Central European Vegetation

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Plant Formations in the Central European BioProvince

Peter Martin Rhind

Central European Beech Woodlands Beech (Fagus sylvatica) woods form the natural climax over much of Central Europe where the soils are relatively dry and can extend well into the uplands in the more southern zones. In the north, however, around Sweden it is confined to the lowlands. Beech woodlands are often open with a poorly developed shrub layer, Characteristic ground layer species may include various helleborines such as Cephalanthera damasonium, C. longifolia and C. rubra and sedges such as Carex alba, whilst in others, grasses like Sesleria caerlea or Melica uniflora may predominate, but in some of the more acidic examples, Luzula luzuloides is likely to dominate. There are also a number of endemic ground layer species. For example, in Carpathian beech woods endemics such as Dentaria glandulosa (Brassicaceae), Symphytum cordata (Boraginaceae) and the fern Polystichum braunii (Dryopteridaceae) may be encountered. Fine examples of primeaval beech woods can be found in the limestone Alps of lower Austria including the famous ‘Rothwald’ on the southeastern slopes of Dürrentein near Lunz. These range in altitude from about 940-1480 m. Here the canopy is dominated by Fagus sylvatica together with Acer pseudoplatanus, Picea abies, Ulmus glabra, and on the more acidic soils by Abies alba. Typical shrubs include Daphne mezereum, Lonicera alpigena and Rubus hirtus. At ground level the herb layer is very rich supporting possibly up to a 100 species of vascular plants. Examples include Adenostyles alliariae, Asplenium viridis, Campanula scheuchzeri, Cardamine trifolia, Cicerbita alpina, Denteria enneaphyllos, Euphorbia amygdaloides, Galium austriacum, Homogyne alpina, Lycopodium annotinum, Mycelis muralis, Paris quadrifolia, Phyteuma spicata, Prenanthes purpurea, Senecio fuchsii, Valeriana tripteris, Veratrum album and the central European endemic Helliborus niger (Ranunculaceae).

Central European Hornbeam - Oak Woodlands Dominated by Carpinus betulus (hornbeam) and oak (either Quercus petraea or Q. robur) these woods tend to occur on the more acidic soils, although on very acidic soils oak often becomes dominant. The shrub layer is usually well developed and may include Euonymus verrucosa, Ligustrum vulgare, Rhamnus cathartica, Crataegus monogyna, Cornus mas, Ribes rubrum, Rubus caesius, Sambucus nigra, Sorbus torminalis and Staphylea pinnata, while typical ground layer species may include Campanula rapunculoides, Carex michelii, Dentaria bulbifera, Euphorbia amygdaloides, Fragaria moschata, Galium sylvaticum, Hacquetia epipactis, Hepatica nobilis, Impatiens noli-tangere, Lithospermum purpurocaeruleum, Melittis melissophyllum, Primula veris, Stellaria holostea, Viola mirabilis and so on. However, these woodlands can be divided into a number of different associations. In the Czech Republic, for example, several different types have been recognized depending on location. These have been described as Hercynian oak- hornbean woodlands (including three associations characterised by 1. Melampyrum pratensis, 2. Stellaria holostea and 3. Tilia cordata and Betula pubescens), Pannonian oak-hornbeam woodlands (including one association characterised by Primula veris), Carpathian oak-hornbeam woodlands (including one association characterised by Carex pilosa) and Polonian oak-hornbeam woodlands (including one association characterised by Tilia cordata). Oak-hornbean forest also represents the most extensive forest type in the famous, primeval Bialowieza Forest of Poland and Belurus. However, because of the relatively high protortion of Tilia cordata these forests can be described under the subtype Tilia cordata – Carpinus betulus (Tilio-Carpinetum) alliance. This magical, multilayered forest typically has an upper canopy dominated by Picea abies, Quercus robur and Tilia Copyright © 2010 Peter Martin Rhind cordata often accompanied by Acer platanoides. A second tree layer usually comprises Carpinus betulus and Tilia cordata. Canopy cover in summer can reach levels of up to 90%. The reduced sub-canopy light levels prevent much shrub layer development, but it may include Corylus avellana, Daphne mezereum, Euonymus europaea and E. verrucosa. The herb layer, on the other hand, is often exceptionally well developed and can often be differentiated in to two or three sub layers. Both its vertical and horizontal composition shows marked seasonal variation. In spring the flora is dominated by spring geophytes including Allium ursinum, Anemone nemerosa, A. ranunculoides, Corydalis cava, C. solida, Gagea lutea, G. minima, G. spathacea, Isopyrum thalictroides and Ranunculus ficaria. Among the most constant herb layer components are Asarum europaeum, Carex digitata, Galium odoratum, Hepatica nobilis, Lamiastrum galeobdolon, Milium effusum, Oxalis acetosella, Pulmonaria obscura, Ranunculus lanuginosus, Stellaria holostea and Viola reichenbachiana. Common deciduous plants are Aegopodium podagraria, Dentaria bulbifera, Equisetum pratense and Lathyrus vernus.

