Habitat from a Butterfly's Point of View
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Université catholique de Louvain Biodiversity Research Centre Louvain-la-Neuve, Belgium Habitat from a butterfly’s point of view: How specialist butterflies map onto suitable resources. Ph-D thesis May 2009 Camille Turlure AUTHOR Camille Turlure † ([email protected]) SUPERVISORS Hans Van Dyck † Nicolas Schtickzelle † JURY MEMBERS René Rezsohazy † (President) André Lejeune † Michel Baguette ¥ Klaus Fisher ƒ Jeremy Thomas ‡ † Université catholique de Louvain ƒ Bayreuth University Biodiversity Research Centre Department of Animal Ecology Ecology and Biogeography Unit P.O. Box 101251 Place Croix du Sud, 4 D- 95440 Bayreuth B-1348 Louvain-la-Neuve Germany Belgium ¥ Museum National d‟Histoire ‡ University of Oxford Naturelle Department of Zoology CNRS UMR MNHN 7179 The Thinbergen Building Avenue du Petit Château, 4 South Park Road F-91800 Brunoy Oxford OX1 3PS France England Table of contents Table of contents 1 Table of contents: illustration cover by Camille Turlure 2 Table of contents GENERAL INTRODUCTION 11 The biodiversity crisis and the need to preserve habitat ....................... 13 A history of habitat definitions .............................................................. 15 The resource-based definition of the habitat ......................................... 20 Objectives of this Ph-D thesis ............................................................... 24 General objective 24 Scientific approach 25 Study system .......................................................................................... 28 Study species ......................................................................................... 34 Lycaena helle 36 Lycaena hippothoe 38 Proclossiana eunomia 40 Clossiana selene 42 Boloria aquilonaris 44 Structure of this PhD-dissertation Brief presentation of the objectives of each chapter ............................. 46 CHAPTER I. 49 Resource-based habitat definition, niche overlap and conservation of two sympatric glacial relict butterflies I.1 Introduction ...................................................................................... 53 I.2 Methods ............................................................................................ 56 3 Study species 56 Study site 56 Caterpillars 59 Adults 61 Niche breadth and overlap 62 Association between resources and their distribution 63 I.3 Results ............................................................................................ 64 Caterpillars 64 Adults 69 Niche breadth and overlap 71 Resources association and distribution 74 I.4 Discussion ...................................................................................... 76 Differences in life history traits 76 Niche breadth and overlap 79 Defining habitat for conservation purposes 80 CHAPTER II. 83 Microclimatic buffering and resource-based habitat in a relict butterfly: significance for conservation under climate change. II.1 Introduction ................................................................................... 87 II.2 Methods ......................................................................................... 92 Study species 92 Study sites and populations 92 Modeling caterpillar presence 93 4 Table of contents Sphagnum hummocks covered by the host plant were mostly used by caterpillars 95 Modeling caterpillar density 95 Caterpillar survival relative to temperature and humidity 97 II.3 Results ........................................................................................... 99 Caterpillar presence relative to vegetation composition 99 Caterpillar behaviour and thermal profiles of hummocks 105 Caterpillar density 107 Temperature and caterpillar survival 112 II.4 Discussion ..................................................................................... 113 Sphagnum hummocks determine caterpillar habitat quality 114 Climate change will influence habitat quality through changes in vegetation composition 116 Mitigation effects of climate change for glacial relict species by peat bog management 118 CHAPTER III. 121 On the consequences of aggressive male mate locating behaviour and microclimate for female host plant use in the butterfly Lycaena hippothoe. III.1 Introduction ................................................................................. 125 III.2 Methods ....................................................................................... 129 Study species 129 Study area and quantifying resources 129 Distribution and density of males and females 130 5 Adult behaviour 131 Egg laying in the field 132 Indoor and outdoor egg-laying experiment 135 Egg development and temperature 136 III.3 Results ......................................................................................... 139 Distribution and density of males and females 139 Adult behaviour 142 Egg-laying sites 144 Egg-laying experiment 147 Larval development, survival and temperature 147 III.4 Discussion ................................................................................... 148 CHAPTER IV. 155 Morphology and mobility are affected by resources grain in butterflies IV.1 Introduction ................................................................................. 159 IV.2 Methods ....................................................................................... 163 Study species 163 1. Species and sites 2. Population characteristics and nectar resources used 3. Specific behaviour of females Measures of resource grain, species mobility and morphological traits 169 1. Resource grain 2. Species mobility 3. Morphological traits 6 Table of contents Are spatial resource repartition and morphological traits in relation with mobility? 169 Modelling morphological trait variations between sites 170 IV.3 Results ......................................................................................... 171 Nectar resources use, flight and feeding behaviour 171 Resource spatial dimension and overlap 173 Relation between resource grain and mobility 174 Relation between morphological traits and mobility 175 Modelling morphological traits variations between sites 176 IV.4 Discussion ................................................................................... 184 Chapter V. 189 Estimating population size with a resource-based habitat approach: a test with butterflies V.1 Introduction .................................................................................. 193 V.2 Methods ........................................................................................ 197 Study species 197 Study areas 197 Resource-based definition of larval habitat quality 198 Estimating population size and density 200 Adult emergence rate 201 Larval parasitism 202 Estimating population size from emerging trap data 202 Test of transferability 205 7 V.3 Results .......................................................................................... 206 Population size and density 206 Emergence rate and habitat quality 206 Parasitism 207 Estimating population size from emerging trap data 209 Transferability 210 V.4 Discussion ..................................................................................... 213 Adult population size was linked to availability and quality of larval habitat 213 Transferability of the resource-based definition between populations 215 Application of the resource_based habitat in Population Viability Analysis 216 GENERAL DISCUSSION 219 Definition of the habitat of the five study species ............................... 222 1. Species-specific resources 222 2. Habitat definitions based on host plant or vegetation type are suitable for specialist species only 223 The resource-based definition of the habitat from a butterfly‟s point of view ..................................................................................................... 237 1. Habitat composition 240 2. Habitat configuration 242 3. Resource availability 244 8 Table of contents 4. The combination of resource composition, configuration and availability determines habitat quality 247 5. Habitat scale and boundaries 249 6. Is the resource-based definition universal? 250 Perspectives ......................................................................................... 251 1. Short term perspectives 251 2. Long term perspectives 253 EPILOGUE .................................................................... 255 A short questionnaire… 258 Was our demonstration convincing? 259 REMERCIEMENTS ......................................................... 261 REFERENCES ................................................................ 269 ANNEXES ..................................................................... 309 9 10 General introduction General introduction 11 General introduction: illustration cover and pictures in Figures i.8, i.9, i.10, i.11 and i.12 by Camille Turlure Aerial pictures in Figure i.4 : Région wallone http://cartographie.wallonie.be/NewPortailCarto 12 General introduction The biodiversity crisis and the need to preserve habitat Current extinction rate of species by far exceeds historical ones and reveals that we face a major biodiversity crisis (Western 1990; Morris 1995; Olson et al. 2002). Human development and biodiversity are closely connected; humankind receives products and services of biodiversity and our (over)use has in turn a strong impact on biodiversity. Nowadays, biodiversity is beyond any shadow of doubt threatened by many processes: mostly by fragmentation, loss and degradation of ecosystems (Fischer 2000; Fahrig 2003; Watling and Donnely 2006;