Atlantic European Oak Woods

The dominant trees here are either Quercus petraea (sessile oak) or Quercus robur (pedunculate oak).  They tend to occur on the more acidic soils, but nevertheless, these woodlands are often the main climax vegetation. They can have well-developed shrub and ground layers. One of the more widespread and characteristic endemics of the field layer is Hyacinthoides non-scripta bluebell (Liliaceae), which may be so extensive as to create the illusion of a blue haze in places, especially in the United Kingdom. In some mixed oak woods both sessile and pedunculate oaks may occur together, but these are often on highly acidic soils and as a consequence have poorly developed shrub and field layers. In Sherwood Forest, England, for example, other trees in these mixed forests or woodlands include Betula pubescens, Malus sylvestris and Sorbus aucuparia. The sparse scrub layer typically includes Crataegus monogyna and Sambucus nigra, while the characteristic field layer species include bracken (Pteridium aquilinum) and grasses such as Agrostis capillarie, Descampsia flexuosa and Holcus lanatus.

Atlantic European Coastal Temperate Rain Forest

This type of forest is an extremely rare being confined to just seven regions of the world - the Pacific Northwest, the Valdivian forests of south-western South America, New Zealand, Tasmania, the north-eastern Atlantic (including the West Highlands of Scotland, Ireland, Norway and Iceland), south-western Japan and the eastern Black Sea. Because of its requirement for highly oceanic, humid conditions it is only found on the extreme western fringes of the BioProvince where annual rainfall exceeds 2000mm. These forests are some of the most complex and dynamic ecosystems on Earth supporting numerous mosses, lichens, ferns, fungi and invertebrates.  In fact, the biological diversity indices for some taxa (particularly invertebrates, bryophytes, fungi and soil organisms) compare with tropical rainforest. Also like tropical rainforests, their trees are often festooned with epiphytic species, but rather than orchids or bromeliads, the main species are usually lichens and mosses. Here the trees are typically dominated by Quercus petraea and Betula pubescens, but tree cover can be rather open and canopy height rarely exceeds about 20m. Other tree species tend to be scarce, but Corylus avellana and Sorbus aucuparia may form a scattered understorey. There is rarely a well-developed shrub layer but large ferns such as Pteridium aquilinum and Dryopteris dilitata may form sizeable stands, and there may be occasional dwarf ericoid shrubs such as Caluna vulgaris, Erica cinerea or Vaccinium myrtillus. The field layer may include grasses such as Agrostis capillaris, Anthoxanthum odoratum, Deschampsia flexuosa, Festuca ovina, Holcus mollis, and various herbaceous forbs like Anemone nemorosa, Galium saxatile, Hypericun pulchrum, Luzula pilosa, Luzula sylvatica, Melampyrum pratense, Oxalis acetosella, Potentilla erecta, Primula vulgaris, Stellaria holostea, Succisa pratense, Teucrium scorodonia, Veronica chamaedrys, Viola rivinianum, and the endemic Hyacinthoides non-scripta (Liliaceae). Ferns are also well represented and may include Athyrium filix-femina, Blechnum spicant, Dryopteris aemula, Gymnocarpium dryopteris or Thelypteris limbosperma. However, bryophytes often dominate the ground layer and may far exceeding vascular plants in their diversity and abundance. Some of the most frequent of these are Bazzania trilobata, Campylopus paradoxus, Dicrodontium denudatum, Diplophyllum albicans, Hylocomnium plendens, Hylocomnium umbratum, Isothecium myosuroides, Lepidozia reptans, Leucobryum glaucum, Plagiochila spinulosa, Polytrichum formosum, Pseudosleropodium purum, Rhytidiadelphus triquetrus, Saccogyna viticulosa, Scapania gracilis, Thuidium delicantulum and Thuidium tamariscinum.  Also many bryophyte species found here have distributions centred on the Atlantic zone – these so-called Atlantic species include Acrobolus wilsonii, Adelanthus decipiens, Aphanolejeunea microscopica, Colura calyptrifolia, Dicranum scottianum, Drepanolejeunea hamatifolia, Harpalejeunea ovata, Hyocomium flagellare, Lejeunea mandonii, Plagiochila tridenticulata, Radula carringtonii and Radula voluta.  In fact, several Atlantic species found in these woodlands, such as Plagiochila punctata and Plagiochila atlantica are endemic to the Atlantic European BioProvince. Bryophytes, together with lichens, also provide the largest number of epiphytic species. Tree trunks may be smothered in bryophyte species such as Hypnum mammilatum, Isothecium myosuroides and Scapania gracilis, while the smaller branches typically support Frullania tamarisci, Lejeunea ulicina, Ulota crispa and Ulota phyllanthus. Of the ferns Polypodium vulgare is commonly found as an epiphyte on oak, and the two filmy ferns, Hymenophyllum tunbrigense and H. wilsonii may be found among dripping mats of bryophytes.

