B i B : i ritish and rish utterflies an sland P ersPective Supporting material is available at http://www.cabi.org/openresources/95061. B i ritish and rish B : i utterflies an sland P ersPective roger l.h. dennis and Peter B. hardy CABI is a trading name of CAB International CABI CABI Nosworthy Way 745 Atlantic Avenue Wallingford 8th Floor Oxfordshire OX10 8DE Boston, MA 02111 UK USA Tel: +44 (0)1491 832111 Tel: +1 (617)682-9015 Fax: +44 (0)1491 833508 E-mail: [email protected] E-mail: [email protected] Website: www.cabi.org © Roger L.H. Dennis and Peter B. Hardy 2018. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Names: Dennis, Roger L.H., author. | Hardy, Peter B., author. Title: British and Irish butterflies : an island perspective / Roger L.H. Dennis & Peter B. Hardy. Description: Boston, MA : CABI, [2018] | Includes bibliographical references and index. Identifiers: LCCN 2017050570 (print) | LCCN 2017051602 (ebook) | ISBN 9781786394989 (ePDF) | ISBN 9781786395078 (ePub) | ISBN 9781786395061 (hbk : alk. paper) Subjects: LCSH: Butterflies--British Isles. | Butterflies--Great Britain. | Butterflies--Ireland. | Biogeography--British Isles. | Biogeography--Great Britain. | Biogeography--Ireland. Classification: LCC QL555.B65 (ebook) | LCC QL555.B65 D46 2018 (print) | DDC 595.78/90941--dc23 LC record available at https://lccn.loc.gov/2017050570 ISBN-13: 978 1 78639 506 1 Commissioning editor: Ward Cooper Editorial assistant: Emma McCann Production editor: James Bishop Typeset by SPi, Pondicherry, India Printed and bound in the UK by CPI Group (UK) Ltd, Croydon, CR0 4YY Contents Dedication ix List of Figures xi List of Plates xv Preface xix Acknowledgements xxiii 1. Basics of Island Biogeography 1 The Basic Model of Island Biogeography 1 A Broader View of Island Faunas 5 An Integrative Island Biogeography Model 7 Population and Ecological Changes on Islands 9 Evolutionary Transformations and Novel Species 12 Island Species as Unique Entities 16 2. The British Islands: a Geological and Historical Outline 18 How Many British Islands Are There? 18 The British Islands: Composition, Context and Complexity 19 Glacial–Interglacial Cycles: Ice Landscapes 21 Shifting Sea Levels, Coastal Physiognomy and Island Formation 23 Climate Change During Island Formation 28 Post-Glacial Transformations to Island Landscapes 30 Historical Imperatives 32 3. Island Butterflies: Understanding and Interpreting Records 34 Habitat and Biotope 34 Habitat and Movement 38 Dispersal and Migration: an Island Viewpoint 40 Species Records and What They Imply 44 Bias and Mistaken Records 46 v vi Contents Introductions 47 British and Irish Butterfly Databanks and Sources 48 Selecting Islands for Study 49 4. British and Irish Butterfly Species: Links and Gradients 51 How Unusual Are British Butterflies? 51 Island Environment–Species Gradients 54 A Recording History of Gains and Losses on the British Mainland 57 Predicting British Species 60 Ranges, Distributions and Species Diversity: Detecting Deeper Causes 62 Range Changes in Recent Times 64 Sources and Range Oscillations of Long-distance Migrants 65 5. Island Associations and Species Affinities: Geographical Influences 69 European Islands: the Wider Context 69 Comparing Butterfly Richness Profiles of Britain and Ireland 71 Ireland’s Unusual Case of the Cryptic Wood White 74 The Influence of Geography on Island Species Richness 75 Geography of Archipelagos: the Isles of Scilly 83 Island Assemblages of Butterfly Species 84 Butterfly Associations on British and Irish Islands 86 6. Assessing the Status of Individual Species on Islands: from Geography to Ecology 88 Using Geography to Predict Species Richness 89 Predicting the Incidence of a Butterfly Species on an Offshore Island: the Grayling on Grassholm 90 Predicting Incidences of Butterfly Species on the British and Irish Offshore Islands 92 The Drumlin Islands of Strangford Lough 95 Colonizing Islands: Modelling Individual Probabilities of Success and Failure 97 How Well Does Island Butterfly Incidence Reflect Butterfly Ecology? 