Salt and brackish marshes around the Baltic Sea and adjacent parts of the North Sea: Their vegetation and management

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<ul><li><p>Biological Conservation 51 (1990) 191-209 </p><p>Salt and Brackish Marshes Around the Baltic Sea and Adjacent Parts of the North Sea: Their Vegetation and Management </p><p>Kees S. Dijkema </p><p>Research Institute for Nature Management, PO Box 59, 1790 AB Den Burg, Texel, The Netherlands </p><p>(Received 26 January 1989; revised version received 20 April 1989; accepted 26 April 1989) </p><p>ABSTRACT </p><p>European salt and brackish marshes are in urgent need of protection. This also holds for the many small marsh sites around the Baltic Sea, including the ad/acent parts of the North Sea. Although the individual Baltic shore marshes are small, together they form an important area of salt and brackish marshes in Europe. </p><p>The Baltic' shore vegetation is very complicated because o.1" gradients in salinity, climate, exposure and water levelJtuctuations. There is a wide range of species and communities: from typical brackish communities around the Baltic Sea (with some arctic species in the Bothnian Bay) to more common central Atlantic communities in the western parts of Sweden and Denmark. A well-developed aspect are the natural transitions .from the shore marshes to woodland, bog and heath communities. </p><p>The Baltic shore marshes have a long history of grazing and hay-making, which favour the occurrence of halophytes in brackish marshes. Nowadays, man), Baltic marshes are no longer usedJ~r these purposes, so that reed beds, tall grasses and woodland increase at the cost of halophytes. A general strategy for the management of salt and brackish marshes and proposalsjbr the management of the Baltic' marshes are presented. </p><p>INTRODUCTION </p><p>The salt and brackish marsh sites in northern Europe are often small and of a simple geomorpho log ica l structure. However , the shore vegetat ion as a </p><p>191 </p><p>Biol. Conserv. 0006-3207/90/$03-50 1990 Elsevier Science Publishers Ltd, England. Printed in Great Britain </p></li><li><p>192 Kees S. Dijkema </p><p>substrate geology salt marsh type </p><p>ria bay - - 1 rocky shores loch/fjord head I i = . 1 </p><p>beach head / </p><p>allocht honous barrier-connected </p><p>marine lagoonal sedementary shores foreland 1 </p><p>estuarine i fluvial sedamentary deltaic shores </p><p>emerging flat shore I </p><p>autochthonous emerging skerries I </p><p>peat type </p><p>Fig. I. Coastal salt marsh types in Europe, with the proportional distribution of sites per region, based on site numbers. Each column represents 100% (after Dijkema, 1987b). </p><p>whole is very complicated because of gradients in salinity, climate, exposure and water-level fluctuations. Salt marshes of sedimentary shores hardly occur (except for the southern Baltic Sea and Jutland, Denmark). Most of the shores are rocky, steep and without marshes at all, sometimes with small salt marsh sites in fjords or on skerries (Dijkema, 1987b; Fig. 1). On emerging flat shores near Oulu, Finland, extensive brackish salt marshes are formed. </p><p>In former times salt marshes as a transitional belt between the sea and terrestrial habitats must have been very extensive in Europe, providing a continuous landscape at least along flat coasts of the sedimentary or deltaic type. Large areas of salt marshes have been reclaimed; first, for agriculture, later for other uses connected with human settlement (Dijkema, 1987c). The Council of Europe has published a list and map of more than 600 salt marsh sites in Europe including information on each site (Dijkema et al., 1984). An important conclusion is that to maintain the complete range of halophytic flora and fauna and to ensure the dispersal of halophytic species, all of the remaining salt marsh sites are in urgent need of protection. This