Author: Schneider-Binder, Erika
Date published: January 1, 2010
Journal code: TRSC
Wet habitat types of the alpine level in the Alpine mountains system - the Pyrenees, Alps and Carpathians - are well represented by springs and their surrounding area, alpine rivulets and streams with their alluvial deposits, protosoils and soils composed of different grain sizes, smaller or larger seepage areas, snowmelt depressions, along the edges of glaciers, glacial torrents as well as around the glacial lakes and as bogs and fens.
Given that the Alps have larger and higher mountain peaks as compared to the Carpathians, this habitat type is rather representative of the Alps than of the Carpathians. But despite of all differences the alpine bioregion is well represented in the Carpathians as well, with a large mountainous and subalpine level, the alpine level covering a smaller area in few mountain ranges and peaks exceeding an altitude of 2,000 m.
All these mountains show visible traces of the quaternary glacial period with glaciers on a smaller (Pyrenees) or larger scale (Alps). Glaciers do not exist in the Carpathians, but witnesses of the quaternary glacial period are still existent: glacial lakes, glacier calderas and U-shaped glacial valleys, moraines and numerous glacial relict habitats, arctic-alpine relict plants, plant communities and fauna species. Glacial géomorphologie forms and their specific habitats mainly occur in the North- Western part of the Carpathians i.e. in the Slovakian Tatra Mountains, the Southern Carpathians (Fägäras, Cindrel, Parâng, Retezat and Tarcu-Godeanu Mountains) called "Transylvanian Alps" in older scientific papers, as well as in the Rodna Mountains of the Eastern Carpathians (Ozenda, 1988, 1994; Oancea et al., 1987; Drägulescu, Schneider and Benedek, 2007).
The Interpretation Manual of European Union habitats EUR 27 (2007) lists the Alpine pioneer formation of Caricion bicoloris-atrofuscae under number 7240 (Palearctic classification 54.3), a priority habitat with Alpine, peri-Alpine and Northern British communities. They are colonising neutral to slightly acid gravel, sandy, stony, sometimes slightly argillaceous or peaty substrates that are soaked by cold water in moraines and along the edges of springs, rivulets, glacial torrents of the alpine or sub-alpine levels of pure, cold, slowly running rivers and calm backwaters. Permanent or ongoing ground frosts that persist for a longer period of the year are essential for the existence of this habitat type.
The low Vegetation of this habitat type is mainly composed by species of Carex and Juncus of the Caricion bicoloris-atrofuscae alliance. Characteristic plants of this habitat type are Carex atrofusca, Carex bicolor, Carex maritima, Carex microglochin, Carex vaginata, Juncus alpino-articulatus, Juncus arcticus, Juncus castanaeus, Juncus triglumis, Kobresia simpliciuscula, Typha lugdunensis, Typha minima, Typha shuttleworthii, Tofieldia pusilla.
The corresponding categories of the German classification are "6402 Alpine Schwemmböden mit niedriger Vegetation" (Alpine alluvial soils with low vegetation), for the Northern vegetation types "3422 Carex atrofusca-Drepanocladus revolvens-typ" and "3423 Carex saxatilis-Drepanocladus revolvens-typ" . The habitat is mentioned as being associated to humid, extensively managed meadows and communities of the Caricion davallianae alliance that are part of the same Tofieldietala order as is the alliance Caricion bicoloris atrofuscae Nordh. 1939 (= Caricion maritimae Br.-Bl-ap. Volk, 1939).
The comprehensive habitats work on all Romanian habitats, considering not only those of community interest (Donifä et al., 2005) mentions the habitat type 7240 Alpine pioneer formations of Caricion bicoloris-atrofuscae for Romania (Romanian habitat number R5403). However, this habitat type title comprises following the above mentioned authors the SouthEastern Carpathian meso-oligothrophic bogs and fens with Carex rostrata and Sphagnum recurvum and not pioneer habitats of alpine alluvial sites, initial phases of fens as well as transition stages to vegetation of snowmelt depressions. Schneider and Drägulescu (2005) mention the habitat type for Romania noting that the characteristic species are present, but that its existence has to be confirmed and that the repartition of this habitat type still has to be clarified by additional studies.
