Description

Book Synopsis
Since its discovery Antarctica has held a deep fascination for biologists. Extreme environmental conditions, seasonality and isolation have lead to some of the most striking examples of natural selection and adaptation on Earth. Paradoxically, some of these adaptations may pose constraints on the ability of the Antarctic biota to respond to climate change. Parts of Antarctica are showing some of the largest changes in temperature and other environmental conditions in the world. In this volume, published in association with the Royal Society,leading polar scientists present a synthesis of the latest research on the biological systems in Antarctica, covering organisms from microbes to vertebrate higher predators. This book comes at a time when new technologies and approaches allow the implications of climate change and other direct human impacts on Antarctica to be viewed at a range of scales; across entire regions, whole ecosystems and down to the level of species and variation within t

Trade Review

“Overall, this book provides a comprehensive overview of Antarctic ecosystems and the open access approach to publication means this volume serves as an easy entre to that literature – many ecologists will benefit from this compilation.” (Austral Ecology, 1 October 2013)

“As an institutional library purchase, I would recommend this book.” (Frontiers of biogeography, 5 January 2013

“This timely summary of the state of Antarctic ecological science provides a springboard for an exciting future of Antarctic research.” (The Quarterly Review of Biology, 1 June 2013)

“Overall, I appreciated the book and found it to be a very good synthesis especially of the marine information.” (Biodiversity and Conservation, 1 October 2012)

“The first of these two books is a good scientific treatise on how snow and ice communities work at the moment, while the second concentrates more on marine environments and their likely future. Both are good and should be in the library.” (British Ecological Society Bulletin, 1 August 2012)

“This book is a must for senior undergraduates, graduate students, and scientists interested in Antarctic ecosystems. Summing Up: Highly recommended. Upper-division undergraduates through professionals.” (Choice, 1 September 2012)



Table of Contents

Contributors, xi

INTRODUCTION: ANTARCTIC ECOLOGY IN A CHANGING WORLD, 1
Andrew Clarke, Nadine M. Johnston, Eugene J. Murphy and Alex D. Rogers

Introduction, 1

Climate change, 2

The historical context, 3

The importance of scale, 3

Fisheries and conservation, 4

Concluding remarks, 6

References, 6

PART 1 TERRESTRIAL AND FRESHWATER HABITATS, 11

1 SPATIAL AND TEMPORAL VARIABILITY IN TERRESTRIAL ANTARCTIC BIODIVERSITY, 13
Steven L. Chown and Peter Convey

1.1 Introduction, 13

1.2 Variation across space, 16

1.2.1 Individual and population levels, 16

1.2.2 Species level, 18

1.2.3 Assemblage and ecosystem levels, 20

1.3 Variation through time, 25

1.3.1 Individual level, 26

1.3.2 Population level, 27

1.3.3 Species level, 29

1.3.4 Assemblage and ecosystem levels, 29

1.4 Conclusions and implications, 30

Acknowledgments, 31

References, 31

2 GLOBAL CHANGE IN A LOW DIVERSITY TERRESTRIAL ECOSYSTEM: THE MCMURDO DRY VALLEYS, 44
Diana H. Wall

2.1 Introduction, 44

2.2 The McMurdo dry valley region, 46

2.3 Above–belowground interactions, 46

2.4 The functioning of low diversity systems, 50

2.5 Effects of global changes on coupled above–belowground subsystems, 51

2.6 Temperature change: warming, 52

2.7 Temperature change: cooling, 54

2.8 Direct human influence: trampling, 54

2.9 UV Radiation, 55

2.10 Concluding remarks, 56

Acknowledgements, 56

References, 56

3 ANTARCTIC LAKES AS MODELS FOR THE STUDY OF MICROBIAL BIODIVERSITY, BIOGEOGRAPHY AND EVOLUTION, 63
David A. Pearce and Johanna Laybourn-Parry

3.1 The variety of antarctic lake types, 63

3.2 The physical and chemical lake environment, 66

3.3 The microbial diversity of antarctic lakes, 66

3.3.1 Methods for exploring Antarctic lake biodiversity, 67

3.3.2 Microbial groups, 69

3.3.3 Protists, 70

3.3.4 Crustacea, 72

3.4 Biogeography, 74

3.4.1 Spatial variation and the global ubiquity hypothesis, 74

3.4.2 Temporal variation and palaeolimnology, 75

3.5 Evolution, 76

3.5.1 Prokaryote physiology, 76

3.5.2 Eukaryote physiology, 77

3.6 Future perspectives, 78

3.7 Acknowledgement, 78

References, 78

PART 2 MARINE HABITATS AND REGIONS, 91

4 THE IMPACT OF REGIONAL CLIMATE CHANGE ON THE MARINE ECOSYSTEM OF THE WESTERN ANTARCTIC PENINSULA, 93
Andrew Clarke, David K. A. Barnes, Thomas J. Bracegirdle, Hugh W. Ducklow, John C. King, Michael P. Meredith, Eugene J. Murphy and Lloyd S. Peck