also holds for the Baltic salt and brackish marshes. Isolated salt marsh and salt steppe areas (e.g. arctic islands, Gulf of Bothnia, northern Adriatic, inland saline areas of western, central and eastern Europe) have a special conservation value because of their potential for genetic deviation and halophyte dispersal (Beeftink, 1984). Apart from its isolation, the low salinity of the Baltic Sea is of evolutionary interest (Russell &amp; Thomas, 1988). An initial proposal for the sites to be included in a European network of biogenetic reserves has been prepared (Dijkema, 1987a). </p></li><li><p>Baltic brackish marshes 193 </p><p>As part of the Council of Europe study reviews have been made of the extensive literature on the salt marsh vegetation around the Baltic Sea and adjacent parts of the North Sea (e.g. Leivisk~, 1908; Dahl &amp; Hadac, 1941; Dahlbeck, 1945; Mikkelsen, 1949; Gillner, 1960, 1965; Tyler, 1969a, 1969b; Siira, 1970; Gravesen, 1972; Ericson &amp; Wallentinus, 1979; Vartiainen, 1980). In 1982 the marsh sites along the Baltic coasts were visited and data on area, vegetation, land use and protection status, collected by means of a questionnaire (Dijkema et al., 1984). The purpose of this compilation is to show the uniqueness of the Baltic shore marshes and the need to preserve them. </p><p>ECOLOGICAL CONDITIONS </p><p>The salinity of submerging water is an important factor in the distribution of halophytic species. Western and Southern Norway are bordered by the Norwegian Sea and the North Sea, which have normal saline conditions (Table 1). The Baltic Sea is a vast brackish sea, connected to the North Sea by a narrow transition area between Denmark and southwestern Sweden. The transition area has strong gradients and fluctuations in salinity. The Baltic Sea can be divided into several basins with different ecological conditions (Table 1): the Baltic Sea proper and the Gulfs of Riga, Finland and Bothnia, the last including the Bothnian Sea and the Bothnian Bay. Compared to the </p><p>TABLE 1 Salinities in the Basins of the Baltic and the Adjacent North Sea [after Tyler, 1969a </p><p>and Ericson &amp; Wallentinus, 1979). </p><p>Salinity o]open Venice classification coastal water </p><p>(%0 S) </p><p>Open Sea Norwegian Sea 33 30 euhalinicum North Sea 34 32 euhalinicum Skagerak 33-27 eu- and polyhalinicum </p><p>Transition Kattegat 30-18 polyhalinicum area Belt Sea 18-10 ~-mesohalinicum </p><p>Oresund 10-8 /%mesohalinicum </p><p>Baltic Sea Baltic Sea proper 10-5 /~-mesohalinicum Gulf of Riga Gulf of Finland 5~3 oligohalinicurn Gulf of Bothnia </p><p>Bothnian Sea 5-3 ~-oligohalinicum Bothnian Bay 3q3-3 /#oligohalinicum </p></li><li><p>194 Kees S. Dijkema </p><p>transition area the open water of the Baltic Sea has a stable salinity, but at the heads of the Bothnian Bay and the Gulf of Finland, and in bays, fjords and near river mouths, large fluctuations occur. </p><p>Tidal fluctuations are small, from about 30 cm in the Skagerak, decreasing to 15 cm in the Kattegat and 10cm in the Belt Sea. In the Baltic Sea and the Gulf of Bothnia tides scarcely exist (Gillner, 1965; Ericson &amp; Wallentinus, 1979). In the Baltic Sea and the Gulfs, water-level fluctuations are mainly determined by the season--by wind direction and force, air pressure and discharge from rivers--and are lowest in late winter and spring and highest in autumn and early winter. The extremes in fluctuation for these periods are 70 and 150 cm, respectively, and reach up to 180 and 300 cm, respectively, at the heads of the Bothnian Bay and the Gulf of Finland, and even 370 cm at The Polish-German border (Gillner, 1965; Ericson &amp; Wallentinus, 1979). The vertical extent of the marshes increases accordingly from 50-60 cm on the Swedish west coast to 50-130cm around Oulu off the Bothnian Bay (Gillner, 1960; Siira, 1970). </p><p>The average water levels between spring and summer