In the Interpretation Manual for Romania the habitat type is mentioned "with uncertain presence in Romania" (Gafta and Mountford, 2008). This statement is justified by the mentioned authors with the evidence that no corresponding associations have yet been described for Romania, the Caricion bicoloris-atrofuscae alliance has not been indicated for Romania, Carex bicolor is mentioned only for the Rodna Mountains and following recent data Carex atrofusca is (Clocarían, 2009) lacking completely in the Rodna Mountains and in the whole Romanian Carpathians.
The wish to contribute to the clarification of these different facts and opinions gave the impetus to analyse the habitat in its repartition with all characteristic site conditions and species of plants as well as typical macroinvertebrates for such pioneer areas of seepages, springs, smaller and larger water courses, snowmelts, to compare the habitat type 7240 from different area and to find an answer to the question, if the habitat type, even in fragments, can be considered as present in the subalpine and alpine bioregion of the Carpathians.
MATERIALS AND METHODS
The priority habitat type *7240 (European Commission, 2007) has been subject to a comparative analysis of different sites of the alpine bioregion Europe - Alps in Germany, Switzerland, Austria, Italy (Ssymank et al., 1998; Pouchol, 2001; Le réseau Natura 2000 France 2010; Oberdorfer et al., 1998; Ellmauer and Traxler, 2000; Ellmauer, 2005; Provincia Autonoma di Bolzano- Alto Adige, 2011) and the Carpathians with special regard to the Rodna Mountains. It has been effected with special regard to interrelations with neighbouring habitat types in similar site conditions such as seepage area, snow melting area, spring area and alpine rivulet courses with chiono-hygrophytes as part of the Montio-Cardaminetea (Coldea, 1 990) class.
The repartition area of characteristic species for the alliance Caricion bicolorisatrofuscae of the order of Tofildietalia Prsg ap. Oberd. 1949 (=Caricetalia davallianae Br. -Bl. 1949), classe Scheuchzerio-Caricetea nigrae Nordh. 1936 in the bioregion has been considered as well. The points analysed were not only the repartition of characteristic species of the phytocoenological units mentioned, but also the phytocoenological units themselves on the basis of literature data. They were brought together with data of own field experiences in the Alps and Carpathians. Parallel to this data concerning typical fauna elements of the habitat type and generally of the alpine alluvial wetland types, in particular Carabidae species of the Bembidion and Nebria genus have been considered in the analysis of the habitat type. These data were presented in synthetic tables.
RESULTS AND DISCUSSIONS
Ongoing from the general description of the habitat type in the Interpretation Manual (EUR 27/2007), data of countries with alpine bioregions have been analysed to compare these data to those that are indicated as being characteristic of this habitat in each country.
The habitat type represented by the Caricion bicoloris-atrofuscae alliance is well developed in Northern Europe, in the Alps, however, it constitutes a rare glacial relict (Ozenda, 1988). The alliance's eponymous species, i.e. the small sedges Carex bicolor and Carex atrofusca, are both circumpolar arctic-alpine species (Meusel, Jäger and Weinert, 1965). In Central Europe Carex bicolor merely occurs in the Central Pyrenees and disjunctly in the Central Alps and the Northern part of the Carpathians (Meusel, Jäger and Weinert, 1965, p. 108), reaching the Southern border of its distribution in the Alpine mountain system. Few dispersed populations of Carex atrofusca in Central Europe could merely be located in the Central Alps and the Carpathians (Meusel, Jäger and Weinert, 1965, p. 109). Both species are rare (Ozenda, 1988; Adler, Oswald and Fischer, 1994; Oberdorfer, 2001) and occur mainly in alpine, to some extent in subalpine bog and marshland communities or in seepage and high altitude alluvial areas. In these spots Kobresia simpliciuscula, Trichophorum pumilum, Carex maritima, Carex microglochin, Tofieldia pusilla and others may be found all the same (Tab. 1). Both species altogether give the impression that even ecologically speaking they are of arctic provenance (Meusel, Jäger and Weinert, 1965).
