A COMPARATIVE STUDY OF CORYNEPHORUS CANESCENS (L.) P.Beauv. COMMUNITIES OF INLAND SAND DUNES IN ENGLAND AND POLAND ARTHUR GODFREY BLUNT MA MPhil A thesis submitted in partial fulfilment of the requirements of the University of Wolverhampton for the degree of Doctor of Philosophy This research programme was carried out in collaboration with the University of Łódź, Poland November 2006 This work or any part thereof has not previously been presented in any form to the University or to any other body whether for the purposes of assessment, publication or for any other purpose (unless otherwise indicated). Save for any express acknowledgements, references and/or bibliographies cited in the work, I confirm that the intellectual content of the work is the result of my own efforts and of no other person. The right of Arthur Godfrey Blunt to be identified as author of this work is asserted in accordance with ss.77 and 78 of the Copyright, Designs and Patents Act 1988. At this date copyright is owned by the author. Signature………………………………………………………. Date…………………………………………………………….. ABSTRACT Inland sand dunes supporting Grey Hair-grass Corynephorus canescens are a declining European habitat designated for conservation under the EU’s Habitats Directive. In Britain they are confined to a handful of sites in East Anglia and the West Midlands. This study investigated the relationships of the British populations to each other and to populations on five sites in Poland, where C. canescens is still widespread. It also conducted exploratory investigations into factors relevant to the conservation of this ecosystem, particularly in the West Midlands. Data were collected chiefly from 1m2 quadrat samples and direct sampling, which recorded the plants and animals present together with parameters such as vigour and fecundity in C. canescens, amounts of bare sand and litter, and measures of erosion and grazing. These data were variously analysed including by CANOCO multivariate analysis and, for the vegetation, TWINSPAN analysis. 153 taxa of plants and 251 of invertebrates were identified. Though strongly distributed on a regional basis, both flora and invertebrate fauna showed relationships particularly between Polish and West Midlands sites. Analysis of the vegetation suggested that West Midlands vegetation had some associations with C. canescens habitats in Europe and that East Anglian vegetation had links with British coastal C. canescens habitats. The invertebrate fauna showed some complex community relationships in Poland and the West Midlands but less so in East Anglia, while assemblages of invertebrates were associated with various vegetational and abiotic factors. Rabbits and hares were the only vertebrates regularly exploiting C. canescens habitats, which they grazed and, in the former case, produced sand disturbances for colonisation by C. canescens. Ants and to a lesser degree some other invertebrates also produced sand disturbances. Observations made in a preliminary cultivation study in the West Midlands suggested that C. canescens may have a biennial phenology, high fecundity, low germination rates and limited dispersal powers in that region. A trampling investigation suggested that C. canescens may be very sensitive to heavy uncontrolled trampling and to vegetational succession under protection. Stages in succession of the C. canescens community were identified, and suggestions for further study and the conservation of C. canescens were drawn up. i CONTENTS Page Abstract i Table of Contents ii Acknowledgements viii Plate 1: Corynephorus canescens x Chapter 1: Introduction 1.1 Background to this study 1 1.2 Aims of the study 2 1.3 Objectives of the study 3 Chapter 2: Grey Hair-grass Corynephorus canescens: literature review 2.1 Species Description 5 2.2 Distribution 7 2.3 Autecology 12 2.3.1 Soil properties 12 2.3.2 Temperature 13 2.3.3 Sand erosion and accretion 14 2.3.4 Reproduction, survival and longevity 15 2.3.5 Response to competition 17 2.3.6 Grazers and pathogens 18 2.4 Conservation Issues 20 2.4.1 Corynephorus canescens in European legislation: the phytosociological framework 20 2.4.2 Corynephorus canescens conservation in Europe 22 2.4.3 Corynephorus canescens conservation in Britain 25 2.4.4 Case studies: Wangford Warren and Kinver Edge 26 Chapter 3: The Study Sites ii 3.1 Selection of study sites 30 3.1.1 Criteria for selection of study sites 30 3.1.2 Limitations imposed by selection of study sites 32 Page 3.2 List of sites selected for this study 33 3.2.1 Main East Anglian Sites 33 3.2.2 Main West Midlands Sites 33 3.2.3 Polish Sites 34 3.2.4 Other Sites 35 Plate 2: Examples of the situations associated with Corynephorus canescens in this study 36 Chapter 4: General Methods 4.1 Timing and nature of sampling 37 4.2 Authorities followed for identification, nomenclature, fieldwork methods and analytical methods 37 4.3 General