Cover illustrations: Beinn Eighe, with alpine bearberry Arctostaphylos alpinus in the foreground (front cover); a mosaic of burnt heather Calluna vulgaris and bracken Pteridium aquilinum in the Lammermuir Hills (back cover). (Photos: Laurie Campbell) An illustrated guide to British upland vegetation A M Averis and A B G Averis Haddington, East Lothian HJBBirks Botanical Institute, University of Bergen D Horsfield and D B A Thompson Advisory Services, Scottish Natural Heritage, Edinburgh MJMYeo Joint Nature Conservation Committee, Peterborough Pelagic Publishing PO Box 725 Exeter EX1 9QU UK An illustrated guide to British upland vegetation © JNCC 2004 This reprint edition published by Pelagic Publishing 2014 www.pelagicpublishing.com ISBN-13: 978-1-78427-015-5 This book is a reprint edition of ISBN-10: 1-86107-553-7 published by JNCC. Contents List of colour plates 9 Preface 11 Acknowledgments 13 Introduction 15 Key to upland vegetation types 57 Accounts of vegetation types 94 W3 Salix pentandra-Carex rostrata woodland 95 W4 Betula pubescens-Molinia caerulea woodland 98 W7 Alnus glutinosa-Fraxinus excelsior-Lysimachia nemorum woodland 101 W9 Fraxinus excelsior-Sorbus aucuparia-Mercurialis perennis woodland 104 W11 Quercus petraea-Betula pubescens-Oxalis acetosella woodland 107 W17 Quercus petraea-Betula pubescens-Dicranum majus woodland 111 W18 Pinus sylvestris-Hylocomium splendens woodland 115 W19 Juniperus communis ssp. communis-Oxalis acetosella woodland 119 W20 Salix lapponum-Luzula sylvatica scrub 122 W23 Ulex europaeus-Rubus fruticosus scrub 126 W25 Pteridium aquilinum-Rubus fruticosus underscrub 129 M1 Sphagnum denticulatum bog pool community 132 M2 Sphagnum cuspidatum/fallax bog pool community 135 M3 Eriophorum angustifolium bog pool community 138 M4 Carex rostrata-Sphagnum fallax mire 141 M5 Carex rostrata-Sphagnum squarrosum mire 144 M6 Carex echinata-Sphagnum fallax/denticulatum mire 147 M7 Carex curta-Sphagnum russowii mire 151 M8 Carex rostrata-Sphagnum warnstorfii mire 154 M9 Carex rostrata-Calliergonella cuspidata/Calliergon giganteum mire 157 M10 Carex dioica-Pinguicula vulgaris mire 160 M11 Carex viridula ssp. oedocarpa-Saxifraga aizoides mire 163 M12 Carex saxatilis mire 166 M15 Trichophorum cespitosum-Erica tetralix wet heath 169 M16 Erica tetralix-Sphagnum compactum wet heath 175 M17 Trichophorum cespitosum-Eriophorum vaginatum blanket mire 178 M18 Erica tetralix-Sphagnum papillosum raised and blanket mire 183 M19 Calluna vulgaris-Eriophorum vaginatum blanket mire 186 M20 Eriophorum vaginatum blanket and raised mire 191 M21 Narthecium ossifragum-Sphagnum papillosum valley mire 194 M23 Juncus effusus/acutiflorus-Galium palustre rush-pasture 197 M25 Molinia caerulea-Potentilla erecta mire 200 5 Contents M26 Molinia caerulea-Crepis paludosa mire 204 M27 Filipendula ulmaria-Angelica sylvestris tall-herb fen 207 M28 Iris pseudacorus-Filipendula ulmaria mire 210 M29 Hypericum elodes-Potamogeton polygonifolius soakway 213 M31 Anthelia julacea-Sphagnum denticulatum spring 216 M32 Philonotis fontana-Saxifraga stellaris spring 219 M33 Pohlia wahlenbergii var.glacialisspring 222 M34 Carex viridula ssp. oedocarpa-Koenigia islandica flush 225 M35 Ranunculus omiophyllus-Montia fontana rill 228 M37 Palustriella commutata-Festuca rubra spring 230 M38 Palustriella commutata-Carex nigra spring 233 H4 Ulex gallii-Agrostis curtisii heath 236 H8 Calluna vulgaris-Ulex gallii heath 239 H9 Calluna vulgaris-Deschampsia flexuosa heath 242 H10 Calluna vulgaris-Erica cinerea heath 246 H12 Calluna vulgaris-Vaccinium myrtillus heath 250 H13 Calluna vulgaris-Cladonia arbuscula heath 254 H14 Calluna vulgaris-Racomitrium lanuginosum heath 257 H15 Calluna vulgaris-Juniperus communis ssp.nanaheath 260 H16 Calluna vulgaris-Arctostaphylos uva-ursi heath 263 H17 Calluna vulgaris-Arctostaphylos alpinus heath 266 H18 Vaccinium myrtillus-Deschampsia flexuosa heath 269 H19 Vaccinium myrtillus-Cladonia arbuscula heath 274 H20 Vaccinium myrtillus-Racomitrium lanuginosum heath 277 H21 Calluna vulgaris-Vaccinium myrtillus-Sphagnum capillifolium heath 280 H22 Vaccinium myrtillus-Rubus chamaemorus heath 284 MG2 Arrhenatherum elatius-Filipendula ulmaria tall-herb grassland 288 MG3 Anthoxanthum odoratum-Geranium sylvaticum grassland 291 MG5 Cynosurus cristatus-Centaurea nigra grassland 294 MG8 Cynosurus cristatus-Caltha palustris grassland 297 MG10 Holcus lanatus-Juncus effusus rush-pasture 300 CG9 Sesleria caerulea-Galium sterneri grassland 303 CG10 Festuca ovina-Agrostis capillaris-Thymus polytrichus grassland 307 CG11 Festuca ovina-Agrostis capillaris-Alchemilla alpina grass-heath 311 CG12 Festuca ovina-Alchemilla alpina-Silene acaulis dwarf-herb community 314 CG13 Dryas octopetala-Carex flacca heath 317 CG14 Dryas octopetala-Silene acaulis