4.1 Introduction, 93

4.1.1 The oceanographic setting, 96

4.1.2 The historical context, 97

4.2 Predicted environmental changes along the western antarctic peninsula, 98

4.3 Environmental variability and ecological response, 100

4.3.1 Biotic responses to climate change: some general points, 102

4.4 Responses of individual marine species to climate change, 102

4.4.1 Acclimation and evolutionary responses to environmental change in antarctic marine organisms, 104

4.5 Community level responses to climate change, 106

4.6 Ecosystem level responses to climate change, 107

4.7 What biological changes have been observed to date?, 109

4.8 Concluding remarks, 110

Acknowledgements, 110

References, 111

5 THE MARINE SYSTEM OF THE WESTERN ANTARCTIC PENINSULA, 121
Hugh Ducklow, Andrew Clarke, Rebecca Dickhut, Scott C. Doney, Heidi Geisz, Kuan Huang, Douglas G. Martinson, Michael P. Meredith, Holly V. Moeller, Martin Montes-Hugo, Oscar Schofield, Sharon E. Stammerjohn, Debbie Steinberg and William Fraser

5.1 Introduction, 121

5.2 Climate and ice, 123

5.2.1 Surface air temperature, 123

5.2.2 Sea ice, 123

5.2.3 Climate co-variability, 125

5.3 Physical oceanography, 127

5.4 Nutrients and carbon, 130

5.4.1 Nutrients and UCDW intrusions, 130

5.4.2 Carbon cycle, 131

5.4.3 Dissolved organic carbon, 132

5.4.4 Sedimentation and export, 133

5.5 Phytoplankton dynamics, 134

5.5.1 Seasonal scale dynamics, 134

5.5.2 Role of light, 134

5.5.3 Role of nutrients, 136

5.5.4 Annual variability in phytoplankton, 137

5.6 Microbial ecology, 138

5.7 Zooplankton, 140

5.7.1 Community composition and distribution, 140

5.7.2 Long-term trends and climate connections, 142

5.7.3 Grazing and biogeochemical cycling, 142

5.8 Penguins, 143

5.8.1 Contaminants in penguins, 145

5.9 Marine mammals, 146

5.10 Synthesis: food webs of the wap, 147

5.11 Conclusions, 148

Acknowledgements, 149

References, 149

6 SPATIAL AND TEMPORAL OPERATION OF THE SCOTIA SEA ECOSYSTEM, 160
E.J. Murphy, J.L. Watkins, P.N. Trathan, K. Reid, M.P. Meredith, S.L. Hill, S.E. Thorpe, N.M. Johnston, A. Clarke, G.A. Tarling, M.A. Collins, J. Forcada, A. Atkinson, P. Ward, I.J. Staniland, D.W. Pond, R.A. Cavanagh, R.S. Shreeve, R.E. Korb, M.J. Whitehouse, P.G. Rodhouse, P. Enderlein, A.G. Hirst, A.R. Martin, D.R. Briggs, N.J. Cunningham and A.H. Fleming

6.1 Introduction, 160

6.2 Oceanography and sea ice, 163

6.2.1 Upper-ocean circulation and characteristics in the Scotia Sea, 163

6.2.2 Physical variability and long-term change, 167

6.3 Nutrient and plankton dynamics, 168

6.4 Krill in the scotia sea food web, 171

6.4.1 Krill distribution in the Scotia Sea, 171

6.4.2 Krill growth and age in the Scotia Sea, 173

6.4.3 Krill reproduction and recruitment in the Scotia Sea, 174

6.4.4 Krill – habitat interactions in the Scotia Sea, 177

6.4.5 Krill population variability and change in the Scotia Sea, 180

6.4.6 Krill in the Scotia Sea food web, 183

6.5 Food web operation, 184

6.5.1 Trophic links, 184

6.5.2 Spatial operation of the food web, 189

6.6 Ecosystem variability and long-term change, 192

6.7 Concluding comments, 195

Summary, 196

Acknowledgements, 197

References, 197

7 THE ROSS SEA CONTINENTAL SHELF: REGIONAL BIOGEOCHEMICAL CYCLES, TROPHIC INTERACTIONS, AND POTENTIAL FUTURE CHANGES, 213
Walker O. Smith, Jr., David G. Ainley, Riccardo Cattaneo-Vietti and Eileen E. Hofmann