methods: physical characteristics 40 4.3.1 Bare sand 40 4.3.2 Litter 40 4.3.3 Sand compaction 41 4.3.4 Grazing impact 41 4.3.5 Erosion impact 42 4.4 General methods: vegetation sampling 42 4.4.1 General inspection 42 4.4.2 1m2 quadrat samples – cover abundance 43 4.4.2.1 Vegetation height 43 4.4.2.2 Number of inflorescences 44 4.4.3 Issues arising from vegetation sampling 45 4.5 General methods: faunal sampling 46 4.5.1 Direct search 47 4.5.2 1m2 quadrat samples 47 4.5.3 Cone traps 48 iii 4.5.4 Inflorescence samples 48 4.5.5 Issues arising from faunal sampling 48 Page 4.6 Analytical methods 52 4.6.1 Canonical correspondence analysis (CANOCO) 52 4.6.2 Two-way Indicator Species Analysis (TWINSPAN) 55 Chapter 5: Preliminary field investigations 5.1 Selection of preliminary field investigations, Aims and Objectives 57 5.2 Phenology of Corynephorus canescens in cultivation in the West Midlands 60 5.2.1 Methods 60 5.2.2 Results 61 5.2.3 Discussion 68 5.2.4 Summary of observations from the preliminary investigation into the phenology of a small population of Corynephorus canescens in cultivation in the West Midlands 73 5.2.5 Suggestions for further study and conservation implications 74 5.3 Trampling investigation 76 5.3.1 Methods 76 5.3.1.1 Layout of the investigation 76 5.3.1.2 Determining the rate of uncontrolled trampling on the new path 79 5.3.1.3 Quadrat sampling 80 5.3.2 Results 81 5.3.2.1 Site KE3 Area A (Vegetated-Uncontrolled Trampling) 81 5.3.2.2 Site KE3 Area B North (Vegetated-Controlled Trampling) 88 5.3.2.3 Site KE3 Area B South (Bare-Controlled Trampling) 94 5.3.3 Discussion 100 iv 5.3.4 Characterising successional stages in the Corynephorus canescens habitat 105 Page 5.3.5 Summary of observations from the trampling investigation 107 5.3.6 Suggestions for further study 109 5.3.7 Potential conservation implications 110 Plate 3: Images from preliminary field investigations 111 Chapter 6: The Corynephorus canescens community (floral) 6.1 Aims and Objectives 112 6.2 Methods 113 6.3 Results 114 6.3.1 Taxonomic composition of the vegetation of the study sites 114 6.3.2 TWINSPAN analysis 119 6.3.3 CANOCO analysis 125 6.4 Discussion 128 6.4.1 Taxonomic composition of the vegetation of the study sites 128 6.4.2 Phytosociological analysis 129 6.4.2.1 Syntaxonomies including Corynephorus canescens 129 6.4.2.2 Polish sites 132 6.4.2.3 East Anglian sites 134 6.4.2.4 West Midlands sites 136 6.4.3 Summary of conclusions from the investigation of the floral community of Corynephorus canescens sites 140 6.4.4 Suggestions for further study 141 Chapter 7: The Corynephorus canescens community (faunal) 7.1 Aims and Objectives 143 7.2 Methods 144 7.3 Results 146 7.3.1 Taxonomic composition of the invertebrate fauna 146 v 7.3.2 Relationship of the invertebrate fauna to sites 157 7.3.3 Relationship of the invertebrate fauna to vegetation 163 Page 7.3.4 Relationship of the invertebrate fauna to environmental factors 174 7.3.5 Interrelationships among the invertebrate fauna 182 7.3.6 The vertebrate fauna 190 7.4 Discussion 193 7.4.1 Faunal composition of Corynephorus canescens sites 193 7.4.2 Relationship of the fauna to vegetational and environmental factors 198 7.4.3 Impact of the fauna on sand erosion and accretion 202 7.4.4 Summary of conclusions from the investigation of the faunal community of Corynephorus canescens sites 204 Plate 4: Impact of man and animals on the Corynephorus canescens habitat 208 Chapter 8: General conclusions and discussion 8.1 Extent to which the Aims of this study are realised 209 8.2 Some main conclusions from this study 212 8.3 Is Corynephorus canescens native in its West Midlands sites? 214 8.4 Conservation implications 215 8.4.1 The classification of Habitat 2330 in Britain 215 8.4.2 Practical conservation measures for Corynephorus canescens: a prescription 216 8.5 Issues for further study 218 References 221 Appendix A: Calculation of trampling pressure on Site KE3 Area A 246 vi Page Appendix B: Detailed descriptions of sites chosen for this study 248 B.1 East Anglian sites: general information 250 B.1.1 Wangford Warren (Site WW) 251 B.1.2 Lakenheath (Site LA) 253 B.2 Kinver Edge: general information 254 B.2.1 Kinver Edge Site 1 (Site KE1) 255 B.2.2 Kinver Edge Site 2 (Site KE2) 257 B.2.3 Kinver Edge Site 3 (Site KE3) 258 B.3.1 Devil’s Spittleful and Rifle Range (Site DS) 259 B.4.1 Burlish Top (Site BT) 261 B.5.1 Alveley Experimental Plot (Site AL) 264 B.6 Bromierzyk sites: general information 265 B.6.1 Bromierzyk Site 1 (Site BR1) 266 B.6.2 Bromierzyk Site 2 (Site BR2) 268 B.6.3 Bromierzyk Site 3 (Site BR3) 270 B.7.1 Bolimów Landscape Park (Site BO) 272 B.8.1 Ciosny Reserve (Site CR) 274 Appendix C: Summary of fieldwork carried out during the study 278 vii ACKNOWLEDGEMENTS This study would not have been possible without the help of many friends and colleagues, and I wish to express my great thanks to them all.