Koroleva N., Polar alpine botanical garden, Kirovsk, Russia 184236.
Coastal vegetation is a subject of continuous interest to geobotanists. Comprehensive studies have have described the composition of plant communities in mid and northern Europe (Nordhagen 1940, 1955; Tüxen 1966, 1967; Passarge & Passarge 1973; Olsson 1974; Thannheiser 1974, 1982, 1987, 1995) and portions of the circumpolar Arctic (Thannheiser 1991; Kojima 1991; Daniels & Molenaar 1993). However there are few studies which have addressed similar plant communities in the Kola Peninsula. The purpose of this study is to describe the coastal vegetation of the south-eastern part of the Kola Peninsula and to present data on the responses of coastal vegetation to some aspects of human influence. This study compares the processes and patterns of vegetation recovery with those reported from seashore regions of northern Europe.
The study area.
The study area is located on the White Sea shore of the Kola Peninsula and lies between 66° 17' and 66° 30' N latitude and 36° 55' and 40° 47' E longitude. This part of the shore of the White Sea , i.e. to the east of the mouth of the Umba River to the mouth of the Ponoi River is known as Tersky bereg. The almost straight coastline and broad sea terrains were formed as a result of sea abrasion are characteristic features of landscape of Tersky Bereg. All sites investigated are underlain by crystalline Archeic and Proterozoic rocks, in some places by Paleozoic red sandstones and are covered by drift material with fluvioglacial and marine deposits. Alluvial deposits are of little importance, though in the mouths of large rivers these deposits take part (Geologicheskoe... 1958).
The substrates consist primarily of Pleistocene sands underlain by marine deposits. Unsorted cobble-stones can occasionally be found mixed in the sand, as well as sandstone flags, which form extensive flat fields in the dune area. In heathlands and grasslands the humus horizon is well-developed, but normally thin, and the podzol layer is poorly differentiated. A slight peat layer is present in wet depressions, as well as gleyish lenses in the transitional zone to the bedrocks. Permafrost doesn’t occur in study area (Kazakov 1995).
Aeolian sands occupy only a small part of the shore, being scattered and mainly limited near river deltas. But in some areas (in the Varzuga river mouth) extensive sand fields and dune system occur.
The climate of the area studied is marine, with moderately cold, snowy winters, short, cold summers and long spring and autumn periods. The year-point meteorological data are presented from meteorological station Kuzomen' (66°17' N, 33°55' E) (Anon. 1965, 1968). The annual average air temperature is -0.2°C, for July 12.6°C, and for January-February -11.7°C. The averaged growing season lasts 115 days. Snow cover lies about 200 days and its distribution is strongly affected by wind and topography. The annual average precipitation is 565 mm, that for August is maximal (62-75 mm), for July minimal (24-29 mm). Annual average wind velocity is 5-6 m/sec, that is maximal for November and December (28-30 m/sec). The prevailing wind direction in January is SW, and in July - SE , E and N. (Jakovlev 1961). It has been determined the velocity of winds with aeolic capacity is about 4-7 m/sec; velocity 8-14 m/sec erodes the ground intensively and wind of velocity more than 15 m/sec causes a sand blizzard in dune areas (Kazakov 1995).
