Foreword by Dr Martin Warren viii
Preface ix
Acknowledgements xi
1 WHAT IS A HABITAT? AN AWKWARD QUESTION 1
Definitions of habitat 1
Distinguishing habitat from biotope and vegetation units 2
2 A SIMPLE MODEL FOR BUTTERFLY HABITATS 9
Habitat model 9
Key issues in the habitat model 14
The matrix or so-called empty space 14
Movement in and between habitats 14
Open versus closed populations and species 14
Qualifying resource outlets 15
Consumables 23
Larval hostplants and herbivory 23
Nectar sources and adult food 25
Utilities 27
Adult basking sites and behaviour 29
Mate location sites substrates and behaviour 31
Egg-laying sites and substrates 34
Adult rests and roosts 35
Larval sites for resting and moulting 37
Pupation sites 38
Parasitoids and predators in the resource zones 39
Symbionts and enemy-free space 41
Hibernation and aestivation sites 42
Conditions and conditioners 44
Climatic agents as conditioners 44
Edaphic agents as conditioners 51
Resource database 52
3 BASIC PRINCIPLES FOR BUTTERFLY HABITATS 53
Describing variation in resources 53
Resource composition 53
Resource physiognomy 56
Resource connectivity 59
Resource variation in the habitat space 59
General principles of resource composition 59
General principles of resource physiognomy 60
General principles of resource connectivity 61
Resource dynamics within habitats 62
General principles of resource dynamics 63
General principles of resource composition 66
General principles of resource physiognomy 67
General principles of resource connectivity 67
Habitats butterfly resources and population status 68
Resource dynamics population status and life cycle strategies 74
Principles relating to population size and density 75
Principles relating to stage appearance 75
Resources movements and dispersion patterns inside the habitat 77
4 EXPLOITING INDIVIDUAL RESOURCES 79
Patterns and agents in resource use 79
Some principles relating to single resource use 79
Principles relating to spatial variation in a resource type 80
Principles relating to temporal variation in single resource types 84
Principles relating to individual preferences and behaviour 86
Distribution of individuals in relation to the distribution of resources 87
Distribution of individuals on single resource patches 90
Placement of individual butterflies on single resource items 94
Manipulation of the micro-landscape: micro-architecture 95
Foraging: theory and practice 97
5 BUTTERFLY HABITATS: SEARCHING FOR ORDER 100
Biotope distinctions among British butterflies 101
Biotope associations 101
Principles of biotope properties 103
Principles linking butterflies to biotopes 106
Principles relating to observations made in biotopes 108
Biotopes environmental conditions and niche parameters 108
Principles relating to biotopes over time 112
Principles relating to vegetation succession and regeneration cycles 117
Communities niches and invasibility 120
Ecological classification of British butterflies 121
Hostplant strategies and butterfly habitats 122
Searching for ecological order in butterfly life history and resource use 127
6 THE HABITAT CONTEXT FOR BUTTERFLY POPULATIONS 129
From populations to metapopulations 129
Basic principles of metapopulations 130
The link between structure and dynamics in metapopulations 135
Empirical studies of butterfly metapopulations in Britain: habitat quality matters 136
Metapopulations and a resource view of the matrix 142
Boundary issues between patch and matrix 143
Matrix resources and movements 146
Topology for resource use and movement 152
Principles of movements and resource use in real matrix situations 157
Corridors barriers and aggregations 159
From metapopulations to an entire landscape approach 162
7 LANDSCAPE INFLUENCES ON BUTTERFLY HABITATS 165
Landscape-scale studies 165
Landscape components and their influence on butterfly habitat distributions 167
Substrate chemistry and butterfly habitats 167
Substrate moisture and butterfly habitats 168
Substrate exposure and butterfly habitats 177
Vegetation succession and butterfly habitats 177
Light warmth and butterfly habitats 177
Hierarchy and scale recurrence in factors influencing butterfly habitats 178
Influence of landscape and landform elements on butterfly habitats and resources 179
Landform and landscape features 179
Principles of landform and landscape influences on butterfly biology 190
Butterfly landscape divides at the British Isles scale 192
Landscape refuges for butterfly habitats 192
Case examples of the impact of landscape features on butterfly resources 193
Hill tops and hill-topping: a special case? 193
The significance of slopes and their aspects for butterfly habitats and resources 196
Rural 'architecture' and 'furniture' and their impact on butterfly resources 200
Translating concepts from the habitat to a landscape scale 204
Landscape-scale studies on butterflies 205
Cautionary principles for landscape-scale studies 205
Empirical findings of remote landscape-scale butterfly studies 209
Landscape modelling approaches 210
8 HABITAT ISSUES IN BUTTERFLY GEOGRAPHICAL RANGES 211
Components of geographical ranges 211
Definitions of range and distribution 211
Measuring geographical ranges 214
Ecological factors underlying ranges and distributions 215
Species' richness: trends and climate 216
Local population abundance-range size rule 217
Contrasts for species' geographical ranges: ecological and life history influences 218
Altitudinal limits: upper and lower limits of tolerance 224
The importance of range for sourcing islands with butterfly species 228
Butterfly species in cities and conurbations: changes in diversity and incidence 228
Range changes before records 232
Origins 232
Establishment 236
Present and future distributions: climate and land use changes 238
Recent and future range and distribution changes in Britain: basic habitat issues 243
Range and distribution changes: response to specific agents of change 245
Principles associated with climate change 246
Principles associated with habitat destruction pollution and accident 248
9 HABITATS IN BUTTERFLY CONSERVATION 256
Approaches to conservation and conserving butterflies 256
Species or 'habitat' approaches 257
'Habitat' ( npatch) versus landscape approach 258
Single species versus multispecies approach 261
The single site in butterfly conservation 262
Basic principles for within habitat conservation 262
Management and development of existing sites 264
Mapping and predicting butterfly habitats 265
Multiple sites in single and multispecies approaches 268
Basic principles for conserving butterfly habitats at the landscape scale 271
Broad principles for conserving landscapes for multispecies 273
Management and development of existing landscapes 273
Guiding principles for landscape restoration 277
Size shape and placement issues for single habitat patches 278
Internal habitat issues for single patches 284
Principles for patchwork creation 285
Principles for creating networks hierarchies and surfaces 286
Introductions 288
Butterflies as indicators and flagship species 292
Appendices
1 Biodiversity Action Plan (BAP) status legal protection and taxonomic relationships for British butterflies 295
1a: Resident and recently extinct species 295
1b: Rare migrants introductions and/or long-extinct species 297
1c: Taxonomic affinities of British butterflies 298
2 Larval hostplants for British butterflies 299
2a: Status of hostplants 299
2b: Hostplant families range of butterfly herbivory and hostplant phenology 312
2c: Larval hostplant biotopes phenology growth forms environments and life history strategies 314
3 Nectar sources of British butterflies 320
3a: Key flowering nectar plants used by butterfly species 320
3b: Nectar plants supporting 10 or more butterfly species 332
3c: Nectar plant families supporting six or more butterfly species 333
3d: Nectar plants used by butterflies more often or less often than expected 334
3e: Adult feeding: nectar and non-nectar sources 335
4 Statistics on larval host use and adult feeding in British butterflies 337
5 Utility resources and life history data on British butterflies 339
5a: Adult environment 339
5b: Egg environment 341
5c: Larval environment 343
5d: Pupal environment 345
5e: Life history 346
6 Adult and larval behaviour in British butterflies 348
6a: Adult behaviour 348
6b: Larval behaviour 350
7 Biotopes for British butterflies 352
References 354
Index 389
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