101 Using Species’ Ecological Profiles to Examine Inter-island Relationships 103 7. Changing Patterns of Species Incidences on Islands 109 Ecological Bases for Stable and Changing Island Faunas 110 Historical Signals in Geographical and Ecological Data 113 Order and Change in Island Faunas 114 Turnover and Trends: the Case of Hilbre and the Farne Islands 117 Desert Islands: the Perpetual Turnover of Adventives (Itinerants) 121 Big Changes on Big Islands: Uncertain Visits and Problematic Extinctions on the Isle of Man 122 Rare Migrants: the Future Colonists? 124 Contents vii 8. British and Irish Islands: an Evolutionary Vantage 128 Pulses of Loss and Gain: Return Periods and Recurrence Intervals 129 Outlets of Change: Ecological Release and Marginality 132 Glacial Refuges and the Post-glacial Invasion and Colonization Process of Britain and Ireland 133 Butterfly Space–Time Responses to Changing Conditions in Post-Devensian Britain and Ireland 138 Shrinking Habitats and Ecological Islands in Holocene Britain 139 Variation: Genes, Phenes and Plastic Traits 141 Models of Differentiation and Adaptation 144 Adjusting to Living on Small Islands 149 9. Island Studies: a Glance Back and the View Ahead 153 Islands: a Pattern of Intricate and Endless Changes 154 Lessons from Island Studies: the Need to Conserve Sources 157 Directions for Future Research of Island Butterflies 158 Plates 161 1. Island Landscapes 163 2. Images of British Butterfly Species 175 Appendices: Records of Butterflies from British and Irish Islands 187 1. Checklist of British and Irish Butterfly Species 189 2. Butterflies Occurring on the British and Irish Islands as Adventives (Accidental and Deliberate Introductions) 197 3. List of Butterflies on British and Irish Offshore Islands 200 4. Sources of Personal Observations on Island Records 232 5. Regional Atlases of British and Irish Butterflies 236 6. Migration Records and Measures for Butterfly Species Resident on the British and Irish Islands 238 7. Migration Capacity and Colonization Ability Measures for British Butterflies 244 8. Guide to Island Studies: Observations and Analysis 247 9. Predictions for Resident Butterfly Species in the British Isles 256 10. Factors Affecting Butterfly Species Richness on the Offshore British and Irish Islands 258 11. Summary of Predictions for Species Incidences on British and Irish Offshore Islands 264 12. Butterflies Recorded on Strangford Lough Islands, Northern Ireland, During the Butterfly Conservation UK Big Butterfly Counts 274 13. Probabilities of Individual Butterfly Transfers to Islands and Locating a Resource on the Island 278 14. Species Richness and Species Occurrence Predictions for the Isle of Man 285 15. Modelling Evolutionary Changes on a Fossil Island: the Great Orme’s Head, North Wales 290 viii Contents Glossary of Terms 297 Bibliography 317 Index 365 Online Supplementary Appendices Supporting material is available at http://www.cabi.org/openresources/95061. Supplementary Appendices Chapter 3 3.1a A copy of Dennis and Shreeve (1996) Butterflies on British and Irish Offshore Islands: Ecology and Biogeography. Gem Publishing Company, Wallingford, Oxon 3.1b The main data file for British and Irish offshore islands 3.2 Basic ecological and life history data used to build the indices for migration capacity and colonization ability Supplementary Appendices Chapter 4 4.1 Contemporary Geography Study of British Butterflies: Data 4.2 Contemporary Geography Study of British Butterflies: Analyses Supplementary Appendices Chapter 5 5.1a The European Islands Data File: Recent Sources 5.1b The European Islands Data File: Butterfly Records 5.1c The European Islands Data File: Geographical Data 5.2 Comparison of British and Irish Species Distributions Supplementary Appendices Chapter 6 6.1a Species Incidences on Offshore Islands: Logit Regression Analyses 6.1b Species Incidences on Offshore Islands: Discriminant Function Analyses 6.2 Species Richness and Incidences on Offshore Islands: Predictions Supplementary Appendix Chapter 7 7.1 Records and Data for the Isle of Man This work is dedicated to all the butterfly recorders who have kindly contributed to the records and observations for the British and Irish islands, the Isle of Man and the Channel Islands. ix