Human influence
Humans have occupied Tersky bereg for a very long time: there is archeological evidence from inhabited sites in the Lower Varzuga River dated from the X-XII centuries. The first settlements were established by Novgorod's traveling people, who sought new territories to join to the State Velikiy (Great) Novgorod. They settled near the mouths of rivers and practiced mainly fishing and hunting, especially sea mammals. The impact of such a sparse and uneven population on the terrestrial ecosystem was not very great. With settlement enlargement in the last century reindeer herding and cattle farming was established, as well as some agriculture. As referred to the statistical records of 1876, the largest village on the Tersky bereg was Kuzomen’, situated near the mouth of the Varzuga river. It had a population of 780 people, and 1700 animals (including cows, sheep and reindeer) (Ushakov 1972). There were few open pastures, but cattle grazed freely on the tundra and forest around the villages. In winter the animals were kept in barns, but reindeer were employed for transport and sometimes spent nights grazing around trees, to which they were tethered. Reindeers cratered through the snow and ate all of the plant cover, including small trees, in a radius of about 15 m. These patches of bare soil joined and marginal closed woodlands gradually transformed to sparse, open trees with destroyed undergrowth and ground layers. This regime of animal husbandry brought a significant change in coastal heathlands and marginal forests. In addition, by the end of last century marginal forests were simultantially burnt (Kazakov 1997). Vegetation was destroyed and wind blown sand had accumulated in the Varzuga, where it had threaten the spawning grounds of the best and largest salmon population in the White Sea region. Woody fences were used as a protection of the river against shifting sand.in the end of last century. Grasses were planted near the Kuzomen' in 1926, but destroyed by the wind. Erosion peaked in 1950’s, as a result of increasingly intensive grazing and agriculture on the sandy soils. The Kuzomen' became desolate and, as a result, most of people moved to other villages on the Tersky bereg. Though human impact on the vegetation ceased, sand eroded continuously. In the 1970’s, experimental planting of juvenile pine (Pinus friesiana) and dune grasses (including.Leymus arenarius) had begun. Cultivation of these species occured in a limited area and is still in an experimental stage.
Material and methods
65 relevés of both disturbed and undisturbed vegetation were made. during the July-August, 1990 and included plots described along a transect, crossing the dune area, plots in coastal heathlands, abandoned dwelling sites and cattle-stalls. Plot sizes were from 2x2 min heathlands and grasslands and dunes but 1 m x 15 m in the vegetation alongside the beach and 0.5x0.5 m in the primary disturbed sites. In each plot vascular and non-vascular vegetation was sampled for composition and cover-abundance using a Braun-Blanquet scale.
The vegetation data were classified following the Braun-Blanquet approach with modifications (Daniels 1982) resulting in phytosociological tables with groups of similar relevés interpreted as communities characterized by diagnostic species, which were either characteristic taxa (Ch) or differential taxa (D).
Nomenclature for vascular plants follows Cherepanov (1995), for mosses Ignatov & Afonina (1992), for liverworts Konstantinova et al (1992) and for lichens Santesson (1993).
Results
Dune area
An extensive desert-like landscape formed as a result of human impact and destruction of the plant cover on the sandy marine deposits,. Individuale blow-outs (from 0.5 m2 to 1000 m2) occur everywhere in the forest and heathlands on the shore. They comprise the first stage of wind erosion on sandy terrain. Even when the bottom of the eroded hollow reaches the ground water level, walls are still exposed to strong deflation. Patches of eroded substrata can enlarge and join together, forming large hollows which cover hundreds of square metres. Extensive sandstone fields that were formed as a result of sand transfer are completely unvegetated, with exception of isolated stems of Agrostis straminea. A plant cover dominated by liverworts and dwarf shrubs establishes wherever the water table is near the eroded sand surface.
Embryonic dunes are formed along the perimeter of the hollow as a result of accumulation of aeolian sand. Strong, prevailing winds promote further erosion, primary dunes reach 3 meters in height, and migrating dune velocity reaches 6 m per year. Buried humus-layer as well as dwarf shrubs remnants can be exposed, when mobile sand is transported from the windward side to the leeward side of the dune. Sand accumulation on the leeward side of dune quickly buries the dwarf shrubs at the margin of the Pine forest and in the adjacent heathlands.
Secondary and tertiary dunes are lower, about 1-1.5 m.They are often connected to the chains and are more stable because of the settlement of dune grasses (Leymus arenarius, Festuca arenaria) and some herbs (Solidago lapponica). Depressions in the dune area are colonized mainly by Agrostis straminea.
Plant communities
Biogeographically the area comprises a transitional zone between tundra and boreal forest and southern limit of hypoarctic tundra (Alexandrova 1971, Gribova 1980). Coastal plant communities observed include beaches, dune areas, heathlands and grasslands, old dwellings and stall sites. They occur on upper geolittoral, epilittoral and marine terrain. They are relatively dry with exception of those which occupy depressions with high water table. They are not saline, since the climatic and geomorphological conditions do not permit the development of marshes. Coastal vegetation types often occur in a distinct, though simple, zonation pattern. In dune areas, the differentiation of plant communities can be ascribed to the character of dunes.