Central European Black Alder-Ash Swamp Woodlands Contrary to popular belief woodlands in which black alder (Alnus glutinosa) forms a major component are not restricted floodplains but can extend outside stream and river valley systems where reasonably wet condition prevail. For example, the Spree Forest (Spreewald), a Biosphere Reserve southeast of Berlin and probably the most important alder woodland in Central Europe, extends for some 48000 ha over marshy land. The majority of black alder woods in Central Europe are regarded as forms of the black alder – ash (Fraxinus excelsior) association. In the upper Spreewald the main tree layer is dominated by Alnus glutinosa and Fraxinus excelsior together with Acer pseudoplatanus, Prunus padus, Quercus robur, Tilia cordata and Ulmus laevis. Typical shrubs include Euonymus europaeus, Frangula alnus, Humulus lupulus, Ribes nigrum, R. rubrum, Rubus idaeus, Salix cinerea, S. pentandra and Sambucus nigra. The surprisingly rich herb layer may include depending on degree of wetness species such as Lamium maculatum, Lysimachia thyrsiflora, Peucedanum palustre, Stachys palustris, Thelypteris palustris, etc and a variety of grasses and sedges. Black alder-ash woodland also forms a significant component of the primeval Bialowieza Forest in Poland and Belarus. Here they are a common feature of streamsides subject to periodic inundation but often give way to ash- elm forest where the ground water is lower. The best example can be seen along the course of the Orlowka Stream. In addition to black alder and ash, spruce is also always present here as a canopy or undergrowth tree. Other undergrowth species include Corylus avellana, Euonymus europaea, Prunus padus, Ribes nugrum and Ribes rubrum as well as alder and ash. Composition and structure of the exuberant herb layer displays major seasonal variation, although contribution from spring geophytes is mainly limited to mass displays of Anemone nemorosa. In summer tall perennials up to a metre tall predominate with species like Chaerophyllum hirsutum, Cirsium oleraceum, Filipendula ulmaria and Urtica dioica. Below these are tiers of other species such as Caltha palustre, C. cornuta, Chrysosplenium alternifolium, Equisetum sylvaticum, Hepatica nobilis and Lamiastrum galeobdolon.

Central European Pinus nigra (black pine) Forest These forests in Central Europe are confined to the dry dolomitic soils of Austria where black pine is represented by the near endemic Pinus nigra subsp. nigra (Austrian pine). The shrub layer typically includes Amelanchier ovalis, Cotoneaster integerrimus, C. nebrodensis and Sorbus aria, while characteristic field layer species include Biscutella laevigata, Daphne cneorum, Erica herbacea, Globularia cordifolia, Polygala chamaebuxus, Sesleria albicans and the Austran endemic Callianthemum anemonoides (Ranunculaceae). The pine trees may be parasitized by the unusual endemic or near

Central European Pinus mugo (mountain pine) Scrub Dwarf mountain pine (Pinus mugo) is a bush-forming shrub growing no more than about 3m high. It usually forms the highest coniferous zone in the sub-alpine region reaching altitudes of 2000m, and is especially characteristic of limestone mountains. Other associated shrubs include Daphne striata, Erica herbacea, Rhododendron hirsutum, Sorbus chamaemespilus and in the Carpathian Mountains the endemic Ribes petraeum var. carpaticum (Grossulariaceae).