The data on the Carpathians Carex bicolor regard the Rodna Mountains in the Northern part of the Eastern Carpathians, where the species originating from the Izvoru Mare Valley occurs on alpine, wet gravel area (Nyarády, 1966). This specification has also been made by Oprea (2005) and Ciocârlan (2000, 2009). Two further locations have been recorded in the Rodna Mountains, in particular Obârsia Rebrei (springs of the Rebra Rivulet) and the Izvorul Cailor-Mountains, where they have been found at an altitude of 1,670 m (Oprea, 2005, pursuant to studies by I. Resmerita 1973 and 1975-1987).
Carex atrofusca has also been mentioned for the Rodna Mountains/ Carpathians (Nyárády, 1966, p. 804-807), however, following newer data, Carex atrofusca Schkuhr does not occur in the Carpathians (Ciocârlan, 2000, 2009), only Carex atrofusca auct. non Schkuhr = Carex atrata L. In this context Oprea (2005) observes, that Carex atrofusca Schkuhr is mentioned in the online data bank of Flora Europaea (www.euromed.org.uk) and notwithstanding this Ciocârlan (2000) states its nonoccurrence.
Habitat type 7240 is listed among the alpine habitat types of Germany (Ssymank et al., 1998), however, it merely occurs in an impoverished form. Juncus alpinus is considered as dominant species, characteristic species are Carex microglochin, Equisetum variegatum, Juncus alpinus, Juncus triglumis, Kobresia simpliciuscula, Tofieldia pusilla, Typha minima and Typha shuttleworthii. The following associations belong to this habitat type and are represented by the Caricion-bicoloris atrofuscae alliance (= Caricion maritimae): Asterò bellidiastro-Kobresietum simpliciusculae (Br.-Bl. ap. Nadig, 1942) Dierssen 1982 Caricetum maritimae Br.-Bl. 18, Juncetum alpino-articulatae (Oberd. 57) Philippi 1960 and Equiseto-Typhetum minimae Br.-Bl. ap. Volk 49 (Ssymank et al., 1998).
In France the habitat type occurs in altogether 20 areas situated in the Central Pyrenees, the areas of Provence- Alpes-Côte D'Azur and Rhône-Alpes (Le réseau Natura 2000 en France 2010, ht1p://na1ura2000.envirormient.gouv.frmabitats/HAB7240.htrril). For France, the associations belonging to this habitat are the following: Juncetum arctici, Junco triglumisCaricetum bicoloris caricetosum maritimae, Junco triglumis-Caricetum bicoloris caricetosum bicoloris, Caricetum microglochinis caricetosum microglochinis, Caricetum microglochinis caricetosum scirpetosum purnili, Caricetum microglochinis caricetosum kobrietosum simpliciusculae, Caricetum atrofusco-vaginatae.
For the Swiss Alps the alpine formations of the Caricion bicoloris-atrofuscae FFH habitat type 7240 have been summarized in an outline (Pouchol, 2010; WWFSwitzerland). In the Central Alps (Switzerland) the vegetation of the Caricion bicolorisatrofuscae alliance, which is now contained as habitat type 7240 in the list of FFH habitat types has been proven with characteristic communities: Kobresietum simpliciusculae Br.-Bl. ap. Nadig 1942, Caricetum maritimae Br.-Bl. 1918, community of Juncus arcticus, Caricetum frigidae Rüb. 1912, and Juncetum alpini (Oberd, 1957) Phil. 1960 (Görs, 1998; Steiner in Grabherr and Mucina, 1993).