Classification.
Class Honckenyo-Elymetea arenariae Tx.66
Alliance Honckenyo-Elymetalia arenariae Tx.66
Order Honckenyo-Elymion arenariae (Fernandez-Galiano 54) Tx.66
Ass. Honckenyo diffusae-Elymetum arenarii (Regel 28) Tx. 66 (Tab.1)
DS: Honckenya oblongifolia (Ch), Lathyrus aleutica (Ch), Mertensia maritima (Ch).
Isolated stands of Honckenya oblongifolia, Mertensia maritima with Leymus arenarius occupy the sandy upper geolittoral zone. These habitats are sheltered by wood and logs cast ashore, and enriched by organic matter (tidal deposits).
Table 1
Synoptic table of plant communities of the dune area and epi- and geolittoral zone. Roman numbers indicate presence degree, arabic numbers indicate range of cover values.
1: Agrostis straminea - community,
2: ass. Elymo-Festucetum arenariae,
3: ass. Honckenyo diffusae-Elymetum arenarii
| com./association no. |
1 |
2 |
3 |
| number of releves |
10 |
10 |
9 |
|
DS of com. no. 1 |
|||
| Agrostis straminea |
III+-1 |
- |
- |
|
DS of ass. no. 2 |
|||
| Rumex graminifolius |
III+-1 |
IV+-1 |
I+ |
| Festuca arenaria |
- |
IV+-1 |
I |
|
DS of ass. no. 3 |
|||
| Honckenya peploides |
- |
- |
IV+ |
| Mertensia maritima |
- |
- |
II+ |
| Lathyrus aleuticus |
- |
- |
II+-1 |
|
other taxa |
|||
| Leymus arenarius |
I1 |
V+-3 |
V+-1 |
| Polytrichum piliferum |
I1 |
II1-3 |
I+ |
| Solidago lapponica |
- |
II+-1 |
II+ |
| Luzula spicata |
I1 |
I+ |
- |
| Empetrum hermaphroditum |
- |
I+-1 |
- |
| Polytrichum juniperinum |
- |
II+-1 |
- |
Ass. Elymo-Festucetum arenariae (Regel) Nordh.55( Tab. 1)
DS: Festuca arenaria and Rumex graminifolius are differential taxa against previously observed associations. Open and closed poor species communities of this association are common for the all dune area. Leymus arenarius is the clear dominant of the homogenous single- or two-layered vegetation. Sparse grasses and dwarf shrubs can to enlarge the coverage in secondary and tertiary dunes. Polytrichum piliferum and P. juniperinum form a weak ground layer. On the abandoned dwellings these association include synusiae of Ceratodon purpureus and Isopaches bicrenatus, which are common on disturbed sites.
Agrostis s traminea- com (Tab. 1)
DS:Agrostis straminea is a differential taxon against both previously observed syntaxa. Stands are common in wet depressions inside the dune area.
Class Cetrario-Loiseleurietea Suzuki-Tokyo & Umezu apud Suzuki-Tokyo 64
Order Cetrario-Loiseleurietalia Suzuki-Tokyo & Umezu apud Suzuki-Tokyo 64
Alliance Loiseleurio-Diapension Braun-Blanquet, Sissingh &Vlieger 39 ex Daniels 82. (Tab. 2)
Ass. Arctostaphylo alpini- Empetretum hermaphroditi (Zinserling 1935) Koroleva 1994 (Tab. 2 )
DS: Arctous alpina, Ledum palustre, Pleurozium schreberi, Ptilidium ciliare, Dicranum fuscescens, that are differential against other syntaxa. Communities of this association form a monotonous dense carpet, sometimes with a shrub layer of Juniperus sibirica. Empetrum hermaphroditum and Arctous alpina are dominants of the field layer; the ground layer is scattered, with mosses Ptilidium ciliare, Pleurozium schreberi and Dicranum fuscescens alternating with patches of lichens Cladonia spp., Cladina spp., Cetraria spp.