vary about 15 cm for the transition area (Gillner, 1960) and 23 cm for the central Baltic Sea (Tyler, 1969a). A low water level in spring is of particular importance for the vegetation of the low marsh zones. For example, seedlings of Scirpus tabernaemontani, Eleocharis palustris and Eleocharis uniglumis make a rapid appearance during years with comparatively low spring water levels (Ericson &amp; Wallentinus, 1979). Year-to-year changes are considerable and show a rather regular shift between low- and high-water years (Ericson, 1981). </p><p>An important ecological condition is the uplift of northern Europe, which increases from zero in southern Sweden, southern Norway and northern Denmark to more than 0"9 cm year-1 in the Bothnian Bay (Devoy, 1987). Land-uplift causes the vegetation zones to move seaward in a long-term process. Annual fluctuations in water level are more important in the short term (&lt; 10 years) (Schwanck, 1974; Ericson &amp; Wallentinus, 1979; Ericson, 1980; Vartiainen, 1980; Cramer &amp; Hytteborn, 1987). The expected worldwide rise in sea level (20-165 cm for the next century; Robin, 1986) might reach higher values than the present rate of land-uplift. </p><p>From the southern Baltic Sea to the northern Bothnian Bay the length of the growth period is reduced by about a third, and the duration and thickness of the ice cover increase strongly. In the Bothnian Bay ice may have severe damaging effects on the vegetation and thus prevent the occurrence of reeds (Siira, 1970; Ericson &amp; Wallentinus, 1979). </p><p>From the open sea into the archipelagos there is a decreasing gradient in salinity, wave action and sunshine, whereas sedimentation, duration and thickness of ice, and precipitation increase (Ericson &amp; Wallentinus, 1979). </p></li><li><p>Baltic brackish marshes 195 </p><p>VEGETATION AND DISTRIBUTION </p><p>In Tables 2, 3 and 4 the vegetation communities of the Baltic shore marshes and the salt marshes of the adjacent parts of the North Sea are compared by listing the common plant species for some important regions arranged according to the gradients in climate and salinity. The communities have been arranged in zones related to the mean water level, comparable to the zones used for North Sea salt marshes (Dijkema et al., 1984; cf. Beeftink, 1977a). Due to the small vertical range in the zonation of the Baltic shore marshes the zones are close together and often difficult to distinguish from each other. </p><p>Communities of emerging flats </p><p>In this pioneer zone below mean water level a striking shift from halophytic to freshwater communities occurs. Due to decreasing salinity the Salicornia dolichostachya community on Danish, German and western Swedish sites is replaced by a brackish reed belt (Scirpus maritima and S. tabernaemontani) in the inner transition area and the Baltic Sea and Gulfs. Aster tripolium is an important species in this zone. In the southern Baltic Sea area a Juncus maritimus community may occur (Voderberg, 1955; Fukarek, 1961; Gillner, 1965; Piotrowska, 1974). Towards the least saline Gulf of Bothnia the non- halophytic Phragmites australis and Eleocharis palustris communities increase. Scirpus tabernaemontani still occurs in brackish water; in freshwater an Equisetumfluviatile community appears. Northwards the reed belts may disappear on exposed sites due to ice erosion. Around the Baltic Sea the halophytic Eleocharis parvula community may occupy bare emerging places, in the Gulf of Bothnia and the Gulf of Finland they are replaced by a non-halophytic Eleocharis acicularis community. </p><p>Communities of lower marshes </p><p>Above mean water level the lower marsh forms a distinct zone, mostly belonging to the Puccinellietum maritimae association in marine conditions and to the Eleocharetum uniglumis association in brackish conditions. However, in areas without grazing, reed beds may extend upwards (e.g. Kauppi, 1967). On the shores of Norway, southwestern Sweden, Denmark and the German Baltic Sea Puccinellia maritima dominates the lower marsh (Puccinellion maritimae alliance). It is accompanied by halophytic species like Aster tripolium, Plantago maritima, Triglochin maritima, Spergularia media, Salicornia europaea and sometimes Limonium humile. With </p></li><li><p>TA</p><p>BLE</p><p> 2 </p><p>Co</p><p>mm</p><p>on</p><p> Plan</p><p>t Sp</p><p>ecie</p><p>s of (</p><p>Ha</p><p>lop</p><p>hyt</p><p>ic) S</p><p>ho</p><p>re C</p><p>om</p><p>mu</p><p>nit</p><p>iesf</p><p>l So</p><p>uth</p><p>ern</p><p> No</p><p>rwa</p><p>y a</p><p>nd</p><p> So</p><p>uth</p><p>we</p><p>st an</p><p>d S</p><p>ou</p><p>the</p><p>ast</p><p> Sw</p><p>ed</p><p>en</p><p>, Ba</p><p>sed</p><p> on</p><p> Lit</p><p>erat</p><p>ure</p><p> and</p><p> Ow</p><p>n </p><p>Ob</p><p>serv</p><p>ati</p><p>on</p><p>s </p><p>Sou</p><p>ther</p><p>n No</p><p>rwa</p><p>y S</p><p>ou</p><p>thw</p><p>est</p><p> Sw</p><p>eden</p><p> Sou</p><p>thea</p><p>st Sw</p><p>eden</p><p> 6%</p><p>00 S (</p><p>Tyle</p><p>r, 1</p><p>96</p><p>9b; W</p><p>alle</p><p>nti</p><p>nus,</p><p> 19</p><p>67</p><p>, 197</p><p>3) </p><p>Tele</p><p>mar</p><p>k 30%</p><p>0 S </p><p>Ost</p><p>oy (</p><p>Os]</p><p>ofjo</p><p>rd) </p><p>Ska</p><p>ge</p><p>rak +</p><p> Kat</p><p>tegat</p><p> O</p><p>resu</p><p>nd 10</p><p>-8%</p><p>o S </p><p>( Vev</p><p>le, 19</p><p>82</p><p>, 198</p><p>5; </p><p>18%</p><p>o S (</p><p>Dah</p><p>l &amp; H</p><p>adac</p><p>, 3</p><p>~2</p><p>0%</p><p>o S (</p><p>Gill</p><p>ner</p><p> (D</p><p>ahlb</p><p>eck,</p><p> 1945</p><p>; O</p><p>pen</p><p> shor</p><p>e Shel</p><p>tere</p><p>d shor</p><p>e H</p><p>ofst</p><p>en &amp; V</p><p>evl</p><p>e, 19</p><p>82) </p><p>1941</p><p>) 19</p><p>60, 1</p><p>965)</p><p> G</p><p>illner</p><p>, 196</p><p>5) </p><p>Emer</p><p>gin</p><p>g fla</p><p>ts </p><p>(Ele</p><p>ochar</p><p>is par</p><p>vula</p><p>) (E</p><p>leoc</p><p>har</p><p>is par</p><p>vula</p><p>) (E</p><p>leoc</p><p>har</p><p>is par</p><p>vula</p><p>) Sal</p><p>icor</p><p>nie</p><p>tum</p><p> stri</p><p>ctae</p><p> S</p><p>uli</p><p>corn</p><p>ia </p><p>dol</p><p>ichos</p><p>tach</p><p>ya (S</p><p>alic</p><p>ornia</p><p> dol</p><p>iehos</p><p>taeh</p><p>ya) Sal</p><p>icor</p><p>nia</p><p> dol</p><p>ichos</p><p>tach</p><p>ya </p><p>Ha</p><p>lo-S</p><p>cirp</p><p>etu</p><p>m </p><p>ma</p><p>riti</p><p>mi </p><p>(Sci</p><p>rpus m</p><p>arit</p><p>imus)</p><p> Sei</p><p>rpus m</p><p>arit</p><p>imus </p><p>(Sci</p><p>rpus m</p><p>arit</p><p>imus)</p><p> Sei</p><p>rpus m</p><p>arit</p><p>imus </p><p>Sci</p><p>rpus m</p><p>arit</p><p>imus </p><p>(Sci</p><p>rpus m</p><p>arit</p><p>imus)</p><p> (S</p><p>cirp</p><p>us ta</p><p>ber</p><p>nae</p><p>mon</p><p>tani)</p><p> (S</p><p>cirp</p><p>us t</p><p>aber</p><p>nae</p><p>mon</p><p>tani)</p><p> (Sei</p><p>rpus t</p><p>aber</p><p>nae</p><p>mon</p><p>tani)</p><p> Sci</p><p>rpus t</p><p>aber</p><p>nae</p><p>mon</p><p>tani </p><p>Phra</p><p>gm</p><p>itio</p><p>n </p><p>(Phra</p><p>gm</p><p>ites</p><p> au</p><p>stra</p><p>lis*)</p><p> Ph</p><p>ragm</p><p>ites</p><p> aust</p><p>ralis</p><p>* (P</p><p>hra</p><p>gm</p><p>ites</p><p> aust</p><p>ralis</p><p>*) </p><p>Phra</p><p>gm</p><p>ites</p><p> aust</p><p>ralis</p><p>* Ph</p><p>ragm</p><p>ites</p><p> aust</p><p>ralis</p><p>* Lo</p><p>wer</p><p> mar</p><p>sh </p><p>(Car</p><p>ex pa</p><p>leac</p><p>ea) </p><p>Car</p><p>e,,: p</p><p>alea</p><p>cea </p><p>(Car</p><p>ex pal</p><p>eace</p><p>a) </p><p>Pucc</p><p>ine</p><p>llie</p><p>tum</p><p> ma</p><p>riti</p><p>ma</p><p>e P</p><p>uec</p><p>inel</p><p>lia mar</p><p>itim</p><p>a- </p><p>Pucc</p><p>inel</p><p>lia mar</p><p>itim</p><p>a- </p><p>Puec</p><p>inel</p><p>lia ma</p><p>riti</p><p>ma</p><p> Pu</p><p>ccin</p><p>ellia</p><p> mar</p><p>itim</p><p>a- </p><p>Plan</p><p>t. m</p><p>ar.-</p><p>Ast</p><p>er trip</p><p>. 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