In the Austrian Alps the habitat type has been proven in the Salzburg area, Carinthia and Tyrol, the broadest and most representative habitats of this type occurring in the Hohe Tauern National Park (Ellmauer and Traxler, 2000; Ellmauer, 2005; www.hohetauern.at/Natur-Wissen/Wissenschaft.Forschung). This is where species-abundant occurrences of Carex bicolor may be found. The typical phytocoenological units represented in the Austrian Alps included in this habitat type are the alliance Caricion atrofuscae-saxatilis Nordh. 1943 (=Caricion bicoloris-atrofuscae Nordh. 1936) with the associations Juncetum castanei Wagner 19665 and Asterò bellidiastro-Kobresietum simpliciusculae (Br.-Bl. in Nadig 1942) Dierßen 1982, and the alliance Caricion davallianae Klika 1934 with the associations Juncetum alpini Philippi 1960 and Equiseto variegati-Typhetum minimae Br.-Bl. in Volk 1940 (Grabherr and Mucina, 1993).
In the Italian part of the Alps, the habitat type 7240 with characteristic species and phytocoenological units included is mentioned in the area of the Nature Park RieserfernerAhrn, Nature Park Sextner Dolomiten and the Nationalpark Stilfser Joch (Provincia Autonoma di Bolzano- Alto Adige, 2011).
No characteristic associations of this alliance have been recorded for the Romanian Carpathians (Sanda, Ollerer and Burescu, 2008). Merely one association of the Swertio perennis-Caricetum chordorrhizae Coldea (1986) 1990 that has been found in the glacier caldera of Gärgäläu in the Rodna Mountains is assigned to the Tofieldietalia order, alliance Caricion davallianae and have some species which occurs also in the Caricion bicoloris-atrofuscae alliance representing the habitat type 7240. In the Tarcu-Godeanu-Mountains/Southern Carpathians is mentioned the order Tofieldietalia, alliance Caricion davalllianae only with one association Carici flavae-Eriophoretum (Boscaiu, 1971).
For the Rodna Mountains phytocoenoses of the Tofieldietalia order (which include the Caricion bicoloris atrofuscae alliance), are quoted as sporadic and small-scale occurrences (Coldea, 1990). However, when analyzing the presence of this habitat's very specific species and their repartition in the Carpathians, more especially in the Rodna Mountains, it becomes apparent that these species occur all the same in characteristic sites of this habitat type in the Rodna Mountains, even though on a small scale and in close interaction with further habitat types. Comparable habitats may be found in the Fägäras, Parâng, Retezat and Tarcu-Godeanu Mountains of the Southern Carpathians also on a small scale. In the Ministerial Order No. 776/6.VI.2007, annexe 4 are listed four sites of community interest including the habitat type 7240. These sites are: 19. Cäliman-Gurghiu Mountains, Eastern Carpathians, 122. Fägäras Mountains, Southern Carpathians 125. Rodna Mountains, Eastern Carpathians and 217. Retezat Mountains, Southern Carpathians (Minsterul Mediului si Dezvoltärii Durabile 2007).
Arctic-alpine species that are characteristic for this habitat type, for example such as Carex bicolor, Juncus castaneus, Juncus triglumis, Kobresia simpliciuscula and Juncus alpinoarticulatus, occur on wet alluvial soils and spring sites, in seepage areas, along brooks and in the marshes of the Rodna Mountains (Ciocârlan, 2009; Nyárády, 1966). The Izvorul Mare-area, where the first Carex bicolor plant have been recorded (Nyárády, 1966), also shelters Juncus castaneus and Juncus triglumis. Further areas of the Rodna Mountains offer sites to specific species of this specific habitat type (Oprea, 2005; Ciocârlan, 2009). Juncus triglumis, Juncus alpinoarticulatus, Kobresia simpliciuscula and Carex atraía are known for various mountain ranges of the Southern Carpathians (Nyárády, 1966; Boscaiu, 1971; Ciocârlan, 2009; Drägulescu, 2010) and may be counted among the habitat type 7240 species.