Atlantic European Beech Woods

Dominated by Fagus sylvatica (beech) these woods tend to be confined to the more shallow, porous soils. They are particularly well established in northwest and central France, southeast England, and Belgium, but also occur as far north as southern Sweden and in the south they occur in the montane regions of northern Spain. Species composition varies from place to place, but on the chalk soils of England, Fagus sylvatica is usually accompanied by Fraxinus excelsior (ash), Prunus avium and Sorbus aria as the main canopy trees, while lower tree levels typically include Ilex aquifolium and Taxus buccata. The low sub-canopy light levels (sometimes down to 2% of incoming radiation) together with heavy leaf litter that is slow to decompose, results in a poor shrub layer and poor but often interesting field layer. Typical shrubs include Buxus sempervirens, Corylus avellana, Euonymus europaeus and Sambucus nigra. In southern England the characteristic field layer includes Aquilegia vulgaris, Sanicula europaea, Helleborus viridis, Polygonatum multiflorum, the orchids Epipactis helleborina, Cephalanthera damasonium, Neottia nidus-avis and the parasite Monotropa hypopitys. In France and Belgium other field layer species may include Helleborus foetidus, Polygonatum odoratum and Vincetoxicum officinale.

Atlantic European Ash Woods

Woodlands dominated by Fraxinus excelsior (ash) are mainly found on limestone or scree in areas north of Atlantic European Beech Woods, while woodlands dominated by the narrow-leaved ash (Fraxinus angustifolia) occur in northern coastal regions of Spain. In England other canopy trees include Sorbus aria, Taxus buccata, Tilia cordata and Ulmus glabra. These are generally open woodlands so have well-developed scrub and field layers. Typical shrubs include Cornus sanquinea, Corylus avellana, Crataegus monogyna, Euonymus europaeus, Rhamnus catharticus, Sambucus nigra and Viburnum lantana. The field layer can be particularly species-rich. Examples include Allium ursinum, Adoxa moschatellina, Aquilegia vulgaris, Campanula latifolia, Galium odoratum, Platanthera chlorantha, while in more northern areas Polemonium caeruleum and Trollius europaeus may be present. Typical field layer species in Fraxinus angustifolia woodland include Ajuga reptans, Euphorbia amygdaloides, Primula vulgaris, Stachys officinalis and the fern Athyrium filix-femina.

Atlantic European Wet Woods

Sometimes referred to as carr these woods of wet and waterlogged areas are usually dominated by Alnus glutinosa (alder), Betula pubescens (downy birch) and various willows such as the local endemic Salix phylicifolia [England, Scotland and Ireland] (Salicaceae). Carr occurs throughout the Atlantic European BioProvince from southwestern Norway to northern Spain. The often-rich shrub layer typically comprises Frangula alnus, Crataegus monogyna, Ligustrum vulgare, Rhamnus catharticus, Viburnum opulus and can include a number fruit bearing bushes such as Ribes nigrum (black currant), R. rubrum (red currant) and R. uva-crispa (gooseberry). Field layers tend to be species-poor but may include Eupatorium cannabinum, Filipendula ulmaria, Iris pseudocorus, Solanum dulcamera, Symphytum officinale, or the fern Thelypterus palustris. Field bindweed (Calystegia sepium) may also cover many of the shrubs. In northern Spain additional field layer species may include Scutellaria minor, Sibthorpia europaea and the royal fern Osmunda regalis.