7.1 Introduction, 213

7.2 Physical setting, 214

7.3 Biological setting, 219

7.3.1 Lower trophic levels, 219

7.3.2 Mid-trophic levels, 225

7.3.3 Fishes and mobile predators, 226

7.3.4 Upper trophic levels, 227

7.3.5 Benthos, 229

7.4 Food web and biotic interactions, 230

7.5 Conclusions, 232

7.5.1 Uniqueness of the Ross Sea, 232

7.5.2 Potential impacts of climate change, 233

7.5.3 Conservation and the role of commercial fishing activity in the Ross Sea, 234

7.5.4 Research needs and future directions, 235

Acknowledgements, 235

References, 235

8 PELAGIC ECOSYSTEMS IN THE WATERS OFF EAST ANTARCTICA (30 E–150 E), 243
Stephen Nicol and Ben Raymond

8.1 Introduction, 243

8.2 The region, 245

8.2.1 The east (80 E–150 E), 245

8.2.2 The west (30 E–80 E), 247

8.3 Ecosystem change off east antarctica, 251

Summary, 251

References, 252

9 THE DYNAMIC MOSAIC, 255
David K.A. Barnes and Kathleen E. Conlan

9.1 Introduction, 255

9.2 Historical and geographic perspectives, 256

9.3 Disturbance, 257

9.3.1 Ice effects, 258

9.3.2 Asteroid impacts, 260

9.3.3 Sediment instability and hypoxia, 261

9.3.4 Wind and wave action, 261

9.3.5 Pollution, 262

9.3.6 UV irradiation, 263

9.3.7 Volcanic eruptions, 263

9.3.8 Trawling, 264

9.3.9 Non-indigenous species (NIS), 264

9.3.10 Freshwater, 265

9.3.11 Temperature stress, 265

9.3.12 Biological agents of physical disturbance, 266

9.4 Colonisaton of antarctic sea-beds, 266

9.4.1 Larval abundance, 266

9.4.2 Hard substrata, 266

9.4.3 Soft sediments, 269

9.5 Implications of climate change, 276

9.6 Conclusion, 279

Acknowledgements, 280

References, 281

10 SOUTHERN OCEAN DEEP BENTHIC BIODIVERSITY, 291
A. Brandt, C. De Broyer, B. Ebbe, K.E. Ellingsen, A.J. Gooday, D. Janussen, S. Kaiser, K. Linse, M. Schueller, M.R.A. Thomson, P.A. Tyler and A. Vanreusel

10.1 Introduction, 291

10.2 History of antarctic biodiversity work, 293

10.3 Geological history and evolution of the antarctic, 294

10.3.1 Indian Ocean, 294

10.3.2 South Atlantic, 294

10.3.3 Weddell Sea, 295

10.3.4 Drake Passage and Scotia Sea, 296

10.4 Benthic composition and diversity of meio-, macro- and megabenthos, 296

10.4.1 Meiofauna, 297

10.4.2 Macrofaunal composition and diversity, 299

10.4.3 Megafaunal composition and diversity, 304

10.5 Phylogenetic relationships of selected taxa, 308

10.5.1 Foraminifera, 308

10.5.2 Isopoda, 308

10.5.3 Tanaidacea, 309

10.5.4 Bivalvia, 310

10.5.5 Polychaeta, 310

10.5.6 Cephalopoda, 310

10.6 Biogeography and endemism, 311

10.6.1 Porifera, 311

10.6.2 Foraminifera, 311

10.6.3 Metazoan meiofauna, 311

10.6.4 Peracarida, 312

10.6.5 Mollusca, 312

10.6.6 Echinodermata, 313

10.6.7 Brachiopoda, 313

10.6.8 Polychaeta, 313

10.6.9 Bryozoa, 313

10.7 Relationship of selected faunal assemblages to environmental variables, 313

10.7.1 Large-scale patterns with depth, 313

10.7.2 Patterns influenced by other environmental or physical factors, 317

10.7.3 Isopoda, 318

10.8 Similarities and differences between antarctic and other deep-sea systems, 318

10.8.1 The environment, 318

10.8.2 A direct comparison between the deep sea of the SO and the World Ocean, 319

10.8.3 Dispersal and recruitment between the SO and the rest of the world, 320

10.8.4 The special case of chemosynthetically-driven deep-sea systems, 320

10.9 Conclusions, 321

Acknowledgements, 321

References, 323

11 ENVIRONMENTAL FORCING AND SOUTHERN OCEAN MARINE PREDATOR POPULATIONS, 335
Phil N. Trathan, Jaume Forcada and Eugene J. Murphy

11.1 Climate change: recent, rapid, regional warming, 335

11.2 Using oscillatory climate signals to predict future change in biological communities, 337

11.3 Potential for regional impacts on the biosphere, 338

11.4 Confounding isues in identifying a biological signal, 339

11.5 Regional ecosystem responses as a consequence of variation in regional food webs, 340

11.6 Where biological signals will be most apparent, 340

11.7 The southwest atlantic, 341

11.8 The indian ocean, 344

11.9 The pacific ocean, 345

11.10 Similarities between the atlantic, indian and pacific oceans, 346

11.11 What ENSO can tell us, 347

11.12 Future scenarios, 349

References, 349

PART 3 MOLECULAR ADAPTATIONS AND EVOLUTION, 355

12 MOLECULAR ECOPHYSIOLOGY OF ANTARCTIC NOTOTHENIOID FISHES, 357
C.-H. Christina Cheng and H. William Detrich III

12.1 Introduction, 357

12.2 Surviving the big chill – notothenioid freezing avoidance by antifreeze proteins, 358

12.2.1 Freezing challenge in frigid Antarctic marine environment, 358

12.2.2 Historical paradigm of teleost freezing avoidance, 360

12.2.3 Paradigm shift I: the ‘larval paradox’, 360

12.2.4 Paradigm shift II: liver is not the source of blood AFGP in notothenioids, 362

12.2.5 Gut versus blood – importance of intestinal freeze avoidance, 363

12.2.6 Non-hepatic source of plasma AFGP, 364

12.2.7 Alterations in environments and dynamic evolutionary change in notothenioid AFGP gene families, 364

12.2.8 Summary comments – antifreeze protein gain in Antarctic notothenioid fish, 367

12.3 Haemoprotein loss and cardiovascular adaptation in icefishes – dr. no to the rescue?, 367

12.3.1 Vertebrates without haemoglobins – you must be kidding!, 367

12.3.2 Haemoprotein loss in icefishes: an evolutionary perspective, 368

12.3.3 Cellular correlates of haemoprotein loss, 370

12.3.4 The icefish cardiovascular system, 371

12.3.5 Compensatory adjustment of the icefish cardiovascular system in a regime of reduced interspecific competition? Enter Dr. NO, 371