Even associations specific for alpine seepages such as e.g. Doronico carpaticiSaxifragetum aizoidis Coldea 1 990 comprise species like Juncus triglumis or Juncus alpinoarticulatus that are also characteristic of habitat type 7240. A close interaction of habitat type 7240 with springs of the Cratoneuro commutati W. Koch 1928 alliance and the interrelation with snowmelt area vegetation become apparent here. When comparing the data of site specific conditions and characteristic species (Tabs. 1, 2 and 3) of the alpine habitat type to the habitat type's characteristic species that occur as well in the Rodna Mountains, it stands to reason that the occurrence, even though fragmentary, is assured for the Rodna Mountains. Except for the Central Alps this habitat type usually occurs in fragments and on a very small scale (Steiner, 1993, in Grabherr and Mucina, 1993). The habitat type's fundamental site factor, i.e. continuous ground frosts that persist for a major part of the year, is given for some places of the subalpine and alpine level of the Rodna Mountains and for all alpine areas of the Carpathians.
For the Northern respectively northwestern part of the Carpathians, especially the High Tatra Mountains, fragmentary occurrences of the habitat type have to be assumed even though they have not been quoted for Slovakia (Seffer and Lasák, 2004). Also in the High Tatra Mountains of Poland, the Northern side of the Tatra it is possible to find fragments of the habitat type 7240 as site conditions as alpine alluvial soils, rivulets, springs, seepage area, snowmelt depressions are know from the area (Mirek and PiekosMirkova, 1992).
In France are mentioned as frequently occuring in the habitat type 7240 also the following species: Polygonum viviparum, Salix foetida, Saliex reticulata, Salix retusa. This list include only the mentioned species in the FFH habitats lists of the countries and the species of the habitat type occuring in Rodna Mountains following one experience in Carpathians/Rodna Mountains and data from literature.
Besides these habitat-specific plant species, certain animal species are just as characterstic of this habitat as they are bound to alpine alluvial sites, seepage areas, snowmelt areas, rivulets, spring areas, initial stades of fens and fens. Moreover, their habitat requires a well determined ground and vegetation structure with specific moisture conditions as well. This mainly concerns ground beetles (Carabidae) species of the Bembidion genus and various Nebria species (Burmeister, 1939; Csiki, 1946; Hurka, 1975).
The data gathered in the Alps of Germany (Ssymank et al., 1998), Switzerland (WWF/CH Pouchol, 2010) and Austria (Elmauer, 2005), - unfortunately no faunistic data were available for France - have been pooled in a comparative table (Tab. 2) together with the data obtained in the Romanian Carpathians/Rodna Mountains (Hurka, 1975). Some species of this habitat type such as Nebria hellwigi Panzer and Nebria germari Heer, occur only in the Alps, others both in the Alps and in the Carpathians as e.g. Bembidion bipunctatum L., Nebria jokischi Sturm, further species such as Bembidion glaciale dacicum Jeannel, Nebria transsylvanica Germ., Nebria carpathica Fuss, Nebria reitteri Ryb. ssp. rodnaensis Horv., Nebria tatrica Mill., Nebria reichi Dej. var. Bissenica E. A. Bielz do only occur in the Carpathians if not merely in the Rodna Mountains (Csiki, 1946; Hurka, 1975, 1997) in typical site conditions of the habitat type 7240 (Burmeister, 1939; Csiki, 1946).
Bembidion glaciale dacicum Jeannel, Bembidion bipunctatum L. ssp. nivale Heer, Nebria transsylvanica Germ, are mentioned from the alpin level of Rodna Mountains (Hurka, 1975) on rivulets, springs, around snowmelt area und seepage area (Csiki, 1946; Hurka, 1975), (Tab. 3).
For the habitat type are mentioned in the Alps (Austria, Switzerland) also a butterfly species Arctiaflavia (Ellmauer, 2005; Pouchol, 2010) and in the Alps in Germany and Austria the spider species Pardosa saturador (Ellmauer, 2005; Ssymank et al., 1998), Pardosa pedestris and Arctosa alpígena (Ssymank et al., 1998).
