Global Climate Change and Terrestrial Invertebrates

Scott N. Johnson is Senior Lecturer in Ecology at the Hawkesbury Institute for the Environment (HIE) at Western Sydney University. T. Hefin Jones is Senior Lecturer in Ecology at the School of Biosciences, Cardiff University, and an Editor of the journals Global Change Biology and Agricultural and Forest Entomology.

List of Contributors xiii

Preface xvii

1 Introduction to Global Climate Change and Terrestrial Invertebrates 1
Scott N. Johnson and T. Hefin Jones

1.1 Background 1

1.2 Predictions for Climate and Atmospheric Change 2

1.3 General Mechanisms for Climate Change Impacts on Invertebrates 2

1.3.1 Direct Impacts on Physiology, Performance and Behaviour 3

1.3.2 Indirect Impacts on Habitats, Resources and Interacting Organisms 3

1.4 Themes of the Book 4

1.4.1 Methods for Studying Invertebrates and Global Climate Change 4

1.4.2 Friends and Foes: Ecosystem Service Providers and Vectors of Disease 4

1.4.3 Multi-Trophic Interactions and Invertebrate Communities 5

1.4.4 Evolution, Intervention and Emerging Perspectives 6

Acknowledgements 7

References 7

Part I Methods for Studying Invertebrates and Climate Change 9

2 Using Historical Data for Studying Range Changes 11
Georgina Palmer and Jane K. Hill

Summary 11

2.1 Introduction 11

2.2 Review of Historical Data Sets on Species’ Distributions 13

2.3 Methods for Using Historical Data to Estimate Species’ Range Changes 15

2.3.1 Measuring Changes in Distribution Size 16

2.3.2 Measuring Change in the Location of Species Ranges 16

2.3.3 An Invertebrate Example: Quantifying Range Shift by the Comma Butterfly Polygonia c-album in Britain 17

2.4 Challenges and Biases in Historical Data 19

2.4.1 Taxonomic Bias 19

2.4.2 Spatial and Temporal Biases 20

2.4.3 Accounting for Temporal and Spatial Biases 21

2.5 New Ways of Analysing Data and Future Perspectives 23

Acknowledgements 24

References 24

3 Experimental Approaches for Assessing Invertebrate Responses to Global Change Factors 30
Richard L. Lindroth and Kenneth F. Raffa