12.3.6 Haemoproteins, NO metabolism, and icefish evolution, 372

12.3.7 Icefishes and erythropoietic gene discovery, 372

12.3.8 Summary comments: haemoprotein loss in Antarctic icefishes, 374

12.4 Concluding remarks, 374

Acknowledgements, 374

Dedication, 374

References, 374

13 MECHANISMS DEFINING THERMAL LIMITS AND ADAPTATION IN MARINE ECTOTHERMS: AN INTEGRATIVE VIEW, 379
Hans O. P€ortner, Lloyd S. Peck and George N. Somero

13.1 Introduction: climate-dependent evolution of antarctic fauna, 379

13.2 Phenomena of thermal specialization and limitation, 382

13.2.1 Molecular and membrane aspects, 383

13.2.2 Genomic aspects: gene expression and loss of genetic information, 390

13.2.3 From molecular to systemic aspects: thermal limitation, 393

13.2.4 From molecular to systemic aspects: thermal adaptation of performance capacity, 397

13.2.5 Ecological implications, 399

13.2.6 Integration of phenomena: concepts, results and perspectives, 405

Acknowledgements, 409

References, 409

14 EVOLUTION AND BIODIVERSITY OF ANTARCTIC ORGANISMS, 417
Alex D. Rogers

14.1 Introduction, 417

14.2 The antarctic biota, 418

14.3 The break-up of gondwana and the evolution of the southern hemisphere biota, 420

14.3.1 Vicariance versus dispersal, 420

14.3.2 Dispersal mechanisms, 421

14.4 The evolution and biodiversity of the terrestrial sub-antarctic and antarctic biota, 423

14.4.1 Plants, 423

14.4.2 Animals, 427

14.5 The marine environment, 432

14.5.1 Biogeography and macroevolution, 432

14.5.2 Notothenioid fish, 432

14.5.3 Birds, 435

14.5.4 Marine invertebrates, 436

14.5.5 The molecular ecology and phylogeography of the marine biota, 437

14.5.6 Patterns of genetic variation in marine species, 448

14.6 Antarctica: a climatic crucible of evolution, 450

14.7 The historical constraints on adaptation to present climate change, 453

14.8 Future directions for research, 453

References, 454

PART 4 CONSERVATION AND MANAGEMENT ASPECTS, 469

15 BIOGEOGRAPHY AND REGIONAL CLASSIFICATIONS OF ANTARCTICA, 471
P. Convey, D.K.A. Barnes, H.J. Griffiths, S.M. Grant, K. Linse and D.N. Thomas

15.1 Introduction, 471

15.2 Historical background, 474

15.2.1 Physical regions in the marine environment, 474

15.2.2 Smaller-scale regionalization within the Antarctic marine environment, 474

15.2.3 Physical regions in the littoral environment, 475

15.2.4 Physical regions in the terrestrial environment, 475

15.3 Data availability, 476

15.4 Different realms in the marine and terrestrial environments, 477

15.4.1 Pelagic realm, 477

15.4.2 Sea ice, 478

15.4.3 Benthic realm, 479

15.4.4 The terrestrial environment, 479

15.4.5 Biogeographical patterns in the terrestrial environment, 480

15.4.6 Biogeographic patterns in the marine environment, 481

15.5 Overview, 485

Acknowledgements, 486

References, 486

16 CONSERVATION AND MANAGEMENT OF ANTARCTIC ECOSYSTEMS, 492
Susie M. Grant, Pete Convey, Kevin A. Hughes, Richard A. Phillips and Phil N. Trathan

16.1 Introduction, 492

16.2 Legal frameworks for conservation and management, 495

16.2.1 Early regulation of marine living resource harvesting, 495

16.2.2 The Antarctic Treaty System, 497

16.2.3 Other (non-ATS) agreements and tools relevant to conservation and management, 500

16.3 Conservation and management measures, 502

16.3.1 Pollution and local disturbance, 502

16.3.2 Biosecurity and non-native species, 505

16.3.3 Conservation and management of marine living resources, 505

16.3.4 Conservation of other individual species, 507

16.3.5 Protected areas, 509

16.4 Conservation science and monitoring, 512

16.5 Future challenges, 515

16.6 Conclusions, 520

Acknowledgements, 521

References, 521

Index, 526

Antarctic Ecosystems

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    A Hardback by Alex D. Rogers, Nadine M. Johnston, Eugene J. Murphy

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      Publisher: John Wiley and Sons Ltd
      Publication Date: 17/02/2012
      ISBN13: 9781405198400, 978-1405198400
      ISBN10: 1405198400