ledge community 321 U1 Festuca ovina-Agrostis capillaris-Rumex acetosella grassland 324 U2 Deschampsia flexuosa grassland 327 U3 Agrostis curtisii grassland 330 U4 Festuca ovina-Agrostis capillaris-Galium saxatile grassland 333 U5 Nardus stricta-Galium saxatile grassland 339 U6 Juncus squarrosus-Festuca ovina grassland 343 U7 Nardus stricta-Carex bigelowii grass-heath 347 U8 Carex bigelowii-Polytrichum alpinum sedge-heath 350 U9 Juncus trifidus-Racomitrium lanuginosum rush-heath 353 U10 Carex bigelowii-Racomitrium lanuginosum moss-heath 356 U11 Polytrichum sexangulare-Kiaeria starkei snow-bed 360 U12 Salix herbacea-Racomitrium heterostichum snow-bed 363 U13 Deschampsia cespitosa-Galium saxatile grassland 367 U14 Alchemilla alpina-Sibbaldia procumbens dwarf-herb community 371 6 Contents U15 Saxifraga aizoides-Alchemilla glabra banks 374 U16 Luzula sylvatica-Vaccinium myrtillus tall-herb community 377 U17 Luzula sylvatica-Geum rivale tall-herb community 381 U18 Cryptogramma crispa-Athyrium distentifolium snow-bed 385 U19 Oreopteris limbosperma-Blechnum spicant community 388 U20 Pteridium aquilinum-Galium saxatile community 391 U21 Cryptogramma crispa-Deschampsia flexuosa community 394 S9 Carex rostrata swamp 397 S10 Equisetum fluviatile swamp 400 S11 Carex vesicaria swamp 402 S19 Eleocharis palustris swamp 404 S27 Carex rostrata-Potentilla palustris tall-herb fen 406 OV37 Festuca ovina-Minuartia verna community 409 OV38 Gymnocarpium robertianum-Arrhenatherum elatius community 412 OV39 Asplenium trichomanes-Asplenium ruta-muraria community 414 OV40 Asplenium viride-Cystopteris fragilis community 416 Brief accounts of other upland habitats and vegetation types 419 References 433 Glossary 440 Index 444 7 List of colour plates Plate 1 The northern Highlands, Scotland Plate 2 The western Highlands and Islands, Scotland Plate 3 Skye and Arran, Scotland Plate 4 Eastern Scotland Plate 5 Southern Scotland and Northumberland Plate 6 The Lake District and the Pennines Plate 7 Mid-Wales Plate 8 South Wales and south-west England Plate 9 Montane (alpine) vegetation Plate 10 Contrasting forms of wetland vegetation Plate 11 Upland woodlands Plate 12 The Cairngorms – affinities with arctic/alpine tundra Plate 13 Upland plants Plate 14 Contrasts in heathland vegetation in the uplands Plate 15 Heather management in the uplands Plate 16 Forestry conflicted with nature conservation interests in the uplands until the mid-1990s 9 Preface The purpose of this guide to upland vegetation is simple – to focus attention on the nature, diversity and importance of habitats which cover almost a third of Great Britain’s land surface and create landscapes so characteristic of its wilder parts. But this book is much more than that; it is also a tribute to the sustained efforts of a small group of people who have studied the vegetation of the uplands. Forty years ago two remarkable young ecologists introduced their classic volume on Plant Communities of the Scottish Highlands (McVean and Ratcliffe 1962) with the fol- lowing statement: ‘Avegetation monograph can be arranged in three ways – ecologically, or according to the main habitat types and altitudinal zonation; systematically, or according to an hierarchical classification of the units distinguished; and physiognomically, or according to the life-form of the dominant species’. It is a great tribute to the two authors, Donald McVean and Derek Ratcliffe, that their book is still a source of raw data on upland vegetation and an inspiring read. Yet in some ways the issues which these two scholars tackled are still taxing us today. Precisely how does one describe the vegetation seen in the field, and to what extent is it possible to classify this in a systematic way? And to what extent does what we see reflect broader environmental influences on plant growth and development? With the publication of volumes 2 and 3 of British Plant Communities (in 1991 and 1992), edited by John Rodwell, and with substantial contributions from ten leading bota- nists, an important line was drawn under the classification and description of the bulk of Great Britain’s upland vegetation. At long last we had an exhaustive treatment of upland vegetation, with a huge amount of detail on the taxonomy, habitats and ecology of each plant community. But as is so often the way with science, time has brought new information to light. Over the past fifteen or so years, a huge amount of fieldwork by just a handful of ecolo- gists has given new insights into the vegetation of the British uplands. While the basic classification of vegetation is much the same, our knowledge of the different communi- ties, notably their distribution, ecology and conservation importance, has advanced considerably. This is due largely to the work of