Sphaerophorus globosus-Empetrum hermaphroditum - community. (Tab. 2)
DS: Ochrolechia frigida (D). Stands occur in blow-outs on the marine terrain and consist of pioneer synusiae of chionophobous lichens, which occupy unstable substrata together with pioneer mosses of Polytrichum spp. and Empetrum hermaphroditum, which thrive onadjacent undisturbed sites. Large hollows are occupied by Juncus trifidus, Festuca spp.-dominated communities with synusiae of the same chionophobous lichens.
Table 2
1: Sphaerophorus globosus-Empetrum hermaphroditum - comm.
2: ass. Cetrario nivalis-Festucetum ovinae
3: ass. Arctostaphylo alpini- Empetretum hermaphroditi
|
subassociation/community no. |
1 |
2 |
3 |
|
|
number of releves |
9 |
10 |
10 |
|
|
DS of All. Loiseleurio-Diapension |
||||
| Cetraria nivalis |
V+-4 |
V+-2 |
II+-2 |
|
| Alectoria ochroleuca |
II+-2 |
II+-2 |
- |
|
| Polytrichum piliferum |
III+-4 |
III+-4 |
- |
|
| Thamnolia vermicularis |
II+ |
I+ |
I+-1 |
|
| Alectoria nigricans |
III+-2 |
II+-1 |
II+-2 |
|
| Sphaerophorus globosus |
III+-3 |
II+-1 |
II+-2 |
|
| Bryocaulon divergens |
IV+-5 |
III+-3 |
I+ |
|
|
DS of comm. no. 1 |
||||
| Ochrolechia frigida |
III+ |
I + |
I+ |
|
|
DS of subass.no. 2 |
||||
| Thymus subarcticus |
I+ |
IV+-2 |
- |
|
| Oxytropis sordida |
- |
III+-4 |
I+ |
|
| Festuca ovina |
I+ |
V2-5 |
IV+-1 |
|
|
DS of subass. no. 3 |
||||
| Arctous alpina |
I1 |
II4 |
IV+-4 |
|
| Ptilidium ciliare |
- |
I+ |
IV+-2 |
|
| Pleurozium schreberi |
- |
- |
III2-5 |
|
| Dicranum fuscescens |
- |
- |
II+-1 |
|
| Ledum palustre |
- |
- |
II+-2 |
|
|
other taxa |
||||
| Empetrum hermaphroditum |
III+-2 |
V+-5 |
V3-5 |
|
| Cetraria islandica |
III+-2 |
V+-5 |
IV+-3 |
|
| C. cucullata |
I+ |
III+-4 |
III+-1 |
|
| Polytrichum juniperinum |
I+ |
I+-1 |
III+-3 |
|
| Vaccinium vitis-idaea |
I+ |
II+-3 |
III+-2 |
|
| V. uliginosum |
II+ |
I+ |
IV+-4 |
|
| Cladonia uncialis |
III+ |
II+-2 |
II+-2 |
|
| C. mitis |
IV+ |
II+-3 |
IV+-2 |
|
| C. rangiferina |
II+ |
I+ |
III+-2 |
|
| C. amaurocraea |
III+ |
I+ |
II+ |
|
| Stereocaulon spp. |
II+ |
II+ |
I+ |
|
| Equisetum arvense |
I+ |
I+ |
II+ |
|
| Cladonia coccifera |
II+ |
I+ |
I+ |
|
| C. pleurota |
I+ |
II+ |
I+ |
|
| C. macroceras |
I+ |
I+ |
I+ |
|
| C. bellidiflora |
I+ |
I+ |
I+ |
|
| C. gracilis |
I+ |
I+ |
I1-2 |
|
| C. cornuta |
II+ |
II+ |
- |
|
| Cephaloziella grimsulana |
I+ |
I1 |
I+ |
|
Ass. Cetrario nivalis-Festucetum ovinae (Nordh. 1943) Dierssen 1992 (Tab. 2)
DS: Thymus subarcticus (D), Oxytropis sordida (D). Festuca ovina is the dominant. The dominant subassociation comprises closed double-layered plant communities with dominance of Festuca ovina and Empetrum hermaphroditum. Lichens Cladonia spp., Cetraria spp. are common in the ground layer, synusiae of Ceratodon purpureus and Leptobryum pyriforme are typical for these sites. Stands occur on the pastures and abandoned cattle-stall.
Class - ?
Order - ?
]
Alliance - ?