      Description

      Book Synopsis
      Since its discovery Antarctica has held a deep fascination for biologists. Extreme environmental conditions, seasonality and isolation have lead to some of the most striking examples of natural selection and adaptation on Earth. Paradoxically, some of these adaptations may pose constraints on the ability of the Antarctic biota to respond to climate change. Parts of Antarctica are showing some of the largest changes in temperature and other environmental conditions in the world. In this volume, published in association with the Royal Society,leading polar scientists present a synthesis of the latest research on the biological systems in Antarctica, covering organisms from microbes to vertebrate higher predators. This book comes at a time when new technologies and approaches allow the implications of climate change and other direct human impacts on Antarctica to be viewed at a range of scales; across entire regions, whole ecosystems and down to the level of species and variation within t

      Trade Review

      “Overall, this book provides a comprehensive overview of Antarctic ecosystems and the open access approach to publication means this volume serves as an easy entre to that literature – many ecologists will benefit from this compilation.” (Austral Ecology, 1 October 2013)

      “As an institutional library purchase, I would recommend this book.” (Frontiers of biogeography, 5 January 2013

      “This timely summary of the state of Antarctic ecological science provides a springboard for an exciting future of Antarctic research.” (The Quarterly Review of Biology, 1 June 2013)

      “Overall, I appreciated the book and found it to be a very good synthesis especially of the marine information.” (Biodiversity and Conservation, 1 October 2012)

      “The first of these two books is a good scientific treatise on how snow and ice communities work at the moment, while the second concentrates more on marine environments and their likely future. Both are good and should be in the library.” (British Ecological Society Bulletin, 1 August 2012)

      “This book is a must for senior undergraduates, graduate students, and scientists interested in Antarctic ecosystems. Summing Up: Highly recommended. Upper-division undergraduates through professionals.” (Choice, 1 September 2012)



      Table of Contents

      Contributors, xi

      INTRODUCTION: ANTARCTIC ECOLOGY IN A CHANGING WORLD, 1
      Andrew Clarke, Nadine M. Johnston, Eugene J. Murphy and Alex D. Rogers

      Introduction, 1

      Climate change, 2

      The historical context, 3

      The importance of scale, 3

      Fisheries and conservation, 4

      Concluding remarks, 6

      References, 6

      PART 1 TERRESTRIAL AND FRESHWATER HABITATS, 11

      1 SPATIAL AND TEMPORAL VARIABILITY IN TERRESTRIAL ANTARCTIC BIODIVERSITY, 13
      Steven L. Chown and Peter Convey

      1.1 Introduction, 13

      1.2 Variation across space, 16

      1.2.1 Individual and population levels, 16

      1.2.2 Species level, 18

      1.2.3 Assemblage and ecosystem levels, 20

      1.3 Variation through time, 25

      1.3.1 Individual level, 26

      1.3.2 Population level, 27

      1.3.3 Species level, 29

      1.3.4 Assemblage and ecosystem levels, 29

      1.4 Conclusions and implications, 30

      Acknowledgments, 31

      References, 31

      2 GLOBAL CHANGE IN A LOW DIVERSITY TERRESTRIAL ECOSYSTEM: THE MCMURDO DRY VALLEYS, 44
      Diana H. Wall

      2.1 Introduction, 44

      2.2 The McMurdo dry valley region, 46

      2.3 Above–belowground interactions, 46

      2.4 The functioning of low diversity systems, 50

      2.5 Effects of global changes on coupled above–belowground subsystems, 51

      2.6 Temperature change: warming, 52

      2.7 Temperature change: cooling, 54

      2.8 Direct human influence: trampling, 54

      2.9 UV Radiation, 55

      2.10 Concluding remarks, 56

      Acknowledgements, 56

      References, 56

      3 ANTARCTIC LAKES AS MODELS FOR THE STUDY OF MICROBIAL BIODIVERSITY, BIOGEOGRAPHY AND EVOLUTION, 63
      David A. Pearce and Johanna Laybourn-Parry

      3.1 The variety of antarctic lake types, 63

      3.2 The physical and chemical lake environment, 66

      3.3 The microbial diversity of antarctic lakes, 66

      3.3.1 Methods for exploring Antarctic lake biodiversity, 67

      3.3.2 Microbial groups, 69

      3.3.3 Protists, 70

      3.3.4 Crustacea, 72

      3.4 Biogeography, 74

      3.4.1 Spatial variation and the global ubiquity hypothesis, 74

      3.4.2 Temporal variation and palaeolimnology, 75

      3.5 Evolution, 76

      3.5.1 Prokaryote physiology, 76

      3.5.2 Eukaryote physiology, 77

      3.6 Future perspectives, 78

      3.7 Acknowledgement, 78

      References, 78

      PART 2 MARINE HABITATS AND REGIONS, 91

      4 THE IMPACT OF REGIONAL CLIMATE CHANGE ON THE MARINE ECOSYSTEM OF THE WESTERN ANTARCTIC PENINSULA, 93
      Andrew Clarke, David K. A. Barnes, Thomas J. Bracegirdle, Hugh W. Ducklow, John C. King, Michael P. Meredith, Eugene J. Murphy and Lloyd S. Peck