Atlantic European Coniferous Forests

Although the moist climate of this BioProvince is not ideal for conifers with their adaptions to drought, there are some important pine forests.  The two main species are Pinus sylvestris (scotts pine) in the north and Pinus pinaster (maritime pine) in the south. There are also a few natural forests of Taxus buccata (yew). Natural Scottish pinewoods can be found in the central and eastern Highlands where they are often associated with mountain slopes. Associated trees may include Alnus glutinosa, Betula pubescens, Populus tremula and Sorbus aucuparia. Typical shrub layer species include Calluna vulgaris, Juniperus communis, Vaccinium myrtillus and Vaccinium vitis-idaea. The field layers can, however, be extremely interesting supporting various uncommon northern species such as Moneses uniflora, Orthilia secunda, Pyrola media, P. minor, P. rotundifolia and Trientalis europaea.

Atlantic European Heath

Typically dominated by Calluna vulgaris (heather) or two or three other ericaceous species, heathland is very characteristic of this BioProvince. Gorse, particularly Ulex europaeus, is often a common associate, but in the west this is often joined or replaced by the endemic Ulex gallii (western gorse) (Fabaceae) while in more southern heaths, in southern England and northwestern France, the endemic Ulex micranthus becomes the more characteristic species. The vegetation is best developed in the more western, oceanic zones, where local endemic ericoids such as Daboecia catabrica (St Dabeoc’s heath), Erica mackaiana (Mackay’s heath and E. vagans (Cornish heath) can be found. In more eastern situations such as the sandy heaths of Norfolk and Suffolk (in England) the rare endemic Scleranthus perennis ssp. perennis (perennial knawl) (Caryophyllaceae) may be encounterd. The coastal cliffs of these western zones are often characterised by the presence of climax maritime heath community dominated by Calluna vulgaris and the endemic Scilla verna (spring squill) (Liliaceae). In the northwest and western uplands the ericoid Vaccinium myrtillus becomes a more conspicuous component of the heathlands, and it is within these heaths that the so-called ‘Northern Atlantic mixed hepatic mats’ can be found. This distinct community dominated by Atlantic liverworts is more or less confined to Britain and Ireland, and often includes up to 13 large leafy liverworts. Typical species include Anastrepta orcadensis, Anastrophyllum donianum, A. joergensenii, Bazzania pearsonii, Jamesoniella carringtonii, Mastigophora woodsii, Pleurozia purpurea, Scapania nimbosa, S. ornithopodioides and the endemic taxa Herbertus aduncus subsp. hutchinsiae and Plagiochila carringtonii. Other northern Atlantic heaths are co-dominated by Calluna vulgaris and Juniperus communis subsp. nana. These occur, for example, on the slopes of Beinn Eighe in Scotland, and it is here beneath the dwarf shrubs that the large yellow-orange cushions of the rare endemic, Atlantic liverwort Herbertus borealis occurs. However, apart from in the more exposed coastal and upland situations, most of the heaths in this BioProvince are maintained by human controlled activities such as tree clearing, grazing or burning.

Atlantic European Mires

These may be acidic in the case of bogs where various bog mosses (Sphagnum) often dominate, or alkaline in the case of fens where various sedges and rushes usually comprise the main species. Mires are particularly well developed in the high rainfall areas of the western Atlantic. Typical bog species are Erica tetralix, Eriophorum angustifolium, Molinia caerulea, the insectivorous plant Drosera rotundifolium, and the widespread endemic Narthecium ossifragum bog asphodel (Liliaceae). Other endemics of bogs are rarely as widespread - the endemic bog moss Sphagnum skyense, for example, is confined to the Isle of Skye on the northwest coast of Scotland. Characteristic sedges and rushes of fens include Cladium mariscus, Carex flacca, Carex nigra, Schoenus nigricans, Juncus articulatus and Juncus subnodulosus. They also include a variety of distinctive herbaceous species like Caltha palustris, Epipactis palustris, Dactylorhiza incarnata, Filipendula ulmaria, Lathyrus palustris, Lychnis flos-cuculi, Lysimachia vulgaris, Lythrum salicaria, Peucedanum palustre and Thalictrum flavum. Another type of mire is the so-called rush pasture typically dominated by Juncus effuses or Juncus acutiflorus. An interesting endemic possibly encountered in rush pasture is the stange whorled caraway Carum verticillatum (Apiaceae), although it can also be found in Molinia mire.