Central European Inundation Grassland One of the best and most undisturbed examples of central European inundation grasslands can be found on the floodplain of the Morava River separating Austria and Slovakia. After the Second World War it became part of the Iron Curtain border zone and as a result was almost completely closed to any human activity. The area comprises several grassland types including ones dominated or characterized by Carex acuta, Glycera maxima, Phragmites australis, Potentilla anserina, Rorippa amphibia and Scirpus lacustris, but the most extensive inundation meadows are descibed as a Alopecurus pratensis – Cnidium dubium community. These are typically inundated at the start of the growing season but as the summer progresses the surface soil gradually dries out. Nevertheless, these have the highest species diversity of all the inundation communities. In addition to Alopecurus pratensis the main grasses include Agrostis stolonifera, Elymus repens and Poa angustifolia, while other characteristic species include Carex praecox, Clematis integrifolia, Cnidium dubium, Galium boreale, Gratiola officinalis, Inula salicina, Serratula tinctoria and Veronica longifolia. Geophytes include Allium angulosum and Iris sibirica. In the most elevated areas, Festuca nigrescens becomes the main character species. Inundation in these areas is often very short and irregular. Other characteristic species include Colchicum autumnalis, Galium verum, Ranunculus auricomus, Sanguisorba officinalis and Serratula tinctoria. In spring the white flowers of Ornithogallum orthophyllum become a feature. Less common species typically include taxa from drier meadows such as Centaurea jacea and Leucanthemum vulgare. Overall these inundation grasslands support some 540-plant species from a variety of different phytogeographical zones. For example, there are boreal (continental) species such as Barbarea stricta, Inula salicina and Veronica longifolia; subcontinental species such as Carex melanostachya, Plantago altissima and Viola pumila; and euroasiatic species such as Cnidium dubium and Erysimum diffusum. Also present are Pannonian endemics or near endemics like Dianthus pontederae (Caryophyllaceae) and Lathyrus pannonicus subsp. pannonicus (Fabaceae).

Central European Steppe Grassland These grasslands are mainly confined to the xerothermic regions of Bohemia, Moravia, Slovakia and the highlands of southern Poland (Little Poland Highland) where they typically occur on dry, sunny south facing slopes on well drained neutral or alkaline soils such as those produced on limestone or gypsum. Grasses such as Festuca pallens, F. rupicola, F. valesiaca, Koeleria macrantha, Stipa capillata and S. pennata are the dominant species. Their peak growth occurs in late spring or early summer when a number of associated perennials forbs come into flower such as Adonis vernalis (yellow adonis) and Potentilla cinerea before the advent of the dry summer when no species flower. These are usually joined by a variety of small annuals like Arabis recta and Arenaria serpyllifolia. In autumn there is typically a second flush of flowering when species such as Aster amellus (European michaelmas daisy) and Odontites lutea (yellow bartsia) come into flower. Among the endemic species associated with these grasslands are

Central European Parasteppic Grasslands of the High Friulian Plain The steppe-like, dry grasslands found to the south of the Carnic and Julian pre-Alps at altitudes up to 300 m are in one of the wettest parts of Europe, but the gravelly soils are so permeable that most of the precipitation rapidly percolates away. The vegetation is known locally as ‘magredi’ (in the Friulian language), which is derived from the Latin term ‘mager’ suggesting an agriculturally poor area. It can be broadly divided in to three successional types: 1. The typical community characterised by Globularia cordifolia and the endemic Centaurea dichroatha (Asteraceae), 2. An intermediate community characterized by Schoenus nigricans and Chrysopogon gryllus and 3. A mature community characterized by Chrysopogon gryllus and the endemic Chamaecytisus purpureus (Fabaceae). These grasslands have considerable phytogeographical interest in that they support eastern (Illyrian and Pontic) elements, Alpic elements and endemic elements. The Illyrian species close to their northern and western limits include Crepis froelichiana, Dorycnium pentaphyllunum, Genista sericea, Knautia illyrica, Potentilla australis, Satureja variegata and Seseli gouanii. Some of the endemic or subendemic taxa include Brassica glabrescens (Brassicaceae), Buphthalmum salicifolium (Asteraceae), Campanula caespitosa (Campanulaceae), Carex mucronata (Cyperaceae), Centaurea nigrescens (Asteraceae), Daphne cneorum (Thymelaeaceae), Euphorbia triflora ssp. kerneri (Euphorbiaceae), Euphrasia cuspidata (Orobanchaceae), Hieracium porrifolium (Asteraceae), Knautia ressmannii (Dipsacaceae), Matthiola carnica (Brassicaceae) and Rhinanthus freynii (Orobanchaceae).