Ass. Gymnocoleo inflatae- Juncetum filiformis ass. nova (Tab.3) Typical réleve - no. 2p.
DS: Juncus filiformis, Gymnocolea inflata. Closed or dense vegetation with variable floristic composition and undetermined sociological degree, that represent a homogenous cover of pioneer liverworts (Gymnocolea inflata, Scapania sp.) with scattered layer of rushes (Juncus filiformis, J. balticus), dwarf shrubs, occurring mainly on low flat hillochs resulting from cryogenic processes - Empetrum hermaphroditum, Vaccinium uliginosum, Calluna vulgaris, Andromeda polifolia, sedges and herbs (Carex maritima, Armeria labradorica, Parnassia obtusifolia). Stands occupy large depressions on the shore with the bottoms being at the level of the water, resulting from the sand removal. Though these depressions represent relatively stable substrata, they don't offer suitable sites for many vascular plants, as stands are inundated by rainwater in the autumn and covered by ice in the winter. Further inland these plant communities are preceded by the vegetation of the dune area, and the ridge-hollow mire complex.
Table 3
Ass. Gymnocoleo inflatae-Juncetum filiformis ass. nova.
Synoptic value for the community (see Table 1) given in col. 11
| Reference no. |
1 |
2 |
3 |
4 |
5 |
6 |
7 |
8 |
9 |
10 |
11 |
| Releve no. |
197 |
205 |
2p |
7p |
9p |
10p |
67' |
66' |
2p |
206 |
|
| Total cover (%) |
95 |
100 |
100 |
75 |
25 |
15 |
10 |
50 |
100 |
100 |
|
| Cover grasses and dwarf shrubs (%) |
75 |
75 |
25 |
25 |
20 |
10 |
10 |
45 |
10 |
10 |
|
| Cover mosses (%) |
15 |
100 |
100 |
55 |
5 |
5 |
1 |
5 |
100 |
100 |
|
| Cover lichens (%) |
- |
1 |
<1 |
<1 |
<1 |
- |
<1 |
- |
<1 |
5 |
|
|
DS of association |
|||||||||||
| Juncus filiformis |
3 |
3 |
1 |
3 |
3 |
1 |
+ |
- |
- |
- |
IV+-3 |
| Gymnocolea inflata |
- |
3 |
3 |
- |
+ |
+ |
- |
- |
4 |
5 |
III+-5 |
|
other taxa |
|||||||||||
| Empetrum hermaphroditum |
1 |
- |
1 |
+ |
- |
+ |
2 |
2 |
1 |
1 |
IV+-2 |
| Polytrichum commune |
1 |
- |
- |
1 |
- |
+ |
+ |
+ |
- |
2 |
IV+-2 |
| Vaccinium uliginosum |
1 |
+ |
1 |
- |
+ |
- |
- |
+ |
1 |
- |
III+-2 |
| Cephalozia bicuspidata |
1 |
2 |
- |
3 |
- |
- |
- |
- |
+ |
3 |
III1-3 |
| Scapania sp. |
1 |
1 |
- |
- |
1 |
- |
- |
- |
+ |
2 |
III1-2 |
| Andromeda polifolia |
- |
1 |
+ |
- |
- |
+ |
- |
+ |
- |
- |
III+-1 |
| Calluna vulgaris |
1 |
- |
1 |
2 |
- |
- |
- |
2 |
- |
- |
II1-2 |
| Carex bigelowii |
- |
1 |
3 |
3 |
- |
- |
- |
- |
- |
- |
II1-3 |
| Vaccinium vitis-idaea |
+ |
- |
+ |
+ |
- |
- |
- |
- |
- |
- |
II+ |
| Juncus balticus |
- |
- |
- |
- |
+ |
1 |
+ |
- |
- |
- |
II+-1 |
| Carex maritima |
- |
- |
+ |
- |
+ |
- |
- |
- |
+ |
- |
II+ |
| Stereocaulon spp. |
- |
- |
+ |
+ |
+ |
- |
- |
- |
+ |
+ |
II+ |
| Polytrichum jensenii |
1 |
1 |
1 |
- |
- |
- |
- |
- |
- |
- |
II1 |
| Armeria labradorica |
- |
+ |
- |
- |
- |
- |
- |
- |
+ |
+ |
I+ |
Additional taxa (occurring with + unless indicated otherwise): Solidago lapponica (2)205; Parnassia palustris 2p,9p;; Pinguicula alpina 9p; Rumex graminifolius 10p; Plantago maritima 2p, 3p; P. schrenkii 3p; Oxycoccus microcarpus 66'; Luzula spicata 10p; Carex concolor 10p; Pohlia nutans 197, 7p; Polytrichum juniperinum 10p, 66'; Aulacomnium palustre 66'; Orthocaulis kunzeanus 2p; Pleurocladula albescens 205,206; Scapania hyperborea 2p; Cladonia coccifera 2p,7p; C. rangiferina 2p; Cetraria cucullata 2p; C. nivalis 7p; C. islandica (1) 2p.