      4.1 Introduction, 93

      4.1.1 The oceanographic setting, 96

      4.1.2 The historical context, 97

      4.2 Predicted environmental changes along the western antarctic peninsula, 98

      4.3 Environmental variability and ecological response, 100

      4.3.1 Biotic responses to climate change: some general points, 102

      4.4 Responses of individual marine species to climate change, 102

      4.4.1 Acclimation and evolutionary responses to environmental change in antarctic marine organisms, 104

      4.5 Community level responses to climate change, 106

      4.6 Ecosystem level responses to climate change, 107

      4.7 What biological changes have been observed to date?, 109

      4.8 Concluding remarks, 110

      Acknowledgements, 110

      References, 111

      5 THE MARINE SYSTEM OF THE WESTERN ANTARCTIC PENINSULA, 121
      Hugh Ducklow, Andrew Clarke, Rebecca Dickhut, Scott C. Doney, Heidi Geisz, Kuan Huang, Douglas G. Martinson, Michael P. Meredith, Holly V. Moeller, Martin Montes-Hugo, Oscar Schofield, Sharon E. Stammerjohn, Debbie Steinberg and William Fraser

      5.1 Introduction, 121

      5.2 Climate and ice, 123

      5.2.1 Surface air temperature, 123

      5.2.2 Sea ice, 123

      5.2.3 Climate co-variability, 125

      5.3 Physical oceanography, 127

      5.4 Nutrients and carbon, 130

      5.4.1 Nutrients and UCDW intrusions, 130

      5.4.2 Carbon cycle, 131

      5.4.3 Dissolved organic carbon, 132

      5.4.4 Sedimentation and export, 133

      5.5 Phytoplankton dynamics, 134

      5.5.1 Seasonal scale dynamics, 134

      5.5.2 Role of light, 134

      5.5.3 Role of nutrients, 136

      5.5.4 Annual variability in phytoplankton, 137

      5.6 Microbial ecology, 138

      5.7 Zooplankton, 140

      5.7.1 Community composition and distribution, 140

      5.7.2 Long-term trends and climate connections, 142

      5.7.3 Grazing and biogeochemical cycling, 142

      5.8 Penguins, 143

      5.8.1 Contaminants in penguins, 145

      5.9 Marine mammals, 146

      5.10 Synthesis: food webs of the wap, 147

      5.11 Conclusions, 148

      Acknowledgements, 149

      References, 149

      6 SPATIAL AND TEMPORAL OPERATION OF THE SCOTIA SEA ECOSYSTEM, 160
      E.J. Murphy, J.L. Watkins, P.N. Trathan, K. Reid, M.P. Meredith, S.L. Hill, S.E. Thorpe, N.M. Johnston, A. Clarke, G.A. Tarling, M.A. Collins, J. Forcada, A. Atkinson, P. Ward, I.J. Staniland, D.W. Pond, R.A. Cavanagh, R.S. Shreeve, R.E. Korb, M.J. Whitehouse, P.G. Rodhouse, P. Enderlein, A.G. Hirst, A.R. Martin, D.R. Briggs, N.J. Cunningham and A.H. Fleming

      6.1 Introduction, 160

      6.2 Oceanography and sea ice, 163

      6.2.1 Upper-ocean circulation and characteristics in the Scotia Sea, 163

      6.2.2 Physical variability and long-term change, 167

      6.3 Nutrient and plankton dynamics, 168

      6.4 Krill in the scotia sea food web, 171

      6.4.1 Krill distribution in the Scotia Sea, 171

      6.4.2 Krill growth and age in the Scotia Sea, 173

      6.4.3 Krill reproduction and recruitment in the Scotia Sea, 174

      6.4.4 Krill – habitat interactions in the Scotia Sea, 177

      6.4.5 Krill population variability and change in the Scotia Sea, 180

      6.4.6 Krill in the Scotia Sea food web, 183

      6.5 Food web operation, 184

      6.5.1 Trophic links, 184

      6.5.2 Spatial operation of the food web, 189

      6.6 Ecosystem variability and long-term change, 192

      6.7 Concluding comments, 195

      Summary, 196

      Acknowledgements, 197

      References, 197

      7 THE ROSS SEA CONTINENTAL SHELF: REGIONAL BIOGEOCHEMICAL CYCLES, TROPHIC INTERACTIONS, AND POTENTIAL FUTURE CHANGES, 213
      Walker O. Smith, Jr., David G. Ainley, Riccardo Cattaneo-Vietti and Eileen E. Hofmann