Summary 30

3.1 Introduction 30

3.2 Experimental Scale: Reductionist, Holistic and Integrated Approaches 32

3.3 Experimental Design: Statistical Concerns 33

3.4 Experimental Endpoints: Match Metrics to Systems 35

3.5 Experimental Systems: Manipulations From Bottle to Field 36

3.5.1 Indoor Closed Systems 36

3.5.2 Outdoor Closed Systems 38

3.5.3 Outdoor Open Systems 39

3.6 Team Science: the Human Dimension 40

3.6.1 Personnel 41

3.6.2 Guiding Principles 41

3.6.3 Operation and Communication 41

3.7 Conclusions 41

Acknowledgements 42

References 42

4 Transplant Experiments – a Powerful Method to Study Climate Change Impacts 46
Sabine S. Nooten and Nigel R. Andrew

Summary 46

4.1 Global Climate Change 46

4.2 Climate Change Impacts on Species 47

4.3 Climate Change Impacts on Communities 48

4.4 Common Approaches to Study Climate Change Impacts 48

4.5 Transplant Experiments – a Powerful Tool to Study Climate Change 49

4.5.1 Can Species Adapt to a Warmer Climate? 50

4.5.2 The Potential of Range Shifts 50

4.5.3 Changes in the Timing of Events 51

4.5.4 Shifts in Species Interactions 52

4.5.5 Disentangling Genotypic and Phenotypic Responses 54

4.5.6 Shifts in Communities 54

4.6 Transplant Experiment Trends Using Network Analysis 57

4.7 What’s Missing in Our Current Approaches? Next Steps for Implementing Transplant

Experiments 60

Acknowledgements 62

References 62

Part II Friends and Foes: Ecosystem Service Providers and Vectors of Disease 69

5 Insect Pollinators and Climate Change 71
Jessica R. K. Forrest

Summary 71

5.1 Introduction 71

5.2 The Pattern: Pollinator Populations and Climate Change 72

5.2.1 Phenology 72

5.2.2 Range Shifts 75

5.2.3 Declining Populations 75

5.3 The Process: Direct Effects of Climate Change 76

5.3.1 Warmer Growing-Season Temperatures 76

5.3.2 Warmer Winters and Reductions in Snowpack 79

5.4 The Process: Indirect Effects of Climate Change 81

5.4.1 Interactions with Food Plants 81

5.4.2 Interactions with Natural Enemies 82

5.5 Synthesis, and the View Ahead 83

Acknowledgements 84

References 84

6 Climate Change Effects on Biological Control in Grasslands 92
Philippa J. Gerard and Alison J. Popay

Summary 92

6.1 Introduction 92

6.2 Changes in Plant Biodiversity 94

6.3 Multitrophic Interactions and Food Webs 94

6.3.1 Warming and Predator Behaviour 97

6.3.2 Herbage Productivity and Quality 98

6.3.3 Plant Defence Compounds 98

6.3.4 Fungal Endophytes 100

6.3.5 Changes in Plant Phenology 101

6.4 Greater Exposure to Extreme Events 102

6.4.1 Changes in Precipitation 102

6.4.2 Drought Effects 103

6.5 Range Changes 103

6.6 Greater Exposure to Pest Outbreaks 104

6.7 Non-Target Impacts 104

6.8 Conclusion 105

Acknowledgements 105

References 105

7 Climate Change and Arthropod Ectoparasites and Vectors of Veterinary Importance 111
Hannah Rose Vineer, Lauren Ellse and Richard Wall

Summary 111

7.1 Introduction 111

7.2 Parasite–Host Interactions 113

7.3 Evidence of the Impacts of Climate on Ectoparasites and Vectors 114

7.4 Impact of Human Behaviour and Husbandry on Ectoparasitism 116

7.5 Farmer Intervention as a Density-Dependent Process 118

7.6 Predicting Future Impacts of Climate Change on Ectoparasites and Vectors 118

Acknowledgements 123

References 123

8 Climate Change and the Biology of Insect Vectors of Human Pathogens 126
Luis Fernando Chaves

Summary 126

8.1 Introduction 126

8.2 Interaction with Pathogens 129

8.3 Physiology, Development and Phenology 131

8.4 Population Dynamics, Life History and Interactions with Other Vector Species 132

8.5 Case Study of Forecasts for Vector Distribution Under Climate Change: The Altitudinal Range of Aedes albopictus and Aedes japonicus in Nagasaki, Japan 134

8.6 Vector Ecology and Evolution in Changing Environments 138

Acknowledgements 139

References 140

9 Climate and Atmospheric Change Impacts on Aphids as Vectors of Plant Diseases 148
James M.W. Ryalls and Richard Harrington

Summary 148

9.1 The Disease Pyramid 148

9.1.1 Aphids 149

9.1.2 Host-Plants 152

9.1.3 Viruses 154

9.2 Interactions with the Pyramid 155

9.2.1 Aphid–Host-Plant Interactions 155

9.2.2 Host-Plant–Virus Interactions 158

9.2.3 Virus–Aphid Interactions 160

9.2.4 Aphid–Host-Plant–Virus Interactions 162

9.3 Conclusions and Future Perspectives 162

Acknowledgements 163

References 164

Part III Multi-Trophic Interactions and Invertebrate Communities 177

10 Global Change, Herbivores and Their Natural Enemies 179
William T. Hentley and Ruth N. Wade

Summary 179

10.1 Introduction 180

10.2 Global Climate Change and Insect Herbivores 181

10.3 Global Climate Change and Natural Enemies of Insect Herbivores 185

10.3.1 Elevated Atmospheric CO2 185

10.3.1.1 Prey Location 185

10.3.1.2 Prey Quality 186

10.3.2 Temperature Change 186

10.3.3 Reduction in Mean Precipitation 188

10.3.4 Extreme Events 190

10.3.5 Ozone and UV-B 190

10.4 Multiple Abiotic Factors 191

10.5 Conclusions 192

Acknowledgements 193

References 193

11 Climate Change in the Underworld: Impacts for Soil-Dwelling Invertebrates 201
Ivan Hiltpold, Scott N. Johnson, Renée-Claire Le Bayon and Uffe N. Nielsen

