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. PEURortner, 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