Central European Scree Vegetation Scree is a widespread feature in the mountains of Central Europe. Despite the unstable nature of this habitat many of the associated plants appear remarkable delicate, but this often belies a well-developed root system, which is frequently accompanied by a mat or tuft forming growth. Nevertheless, much of this vegetation comprises light-requiring plants that would quickly succumb to competition from other plants in less demanding situations. In the Carpathian Mountains, scree is particularly well represented in the Tatras (the highest range of the Carpathians). On calcareous scree one of the most characteristic species is the endemic Papaver alpinum subsp. tatrietum (Papaveraceae). Other species commonly include Cerastium latifolium, Saxifraga cernua and the endemic Cerastium tatrae (Caryophyllaceae). On granitic scree, where more acidic conditions prevail, Oxyria digyna becomes one of the main species, where it is often accompanied by various Carpathian endemics such as Cardaminopsis neglecta and Cochlearia tatrae (Brassicaceae) and Saxifraga carpatica (Saxifragaceae). In the Alps a number of scree vegetation types have been recognized. On limestone and dolomite scree in the high alpine and nival belts Thlaspi rotundifolium often characterizes the vegetation. Other local dominants may include Achillea atrata, Campanula cochlearifolia, Doronicum grandiflorum, Gymnocarpum robertianum, Leucanthemum atrata, Rumex scutatus and Saxifraga oppositifolia. The many alpine endemics or near endemics found in this type of vegetation include Campanula cespitosa (Campanulaceae), Galium helveticum (Rubiaceae), Papaver alpinum (Papaveraceae), Saxifraga biflora (Saxifragaceae), Valeriana supina (Valerianaceae), Viola cacarata and V. cenisia (Violaceae). Scree vegetation characterized by the alpine endemic Leontodon montanus (Asteraceae) occurs on finer more moist, clay-slate screes with quantities of interstitial fine soil. Associate species include Ranunculus parnassifolius and the alpine endemic Saxifraga biflora subsp. macropetala (Saxifagaceae). On the other hand, some of the limestone screes in the subalpine zone with fine-moist soil are characterized by Petasites paradoxus. Other

Copyright © 2010 Peter Martin Rhind characteristic species here include Valeriana montana and the alpine endemic Adenostyles glabra var. calcarea (Asteraceae) and Poa cenisia (Poaceae). Surprisingly acidic, silicate scree tends to be more favourable to plant growth than limestone scree since it usually supports quantities of sandy soil that has a better water holding capacity than the soils of limestone screes. The main plant formation of these acidic screes is characterized by the alpine endemic Androsace alpina (Primulaceae). Other dominants may include Geum reptans and Trisetum spicatum, while typical endemic taxa include Gentiana bavarica var. subacaulis (Gentianaceae) and Saxifraga sequieri (Saxifragaceae). On silicate screes poor in humus, Oxygria digyna together with Adenostyles tomentosa and the alpine endemic Cerastium pedunculatum (Caryophyllaceae) may become conspicuous. Finally, on calcareous slate slopes, which are thought to have characteristic in between limestone and silicate screes, the alpine endemic Draba hoppeana (Brassicaceae) is likely to become the most characteristic species. Others include Artemisia genipi, Crepis rhaetica, Draba fladnicensis and the alpine endemic Pedicularis asplenifolia (Orobanchaceae).