The column Carpathians include the whole high mountains with alpine level/belt apart from Rodna Mountains, which are given in a separate column.
The habitat type 7240 lives on changes and on a certain dynamics. The prerequisite for its long term sustainability is a periodical disturbance of the sites which is mainly assured by the water: flooding of the sites, aggradation and erosion processes, solifluction and alterations caused by frost and thawing. Whenever site conditions stabilize this leads to a suppression of the species occurring on these alpine alluvial sites by more competitive fen species (see also Ellmauer, 2005). Under stable conditions the habitat may thus transform into a calcareous lightly basiphilous or neutrophilous fen and be replaced by succession processes.
The registration of characteristic sites whilst considering not merely plant but also animal species can help provide a more exhaustive illustration of the habitat type.
Both long-lasting ground frosts and periodical disturbances caused by the waters represent this habitat type's major site factors.
Protosoil sites (alluvial soils) that are influenced by cold water, snowmelt runoff and ground frosts, seepage areas and springs are among the characteristic sites of this pioneer habitat and occur in the Rodna Mountains. They have also been recorded as Carex bicolor sites. The occurrence of further arctic-alpine plant species and their sites, all of them characteristic of this habitat type prove the existence of the latter, even though it occurs in fragments and on a small scale. Moreover, what comes along are the various beetle species, arctic-alpine species all the same, that are bound to this kind of pioneer sites. They emphasize also the habitat's relevance as glacial relict.
It may thus be concluded that the biological-ecological relevance of the habitat type altogether depends on its arctic-alpine plant and animal species, i.e. on glacial relicts of special bio-geographical interest. In this respect the Rodna Mountains play a major role as some species such as e.g. Carex bicolor or Juncus castaneus reach the southernmost point of their disjunct repartition here. Also some ground beetles are located only in these specific habitats in the Rodna Mountains (Hurka, 1997).
Even though the habitat type merely occurs on a small scale, or right therefore, its characteristic species and their repartition are of special relevance. The presence of characteristic species - even though not of all of them - constitutes the only and most important identifying feature of this habitat type (Pouchol, 2010).
Given that the Carpathians' alpine belt is not very broad (Ozenda, 1994), its sustainability and that of further alpine habitat types is threatened with extinction by global warming. This is why it deserves our special interest.
A comparative evaluation of the habitat type's occurrence data and its respective phytocoenosis' in the Alps and the Carpathians reveals clearly that fragmentary sites of the habitat type have been considered all the same and that its existence in an area may even be proven on the basis of such fragments, to which great value is attached (Ozenda, 1988; Ssymank et al., 1998; Ellmauer and Traxler, 2000; Ellmauer, 2005).
The whole complex of site factors, habitat structures and species, plants and animals, mainly macroinvertebrates such as spiders, beetles, butterflies and gastropods, is of major importance as for the identification and delimitation of the habitat type. It frequently reveals necessary to consider very small-scale, mosaic-like areas that are in close interaction with further habitats, e.g. springs, snowmelt depressions plant communities, wetland meadows and fens.
The dynamic factor, i.e. the hydrological dynamics and possible changes in the morphological condition as a consequence of hydrological changes after snowmelt as well as changing water courses, is of major importance for a long term sustainability of this habitat type.
Considering the biogeographical importance of the habitat, it reveals fundamental to study the characteristic sites in greater detail and make them undergo a long-term monitoring.
Thanks to my husband Eckbert Schneider, entomologist, for providing the entomological bibliography and collection of data making it possible to have a more detailed view on habitat type 7240, for which, in the Romanian Habitat Manual, entomological data are not mentioned.
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Erika SCHNEIDER-BINDER *
* Karlsruhe Institute for Technology - University of Land Baden- Württemberg and Research Center of the Helmholtz Society, Institute for Geography and Geoecology, Department WWF-Institute for Floodplain Ecology, Josefstr. 1, Rastatt/Germany, D-76437, email@example.com; firstname.lastname@example.org