      7.1 Introduction, 213

      7.2 Physical setting, 214

      7.3 Biological setting, 219

      7.3.1 Lower trophic levels, 219

      7.3.2 Mid-trophic levels, 225

      7.3.3 Fishes and mobile predators, 226

      7.3.4 Upper trophic levels, 227

      7.3.5 Benthos, 229

      7.4 Food web and biotic interactions, 230

      7.5 Conclusions, 232

      7.5.1 Uniqueness of the Ross Sea, 232

      7.5.2 Potential impacts of climate change, 233

      7.5.3 Conservation and the role of commercial fishing activity in the Ross Sea, 234

      7.5.4 Research needs and future directions, 235

      Acknowledgements, 235

      References, 235

      8 PELAGIC ECOSYSTEMS IN THE WATERS OFF EAST ANTARCTICA (30 E–150 E), 243
      Stephen Nicol and Ben Raymond

      8.1 Introduction, 243

      8.2 The region, 245

      8.2.1 The east (80 E–150 E), 245

      8.2.2 The west (30 E–80 E), 247

      8.3 Ecosystem change off east antarctica, 251

      Summary, 251

      References, 252

      9 THE DYNAMIC MOSAIC, 255
      David K.A. Barnes and Kathleen E. Conlan

      9.1 Introduction, 255

      9.2 Historical and geographic perspectives, 256

      9.3 Disturbance, 257

      9.3.1 Ice effects, 258

      9.3.2 Asteroid impacts, 260

      9.3.3 Sediment instability and hypoxia, 261

      9.3.4 Wind and wave action, 261

      9.3.5 Pollution, 262

      9.3.6 UV irradiation, 263

      9.3.7 Volcanic eruptions, 263

      9.3.8 Trawling, 264

      9.3.9 Non-indigenous species (NIS), 264

      9.3.10 Freshwater, 265

      9.3.11 Temperature stress, 265

      9.3.12 Biological agents of physical disturbance, 266

      9.4 Colonisaton of antarctic sea-beds, 266

      9.4.1 Larval abundance, 266

      9.4.2 Hard substrata, 266

      9.4.3 Soft sediments, 269

      9.5 Implications of climate change, 276

      9.6 Conclusion, 279

      Acknowledgements, 280

      References, 281

      10 SOUTHERN OCEAN DEEP BENTHIC BIODIVERSITY, 291
      A. Brandt, C. De Broyer, B. Ebbe, K.E. Ellingsen, A.J. Gooday, D. Janussen, S. Kaiser, K. Linse, M. Schueller, M.R.A. Thomson, P.A. Tyler and A. Vanreusel

      10.1 Introduction, 291

      10.2 History of antarctic biodiversity work, 293

      10.3 Geological history and evolution of the antarctic, 294

      10.3.1 Indian Ocean, 294

      10.3.2 South Atlantic, 294

      10.3.3 Weddell Sea, 295

      10.3.4 Drake Passage and Scotia Sea, 296

      10.4 Benthic composition and diversity of meio-, macro- and megabenthos, 296

      10.4.1 Meiofauna, 297

      10.4.2 Macrofaunal composition and diversity, 299

      10.4.3 Megafaunal composition and diversity, 304

      10.5 Phylogenetic relationships of selected taxa, 308

      10.5.1 Foraminifera, 308

      10.5.2 Isopoda, 308

      10.5.3 Tanaidacea, 309

      10.5.4 Bivalvia, 310

      10.5.5 Polychaeta, 310

      10.5.6 Cephalopoda, 310

      10.6 Biogeography and endemism, 311

      10.6.1 Porifera, 311

      10.6.2 Foraminifera, 311

      10.6.3 Metazoan meiofauna, 311

      10.6.4 Peracarida, 312

      10.6.5 Mollusca, 312

      10.6.6 Echinodermata, 313

      10.6.7 Brachiopoda, 313

      10.6.8 Polychaeta, 313

      10.6.9 Bryozoa, 313

      10.7 Relationship of selected faunal assemblages to environmental variables, 313

      10.7.1 Large-scale patterns with depth, 313

      10.7.2 Patterns influenced by other environmental or physical factors, 317

      10.7.3 Isopoda, 318

      10.8 Similarities and differences between antarctic and other deep-sea systems, 318

      10.8.1 The environment, 318

      10.8.2 A direct comparison between the deep sea of the SO and the World Ocean, 319

      10.8.3 Dispersal and recruitment between the SO and the rest of the world, 320

      10.8.4 The special case of chemosynthetically-driven deep-sea systems, 320

      10.9 Conclusions, 321

      Acknowledgements, 321

      References, 323

      11 ENVIRONMENTAL FORCING AND SOUTHERN OCEAN MARINE PREDATOR POPULATIONS, 335
      Phil N. Trathan, Jaume Forcada and Eugene J. Murphy