Carpathian Mountain Grasslands In the calcareous zones, much of the upland grasslands are dominated by the endemic Festuca versicolor var. vulgaris (Poaceae), while common associates include the two Carpathian endemics Carex sempervirens subsp. tatrorum (Cyperaceae) and Sesleria tatrae (Poaceae). It extends to the highest calcareous summits in the Tatras and is one of the richest communities of these mountains supporting at least 100 species. Some of these include Androsace chamaejasme, Astragalus frigidus, Cerastium lanatum, Hedysarum obscurum, Oxytropis halleri, Saussurea alpina var. bordasii and Senecio capitatus. In areas where the soil is thin and poorly developed a sedge-dominated community - the principal species being Carex firma, replaces these grasslands. Other associates include Dryas octopetala, a number of cushion plants like Minuartia sedoides, Saxifraga caesia, Silene acaulis subsp. pannonica, and dense mat forming plants such as the endemic Oxytropis carpatica (Fabaceae). On the lower slopes the calcareous grasslands are more likely to be dominated by the two endemic species - Festuca tatrae (Poaceae) and Carex sempervirens subsp. tatrorum (Cyperaceae). These grasslands are again very rich in species including both high mountain and lowland species - examples include Allium montanum, Anthyllis alpestris, Carduus glaucus, Dianthus praecox, Sempervivum soboliferum subsp. preissianum, Thesium alpinum, and the endemic Knautia kitaibelii (Dipsacaceae). In gullies and depressions where snow cover tends to persist for longer and where soil moisture levels are higher the grasslands are typically characterised by Calamagrostis villosa, and on some of the more calcareous substrata by the endemic Festuca carpatica (Poaceae). Other associates include Crepis mollis, Linum extraaxillare Phleum michelii, and the endemic Achillea sudetica (Asteraceae). In still wetter places at the foot of cliffs with dripping water, alongside streams or around springs, grasses are largely repleced by a number of tall perennial forbs - dominant among these are Adenostyles alliariae, Doronicum austriacum and Mulgedium alpinum. It is also in these situations that the two rare endemics Aconitum callibotryon var. firmum and Delphinium oxysepalum (Ranuncuaceae) reside. Moving into more acidic areas on granite, gneisses or quartzite, the principal grassland species are Festuca supina (often in its viviparous form), Oreochloa disticha and Juncus trifidus. The latter species often turns reddish in late summer giving whole mountain slopes a red-brown tinge, and is thought to be responsible for the name Czerwone Wierchy (Red Hills) given to a mountain range in the Tatras. Other associates of these acidic grasslands include Avenastrum versicolor, Pulsatilla alba and the endemic Senecio carpaticus (Asteraceae).

Central European Alpine Grasslands on Carbonate Soils (Alps) Most noteworthy of these alpine grasslands are the blue moorgrass (Sesleria albicans) evergreen sedge (Carex sempervirens) formations which is particularly well represented on limestone rocks of the eastern Alps. This formation provides some of the most spectacular flower shows of the alpine and sub alpine zones. Other characteristic species depending on location include the famous Edelweiss (Leontopodium alpinum) together with Anthyllis alpestris, Carduus defloratus, Carex humilis, Erica herbacea, Helianthemum alpestre, H. grandiflorum, Pedicularis verticillata, Phyteuma obiculare, Polygala chamaebuxus and endemic or near endemic species like Alchemilla hoppeana (Rosaceae), Carex austroalpina (Cyperaceae) and Pedicularis rostrato-capitata (Orobanchaceae). In places where the snow clears early exposing areas to cold winds Sesleria albicans can still survive but a number of the other associates are replaced by cushion forming plants. One of the most characteristic of these is the cushion-sedge (Carex firma). Apparently this species is highly economic with the nutrients of its self-made humus contained within its cushions, and can even re-root into its dead remains. Other cushion plants and low-growing rosette species found under these conditions may include Chamorchis alpina, Crepis kerneri, Gentiana clusii and Saxifraga caesia. Other alpine calcareous grasslands types found in the Alps include rusty sedge (Carex ferruginea) meadows in the outer Alps and the violet fescue (Festuca violacea) pastures of the northern Swiss Alps.

Central European Alpine Acid Grassland (Alps) On crystalline rock and other lime-deficient substrates crocked sedge (Carex curvula) often predominates. The olive-green leaves of this species give the sward a perpetual autumn look and it has very low grazing value. Other flowering plants are not particularly well represented but still add welcome colour to the sward with the yellows of Arnica montana, Leontodon helveticus, Potentilla aurea and Senecio incanus ssp. carniolicus, and the blues of Campanula barbata, Veronica bellidioides and species of Phyteuma. In other acidic areas, such as the slopes near Puschlav the alpine, endemic grass Festuca varia (Poaceae) becomes widespread. A common companion is Carex sempervirens. In the lower alpine and sub-alpine belts Nardus strictus (mat grass) often becomes the dominant grass of acidic areas.