Discussion
Thannheiser (1991) says, that it is difficult to put a distinct boundary between arctic and boreal zone in coastal areas. This is the case at the Tersky bereg of the Kola Peninsula. Though the majority of the area studied lies in the boreal zone, the coastal vegetation shows striking similarities to arctic and subarctic seashore plant communities. The most widespread association alongside the beach is Honckenyo diffusae-Elymetum arenariae, that is reported to be typical everywhere in Circumpolar Regions. The prevailing association on coastal terrain, Arctostaphylo alpini-Empetretum hermaphroditi, shows similarity with the seashore vegetation of Northern and Middle Europe, which belong to alliances Empetrion nigrae Böcher 1943, Phyllodoco-Vaccinion myrtilli Nordhagen 1936 и Oxycocco-Empetrion hermaphroditi Nordh.1936. (Smidt 1967, Barendregt 1982). In plant communities on the White Sea shore arctic and hypoarctic species take more essential part (Koroleva, 1994).
The present plant cover of the dune area appears to characterize the succession course and rate in the Low Arctic, with a relatively small number of syntaxa described. Ass. Elymo-Festucetum arenariae is prevailing type of dune vegetation. General succession course proposed for dune area of coastal Europe (boreoatlantic zone) (Tûxen, 1967) is weakly expressed. Even after 30 years after the cessation of activity unassisted recovery and stabilization of the dune area have failed to occur. In spite of the fact that secondary and tertiary dunes have been reported as a more stable kind of substrata, they are destroyed by the wind erosion, and cyclic changes take place.
In contrast, wet sites resulting from sand removal recovered more readily. The most successful invaders were liverworts Gymnocolea inflata, Cephalozia bicuspidata, Scapania obcordata, that occurred in small microsites on the ridge-hollow mire complex, preceding the seashore communities inland. Association Gymnocoleo inflatae-Juncetum filiformis ass. nova.represent first stage of succession on new substrata. Presence of Juncus balticus and Carex maritima shows an affinity of this syntaxa to halotolerant association Carici maritimae-Juncetum baltici Vanden Bergen 1969 (Dierssen, 1992). Groups of dwarf shrubs, mainly concentrated on cryogenic hillocks, reveals a tendency to formation of fragments of oligotrophic and distrophic bogs of alliance Oxycocco-Empetrion hermaphroditi Nordh.1936, which have a northern limit on the White Sea shore. Successional course on new substrata is likely to lead to komplex of bogs on hillocks and halophytic rushes-dominated communities in depressions.
Conclusion
Plant communities on the Tersky Bereg area, which belong to boreal zone, have similar features with subarctic seashore vegetation, where Honckenyo diffusae-Elymetum arenariae is typical association on the beach.
Prevailing association on the Sea terrain is Arctostaphylo alpini-Empetretum hermaphroditi, that shows similarity with mid and northern Europe coastal vegetation.
Successional course in the dune area is presented by small number of syntaxa: association Elymo-Festucetum arenariae prevails on embrionic, secondary and tertiary dunes.
Vegetation arising after the destruction of native plant cover (and native substrata) is presented by association Gymnocoleo inflatae-Juncetum filiformis and is completely different from native plant cover. Further transformation do not appear to lead to the original plant cover - dwarf shrub dominated sea shore plant communities. These communities, as well as marginal forests need special protection in this coastal area.
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