Atlantic European Grassland

With the exception of certain alpine and maritime grasslands, virtually all grasslands in Atlantic BioProvince are semi-natural and maintained through various forms of human activity. Grazing by livestock or cutting regularly as a hay crop for silage is two of the more important controlling mechanisms. Grassland types may be acidic with species such as Nardus strictus and Agrostis capillaries, neutral with species such as Cynosurus cristatus and Dactylus glomerata, or calcareous with species such as Avenula pubescens and Festuca ovina. On the damper sites Deschampsia flexuosa or Molinia caerulea usually becomes the main species. Many of the endemics, such as Alchemilla minima (Rosaceae), Gentianella anglica (Gentianaceae), Helianthemum canum ssp. laevigatum (Cistaceae) and Linum perenne ssp. subciliata (Linaceae) are associated with calcareous grassland. However, some of these are very local. Alchemilla minima, for example, is confined to just two hills in northern England.

Atlantic European Sand Dunes

Atlantic dune systems are largely composed of mobile dune with Ammophila arenaria Marram, fixed dune grasslands with Festuca rubra, and dune slacks with Salix repens. There may also be dune heath present with Calluna vulgaris. These largely natural systems support vast numbers of flowering plants. Some of the endemics include Astragalus baionensis [found in France and Spain], Linaria arenaria [found in semi-fixed dunes in France and northwest Spain], Linaria thymifolia [found in France] (Fabaceae), Arabis brownii (Brassicaceae), Carex trinervis [dune slacks] (Cyperaceae), Dianthus gallicus Jersey pink [found in France and Spain] (Caryophyllaceae), Gentianella amarella subsp. septentrionalis  (Gentianaceae), Omphalodes littoralis [found in dune slacks in France and Spain] (Boraginaceae) and the two orchids Dactylorhiza majalis ssp. cambrensis Welsh marsh orchid [dune slacks] and Epipactis leptochila var. dunensis dune helleborine [dune slacks] (Orchidaceae), but none of these can be described as widespread within the BioProvince.

Atlantic European Saltmarsh

Many of the sheltered estuaries support marshes of halophytic (salt-tolerant) vegetation. On the seawards fringes these usually include a pioneer zone of Suaeda maritima and species Salicornia, while the dominant species in the mid-marsh zones is usually the common saltmarsh grass Puccinellia maritima. Other common species in this zone include Aster tripolium and Atriplex portulacoides. In the landward transition zones the grass Elytrigia atherica is typically present. Spartina maritima Small Cord-Grass (Poaceae) is one of the few endemic species.

Atlantic European Maritime Cliffs

On the seaward fringes, the exposed rocky ledges often support Crithmum maritimum and the endemic Spergularia rupicola (Caryophyllaceae) in the south, and Ligusticum scoticum in the north. Further inland rocky cliff-tops usually give way to maritime grassland with Festuca rubra and on exposed sites this may extend inland for up to 500 m. Endemics within this zone are numerous although many such as Angelica polycarpa (Apiaceae), Anthyllis vulneraria ssp. corbierei (Fabaceae), Centaurium scilloides, C. chloodes (Gentianaceae), Coincya wrightii Lundy cabbage (Brassicaceae), Heriaria ciliolata ssp. ciliolata (Caryophyllaceae), Limonium minutum, L. paradoxum, L. recurvum and L. transwallianum (Plumbaginaceae) and Primula scotica Scottish primrose (Primulaceae), are rare with very local distributions.

References

Chilton, L. 1997. Plant List for the Pyrenees. A Marengo Publication.

Eggeling, W. J. 1956. The flowering plants and ferns of the Isle of May, Firth of Forth, Fife. Transactions of the Botanical Society of Edinburgh, 37: 11-23.

Fuller, R.M. 1975. The Culbin shingle bar and its vegetation. Transactions & Proceedings of the Botanical Society of Edinburgh, 42: 293-305.