Central European Alpine and Subalpine Heaths (Alps) In the Alps these can be divided into at least three separate associations. In the true alpine zone only a few woody species are able to survive. One of the most important of these is the prostrate azalea Loiseleuria procumbens and a few dwarf willows. The herbaceous willow Salix herbacea may occur in protected snow coombs for example. Creeping azalea heaths are often exposed to strong winds and may not gain a covering of protective snow during the winter and so can experience temperatures down to -40o C. In slightly less exposed areas other dwarf shrubs such as Arctostaphylos uva-ursi and Vaccinium uliginosum may be present. At lower levels, in the subalpine zone, Empetrum hermaphroditum (mountain cranberry) and Vaccinium uliginosum (bog bilberry) can form distinct heaths, but these formations are often protected by a covering on snow in winter, and as a result can grow to their full stature during the growing season, but many stands are now heavily grazed. More conspicuous in the sub alpine zone are alpenrose (Rhododendron ferrugineum) heaths with their displays of bright red flowers.

Central European Alpine Naked Rush-Dwarf Shrub Heath In the Balianske Tatry (limestone) Mountains (Western Carpathians) this formation is described as a Carex carpatica-Elyna (Kobresia) myosuroides (naked rush) association. It

Copyright © 2010 Peter Martin Rhind typically occurs on windswept slopes and ridges. In these harsh environments the shallow soils usually dry out in summer and winter temperatures can be extremely low, but snow cover tends to be light due to windstorms. The vegetation is therefore adapted to both dry and cold conditions and technically described as cryo-xerophytic. The characteristic species, Elyna (Kobresia) myosuroides and the Carpathian endemic Carex carpatica (Cyperaceae), may be accompanied by a sparse undergrowth of cushion forming species like Minuartia sedoides, Saxifraga oppositifolia, S. paniculata and Silene acaulis and ground cover of lichens. Other Carpathian endemics found here include Campanula tatrae (Campanulaceae), Carex tatrorum (Cyperaceae), Delphinium oxysepalum (Ranunculaceae), Erigeron hungaricus (Asteraceae), Genteniella tatrae (Gentianaceae), Leontodon pseudotaraxaci (Asteraceae), Oxytropis carpatica (Fabaceae), Poa carpatica (Poaceae), Primula hungaricus (Primulaceae), Saxifraga wahlenbergii (Saxifragaceae), Sesleria tatrae (Poaceae) and Thymus sudeticus (Lamiaceae). Analagous vegetation in the Alps is also characterised by Elyna myosuroides. Characteristics species here include Carex atrata, Dianthus glacialis, Erigeron uniflorus, Potentilla atrata, Saussurea alpina and the endemic or near endemic Oxytropis jacquinii (Fabaceae) and Sesleria sphaerocephala (Poaceae).

Central European Alpine and Subalpine Rock and Boulder Formations Although lichens and certain bryophytes can colonize bare rock, phanerogams are largely restricted to cracks and crevices. These so-called chasmophytic (crevice dwelling) species include a number of different associations depending on rock type and environmental conditions and typically include a rich variety of endemic taxa. In the Alps Potentilla caulescens is often the main character species of carbonate rock in the true alpine zone. Among its associates are various endemic or near endemic species like Carex mucronata (Cyperaceae), Minuartia rupestris (Caryophyllaceae) and Valeriana saxatilis (Valerianaceae). On the hard limestone of the Swiss Jura the endemic Androsace helvetica (Primulaceae) can also become characteristic with associates such as the endemic Draba ladina (Brassicaceae). Potentilla caulescens can also be a feature of subalpine carbonate rocks. Its associates may include Festuca stenantha and Hieracium humile. On silicate rocks in the Alps Androsace vandellii often characterizes the chasmophytic flora together with species such as the endemic or near endemic Erigeron gaudinii (Asteraceae), Eritrichum nanum (Boraginaceae) and Minuartia cherlerioides (Caryophyllaceae). At lower altitudes Primula hirsutum can become one of the main silicate rock fissure species, while the endemic Phyteuma scheuchzeri (Campanulaceae) is likely to be one of its associates. Many other chasmophytic formations have been described particularly in the Carpathian Mountains.

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