Summary 201

11.1 Introduction 201

11.1.1 Soil Community Responses to Climate Change 202

11.1.2 Scope of the Chapter 202

11.2 Effect of Climate Change on Nematodes: Omnipresent Soil Invertebrates 203

11.2.1 Nematode Responses to eCO2 203

11.2.2 Nematode Responses to Warming 205

11.2.3 Nematode Responses to Altered Precipitation Regimes 206

11.2.4 Ecosystem Level Effects of Nematode Responses to Climate Change 207

11.3 Effect of Climate Change on Insect Root Herbivores, the Grazers of the Dark 207

11.3.1 Insect Root Herbivore Responses to eCO2 208

11.3.2 Insect Root Herbivore Responses to Warming 210

11.3.3 Insect Root Herbivore Responses to Altered Precipitation 210

11.3.4 Soil-Dwelling Insects as Modifiers of Climate Change Effects 211

11.4 Effect of Climate Change on Earthworms: the Crawling Engineers of Soil 212

11.4.1 Earthworm Responses to eCO2 212

11.4.2 Earthworm Responses to Warming and Altered Precipitation 214

11.4.3 Climate Change Modification of Earthworm–Plant–Microbe Interactions 214

11.4.4 Influence of Climate Change on Earthworms in Belowground Food Webs 215

11.4.5 Influence of Climate Change on Earthworm Colonization of New Habitats 215

11.5 Conclusions and Future Perspectives 216

Acknowledgements 217

References 218

12 Impacts of Atmospheric and Precipitation Change on Aboveground-Belowground Invertebrate Interactions 229
Scott N. Johnson, James M.W. Ryalls and Joanna T. Staley

Summary 229

12.1 Introduction 229

12.1.1 Interactions Between Shoot and Root Herbivores 231

12.1.2 Interactions Between Herbivores and Non-Herbivorous Invertebrates 232

12.1.2.1 Detritivore–Shoot Herbivore Interactions 232

12.1.2.2 Root Herbivore–Pollinator Interactions 232

12.2 Atmospheric Change – Elevated Carbon Dioxide Concentrations 233

12.2.1 Impacts of e[CO2] on Interactions Mediated by Plant Trait Modification 233

12.2.2 Impacts of e[CO2] and Warming on Interactions Mediated by Plant Trait Modification 234

12.2.3 Impacts of Aboveground Herbivores on Belowground Invertebrates via Deposition Pathways 234

12.3 Altered Patterns of Precipitation 236

12.3.1 Precipitation Effects on the Outcome of Above–Belowground Interactions 236

12.3.1.1 Case Study – Impacts of Simulated Precipitation Changes on Aboveground–Belowground Interactions in the Brassicaceae 237

12.3.2 Aboveground–Belowground Interactions in Mixed Plant Communities Under Altered Precipitation Scenarios 239

12.3.3 Altered Precipitation Impacts on Decomposer–Herbivore Interactions 240

12.3.4 Impacts of Increased Unpredictability and Variability of Precipitation Events on the Frequency of Above–Belowground Interactions 240

12.4 Conclusions and Future Directions 242

12.4.1 Redressing the Belowground Knowledge Gap 243

12.4.2 Testing Multiple Environmental Factors 243

12.4.3 New Study Systems 244

12.4.4 Closing Remarks 245

Acknowledgements 245

References 245

13 Forest Invertebrate Communities and Atmospheric Change 252
Sarah L. Facey and Andrew N. Gherlenda

Summary 252

13.1 Why Are Forest Invertebrate Communities Important? 253

13.2 Atmospheric Change and Invertebrates 253

13.3 Responses of Forest Invertebrates to Elevated Carbon Dioxide Concentrations 254

13.3.1 Herbivores 254

13.3.2 Natural Enemies 259

13.3.3 Community-Level Responses 259

13.4 Responses of Forest Invertebrates to Elevated Ozone Concentrations 263

13.4.1 Herbivores 263

13.4.2 Natural Enemies 264

13.4.3 Community-Level Studies 265

13.5 Interactions Between Carbon Dioxide and Ozone 265

13.6 Conclusions and Future Directions 267

Acknowledgements 268

References 268

14 Climate Change and Freshwater Invertebrates: Their Role in Reciprocal Freshwater–Terrestrial Resource Fluxes 274
Micael Jonsson and Cristina Canhoto

Summary 274

14.1 Introduction 274

14.2 Climate-Change Effects on Riparian and Shoreline Vegetation 275

14.3 Climate-Change Effects on Runoff of Dissolved Organic Matter 277

14.4 Climate Change Effects on Basal Freshwater Resources Via Modified Terrestrial Inputs 278

14.5 Effects of Altered Terrestrial Resource Fluxes on Freshwater Invertebrates 279

14.6 Direct Effects of Warming on Freshwater Invertebrates 280

14.7 Impacts of Altered Freshwater Invertebrate Emergence on Terrestrial Ecosystems 282

14.8 Conclusions and Research Directions 284

14.8.1 Effects of Simultaneous Changes in Resource Quality and Temperature on Freshwater Invertebrate Secondary Production 284

14.8.2 Effects of Changed Resource Quality and Temperature on the Size Structure of Freshwater Invertebrate Communities 284