Fuller, R.M. 1987. Vegetation Establishment on Shingle Beaches. Journal of Ecology, 75: 1077-1089.

Gomez, D., Sese, J. A. & Villar. L. 2003. The Vegetation of the Alpine Zone in the Pyrenees. In: Alpine Biodiversity in Europe. Eds. L. Nagy, G.Grabherr, Ch. Korner; & D.B.A. Thompson. Ecological Studies, Vol. 167. Springer-Verlag Berlin Heidelberg.

Hodgetts, N. 1993. Atlantic bryophytes on the western seaboard. British Wildlife, 4: 287-295.

Lomba, A., Alves, P. & Honrada, J. 2008. Endemic sand dune vegetation of the northwest Iberian Peninsula: diversity, dynamics, and significance for bioindication and monitoring of coastal landscapes. Journal of Coastal Research, 24: 113-121.

Lousley, J. E. 1971. Flora of the Isles of Scilly. David and Charles. Newton Abbot.

Montserrat, P. & Villar, L. 1987. The vegetation and endemic flora of the Spanish Pyrenees. Excursion no. 41. Excursion Guide, XIV International Botanical Congress, Berlin.

Moore, K. 1990. Where Is It and How Much is Left? The State of the Temperate Rainforest in British Columbia. Forest Planning Canada, 6:15.

Oliver, F. W. 1912. The shingle beach as a plant habitat. New Phytologist, 11: 73-99

Packham  J. R. 2001. Ecology and Geomorphology of Coastal Shingle. Westbury Academic & Scientific Publishing.

Polunin, O. & Walters, M. 1985. A guide to the vegetation of Britain and Europe. Oxford University Press.

Poore, M. E. D. & Robertson, V. C. 1949. The vegetation of St Kilda in 1948. Journal of Ecology, 37: 82-99.

Randall, R. E. 1989. Shingle habitats in the British Isles. Botanical Journal of the Linnean Society, 101: 2-18.

Randall, R. E. & Scott, G. A. M. 1997. Communities of Sand and Shingle Beaches. In: Ecosystems of the World. 2C. Dry Coastal Ecosystems. Elsevier.

Ratcliffe, D. A. 1968. An ecological account of the Atlantic bryophytes in the British Isles. New Phytologist, 67: 365-439.

Rhind, P. M. 2003. Comment: Britain’s contribution to global conservation and our coastal temperate rainforest.  British Wildlife, 15: 97-102.

Rich, T. C. G., Hutchinson, G., Randall, R. D. & Ellis, R. G. 1999. List of plants endemic to the British Isles.  BSBI News, 80: 23-27.

Rodwell, J. S. 2000. British Plant Communities Volume 5 - Maritime communities and vegetation of open habitats. Cambridge University Press.

Scott, E. J. 1963. The Ecology of Shingle Beach Plants. Journal of Ecology, 51: 517-527.

Scott, E. J. 1965. The shingle succession at Dungeness. Journal of Ecology, 53: 21-31.

Scott, W., Harvey, P. V., Riddington, R. & Fisher, M. 2002. Rare Plants of Shetland. Shetland Amenity Trust, Lerwick.

Scott, W. & Palmer, R. 1987. The Flowering Plants and Ferns of the Shetland Islands Shetland Times, Lerwick.

Vevers, H. G. 1936. The land vegetation of Ailsa Craig. Journal of Ecology, 24: 424-445.

Villar, L. undated. El Elemento Endémico en la Flora del Pirineo Occidental Español. Instituto Pirenaico de Ecologla. 22700 JACA (Huesca).

Villar, L & Garcia B. 1989. Vers une Banque de Donns des Plantes Vasculaires Endemiques des Pyrenees. Acta Biologica Montana, IX: 261-274

Villar, L., Sesé, J. A., Goni, D., Fernandez, J. V., Guzman, D. & Catalan, P. 1997. Sur La Flore Endémique et Menacée Des Pyrénées (Aragon et Navarre). Lagascalia, 19: 673-684.

Walters, S. M. Endemism in the genus Alchemilla in Europe. In: Taxonomy, Phytogeography and Evolution. Ed. D. H. Valentine. Academic Press.