks under influence of salty sea spray and aerosols), Honckenyo--‐Elymetea arenari i R. Tx. 1966 (communities on sand dunes and sand drifts with dominance of Leymus arenarius ).

Coastal communities, including non--‐saline meadow, were characterized in the process of analysis of local floras of the Russian European North ((including the eastern part of the Barents coast of the Kola Peninsula) by S.V. Chinenko [42, 2008]. Chinenko described the position of com--‐ munities in the landscape, the size of phytocoenoses, composition of vascular plants, their occur--‐ rence and abundance, for coastal meadows on the sandy beaches (psammophytic meadows) and stony beaches (petrophytic meadows).

The rare in the Murmansk Region brackish communities were preliminary related to the al--‐ liance Nanocyperion flavescentis Koch 1926 (class Isoёto--‐Nanojuncetea Br.--‐Bl. et Tx. 1943) as a re--‐ sult of ecological--‐floristic classification in the estuary of river Lavna, on the coast of the Barents Sea [43, 2011]. Following types of communities were described: Agrostis stolonifera --‐ Calliergon cordifolium, Callitriche palustris --‐ Limosella aquatica and Zannichellia palustris --‐ Callitriche palustris (rare in the area), Eleocharis uniglumis --‐ Callitriche palustris Communities included Red Data Book species and were considered as value habitats.

Meadows (mainly coastal meadows) as value types of habitats which include species of The Red Data list were considered during the GAP--‐analyses of valuable natural areas in Northwest Rus--‐ sia [11, p. 101]. Value non--‐saline meadows with high biodiversity are rare in the north of the North--‐West of Russia and include regional Red Data Book species such as Armeria scabra , Thymus subarcticus (on dry coastal meadows), Ligularia sibirica (on coastal meadows on the high shores), Valeriana capitata, Tanacetum bipinnatum (meadows of Murmansk Coast), Saxifraga hieracifolia and Castilleja lapponica (shortgrass early snowbed meadows in tundra zone in Lovozero Moun--‐ tains), Alchemilla alpina (meadows of Rybachii Peninsula), Polemonium boreale , Hedysarum arcti--‐ cum (floodplain meadows of Terskiy Coast).

Conclusion

Development of research of meadows and grasslands in Murmansk Province there were practiced different approaches, various definitions of the meadow community as an object of clas--‐ sification, and changing approaches to classification --‐ from individual associations and formations based on dominants trait to the vegetation surveys with a wide geographical and territorial cover--‐ age on the base of ecology and flora. It reflects a paradigm shift that is characteristic for the de--‐ velopment of the science of vegetation in the XX century. Despite the fact that the natural and ge--‐ ographical conditions of the Murmansk region does not favor the formation of large meadows and grasslands, their diversity and specificity are very high, but the level of knowledge about meadows and grasslands on their northern limit is clearly insufficient, and main researches were carried out in the 30th of the last century. Currently, the main directions of investigations in the meadows of the Murmansk region will be study of the flora and typology of meadows and grasslands, their ecology and production process and the cycle of elements. It is necessary to estimate their total area and biodiversity, to analyze geographic variation of meadow and grassland communities at the northern limit of the range and to identify their position in the circumpolar Arctic.