14.8.3 Effects of Changed Resource Quality on Elemental Composition (i.e., Stoichiometry, Autochthony versus Allochthony, and PUFA Content) of Freshwater Invertebrates 284

14.8.4 Effects of Changed Freshwater Invertebrate Community Composition and Secondary Production on Freshwater Insect Emergence 285

14.8.5 Effects of Changed Quality (i.e., Size Structure and Elemental Composition) of Emergent Freshwater Insects on Terrestrial Food Webs 285

14.8.6 Effects of Climate Change on Landscape-Scale Cycling of Matter Across the Freshwater–Terrestrial Interface 285

Acknowledgements 286

References 286

15 Climatic Impacts on Invertebrates as Food for Vertebrates 295
Robert J. Thomas, James O. Vafidis and Renata J. Medeiros

Summary 295

15.1 Introduction 295

15.2 Changes in the Abundance of Vertebrates 296

15.2.1 Variation in Demography and Population Size 296

15.2.2 Local Extinctions 299

15.2.3 Global Extinctions 299

15.3 Changes in the Distribution of Vertebrates 300

15.3.1 Geographical Range Shifts 300

15.3.2 Altitudinal Range Shifts 301

15.3.3 Depth Range Shifts 302

15.3.4 Food-Mediated Mechanisms and Trophic Consequences of Range Shifts 302

15.4 Changes in Phenology of Vertebrates, and Their Invertebrate Prey 303

15.4.1 Consequences of Phenological Changes for Trophic Relationships 303

15.4.2 Phenological Mismatches in Marine Ecosystems 303

15.4.3 Phenological Mismatches in Terrestrial Ecosystems 304

15.4.3.1 Behaviour and Ecology of the Vertebrates 305

15.4.3.2 Habitat Differences in Prey Phenology 306

15.5 Conclusions 307

15.6 Postscript: Beyond the Year 2100 308

Acknowledgements 308

References 308

Part IV Evolution, Intervention and Emerging Perspectives 317

16 Evolutionary Responses of Invertebrates to Global Climate Change: the Role of

Life-History Trade-Offs and Multidecadal Climate Shifts 319
Jofre Carnicer, Chris Wheat, Maria Vives, Andreu Ubach, Cristina Domingo, S̈oren Nylin, Constantí Stefanescu, Roger Vila, Christer Wiklund and Josep Peñuelas

Summary 319

16.1 Introduction 319

16.2 Fundamental Trade-Offs Mediating Invertebrate Evolutionary Responses to Global Warming 327

16.2.1 Background 327

16.2.2 Mechanisms Underpinning Trade-Offs 328

16.2.2.1 Endocrine Hormone-Signalling Pathway – Antagonistic Pleiotropy Trade-Off Hypothesis 330

16.2.2.2 The Thermal Stability – Kinetic Efficiency Trade-Off Hypothesis 330

16.2.2.3 Resource-Allocation Trade-Off Hypothesis 331

16.2.2.4 Enzymatic-Multifunctionality (Moonlighting) Hypothesis 331

16.2.2.5 Respiratory Water Loss – Total Gas Exchange Hypothesis 332

16.2.2.6 Water-Loss Trade-Off Hypotheses 332

16.3 The Roles of Multi-Annual Extreme Droughts and Multidecadal Shifts in Drought Regimens in Driving Large-Scale Responses of Insect Populations 333

16.4 Conclusions and New Research Directions 337

Acknowledgements 339

References 339

17 Conservation of Insects in the Face of Global Climate Change 349
Paula Arribas, Pedro Abellán, Josefa Velasco, Andrés Millán and David Sánchez-Fernández

Summary 349

17.1 Introduction 349

17.1.1 Insect Biodiversity 349

17.1.2 Insect Biodiversity and Climate Change: the Research Landscape 350

17.2 Vulnerability Drivers of Insect Species Under Climate Change 352

17.3 Assessment of Insect Species Vulnerability to Climate Change 353

17.4 Management Strategies for Insect Conservation Under Climate Change 355

17.5 Protected Areas and Climate Change 357

17.6 Perspectives on Insect Conservation Facing Climate Change 359

Acknowledgements 360

References 361

18 Emerging Issues and Future Perspectives for Global Climate Change and Invertebrates 368
Scott N. Johnson and T. Hefin Jones

18.1 Preamble 368

18.2 Multiple Organisms, Asynchrony and Adaptation in Climate Change Studies 368

18.3 Multiple Climatic Factors in Research 369

18.4 Research Into Extreme Climatic Events 371

18.5 Climate change and Invertebrate Biosecurity 372

18.6 Concluding Remarks 374

References 374

Species Index 379

Subject Index 385