      11.1 Climate change: recent, rapid, regional warming, 335

      11.2 Using oscillatory climate signals to predict future change in biological communities, 337

      11.3 Potential for regional impacts on the biosphere, 338

      11.4 Confounding isues in identifying a biological signal, 339

      11.5 Regional ecosystem responses as a consequence of variation in regional food webs, 340

      11.6 Where biological signals will be most apparent, 340

      11.7 The southwest atlantic, 341

      11.8 The indian ocean, 344

      11.9 The pacific ocean, 345

      11.10 Similarities between the atlantic, indian and pacific oceans, 346

      11.11 What ENSO can tell us, 347

      11.12 Future scenarios, 349

      References, 349

      PART 3 MOLECULAR ADAPTATIONS AND EVOLUTION, 355

      12 MOLECULAR ECOPHYSIOLOGY OF ANTARCTIC NOTOTHENIOID FISHES, 357
      C.-H. Christina Cheng and H. William Detrich III

      12.1 Introduction, 357

      12.2 Surviving the big chill – notothenioid freezing avoidance by antifreeze proteins, 358

      12.2.1 Freezing challenge in frigid Antarctic marine environment, 358

      12.2.2 Historical paradigm of teleost freezing avoidance, 360

      12.2.3 Paradigm shift I: the ‘larval paradox’, 360

      12.2.4 Paradigm shift II: liver is not the source of blood AFGP in notothenioids, 362

      12.2.5 Gut versus blood – importance of intestinal freeze avoidance, 363

      12.2.6 Non-hepatic source of plasma AFGP, 364

      12.2.7 Alterations in environments and dynamic evolutionary change in notothenioid AFGP gene families, 364

      12.2.8 Summary comments – antifreeze protein gain in Antarctic notothenioid fish, 367

      12.3 Haemoprotein loss and cardiovascular adaptation in icefishes – dr. no to the rescue?, 367

      12.3.1 Vertebrates without haemoglobins – you must be kidding!, 367

      12.3.2 Haemoprotein loss in icefishes: an evolutionary perspective, 368

      12.3.3 Cellular correlates of haemoprotein loss, 370

      12.3.4 The icefish cardiovascular system, 371

      12.3.5 Compensatory adjustment of the icefish cardiovascular system in a regime of reduced interspecific competition? Enter Dr. NO, 371

      12.3.6 Haemoproteins, NO metabolism, and icefish evolution, 372

      12.3.7 Icefishes and erythropoietic gene discovery, 372

      12.3.8 Summary comments: haemoprotein loss in Antarctic icefishes, 374

      12.4 Concluding remarks, 374

      Acknowledgements, 374

      Dedication, 374

      References, 374

      13 MECHANISMS DEFINING THERMAL LIMITS AND ADAPTATION IN MARINE ECTOTHERMS: AN INTEGRATIVE VIEW, 379
      Hans O. P€ortner, Lloyd S. Peck and George N. Somero

      13.1 Introduction: climate-dependent evolution of antarctic fauna, 379

      13.2 Phenomena of thermal specialization and limitation, 382

      13.2.1 Molecular and membrane aspects, 383

      13.2.2 Genomic aspects: gene expression and loss of genetic information, 390

      13.2.3 From molecular to systemic aspects: thermal limitation, 393

      13.2.4 From molecular to systemic aspects: thermal adaptation of performance capacity, 397

      13.2.5 Ecological implications, 399

      13.2.6 Integration of phenomena: concepts, results and perspectives, 405

      Acknowledgements, 409

      References, 409

      14 EVOLUTION AND BIODIVERSITY OF ANTARCTIC ORGANISMS, 417
      Alex D. Rogers

      14.1 Introduction, 417

      14.2 The antarctic biota, 418

      14.3 The break-up of gondwana and the evolution of the southern hemisphere biota, 420

      14.3.1 Vicariance versus dispersal, 420

      14.3.2 Dispersal mechanisms, 421

      14.4 The evolution and biodiversity of the terrestrial sub-antarctic and antarctic biota, 423

      14.4.1 Plants, 423

      14.4.2 Animals, 427

      14.5 The marine environment, 432

      14.5.1 Biogeography and macroevolution, 432

      14.5.2 Notothenioid fish, 432

      14.5.3 Birds, 435

      14.5.4 Marine invertebrates, 436

      14.5.5 The molecular ecology and phylogeography of the marine biota, 437

      14.5.6 Patterns of genetic variation in marine species, 448

      14.6 Antarctica: a climatic crucible of evolution, 450

      14.7 The historical constraints on adaptation to present climate change, 453

      14.8 Future directions for research, 453

      References, 454

      PART 4 CONSERVATION AND MANAGEMENT ASPECTS, 469

      15 BIOGEOGRAPHY AND REGIONAL CLASSIFICATIONS OF ANTARCTICA, 471
      P. Convey, D.K.A. Barnes, H.J. Griffiths, S.M. Grant, K. Linse and D.N. Thomas

      15.1 Introduction, 471

      15.2 Historical background, 474

      15.2.1 Physical regions in the marine environment, 474

      15.2.2 Smaller-scale regionalization within the Antarctic marine environment, 474

      15.2.3 Physical regions in the littoral environment, 475

      15.2.4 Physical regions in the terrestrial environment, 475

      15.3 Data availability, 476

      15.4 Different realms in the marine and terrestrial environments, 477

      15.4.1 Pelagic realm, 477

      15.4.2 Sea ice, 478

      15.4.3 Benthic realm, 479

      15.4.4 The terrestrial environment, 479

      15.4.5 Biogeographical patterns in the terrestrial environment, 480

      15.4.6 Biogeographic patterns in the marine environment, 481

      15.5 Overview, 485

      Acknowledgements, 486

      References, 486

      16 CONSERVATION AND MANAGEMENT OF ANTARCTIC ECOSYSTEMS, 492
      Susie M. Grant, Pete Convey, Kevin A. Hughes, Richard A. Phillips and Phil N. Trathan

      16.1 Introduction, 492

      16.2 Legal frameworks for conservation and management, 495

      16.2.1 Early regulation of marine living resource harvesting, 495

      16.2.2 The Antarctic Treaty System, 497

      16.2.3 Other (non-ATS) agreements and tools relevant to conservation and management, 500

      16.3 Conservation and management measures, 502

      16.3.1 Pollution and local disturbance, 502

      16.3.2 Biosecurity and non-native species, 505

      16.3.3 Conservation and management of marine living resources, 505

      16.3.4 Conservation of other individual species, 507

      16.3.5 Protected areas, 509

      16.4 Conservation science and monitoring, 512

      16.5 Future challenges, 515

      16.6 Conclusions, 520

      Acknowledgements, 521

      References, 521

      Index, 526

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