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 Copyright © 2010 Peter Martin Rhind endemic mistletoe Viscum album subsp. austriacum, which usually has yellow rather than white berries. 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