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The Arctic is often portrayed as being isolated, but the reality is that the connectivity with the rest of the planet is huge, be it through weather patterns, global ocean circulation, and large-scale migration patterns to name but a few. There is a huge amount of public interest in the 'changing Arctic', especially in terms of the rapid changes taking place in ecosystems and exploitation of resources. There can be no doubt that the Arctic is at the forefront of the international environmental science agenda, both from a scientific aspect, and also from a policy/environmental management perspective.
This book aims to stimulate a wide audience to think about the Arctic by highlighting the remarkable breadth of what it means to study its ecology. Arctic Ecology seeks to systematically introduce the diverse array of ecologies within the Arctic region. As the Arctic rapidly changes, understanding the fundamental ecology underpinning the Arctic is paramount to understanding the consequences of what such change will inevitably bring about.
Arctic Ecology is designed to provide graduate students of environmental science, ecology and climate change with a source where Arctic ecology is addressed specifically, with issues due to climate change clearly discussed. It will also be of use to policy-makers, researchers and international agencies who are focusing on ecological issues and effects of global climate change in the Arctic.
About the Editor
David N. Thomas is Professor of Arctic Ecosystem Research in the Faculty of Biological and Environmental Sciences, University of Helsinki. Previously he spent 24 years in the School of Ocean Sciences, Bangor University, Wales. He studies marine systems, with a particular emphasis on sea ice and land-coast interactions in the Arctic and Southern Oceans as well as the Baltic Sea. He also edited a related book: Sea Ice, 3rd Edition (2017), which is also published by Wiley-Blackwell.
DAVID N. THOMAS is Professor of Arctic Ecosystem Research in the Faculty of Biological and Environmental Sciences, University of Helsinki. Previously he spent 24 years in the School of Ocean Sciences, Bangor University, Wales, where he is now an Honorary Professor. He studies marine systems, with a particular emphasis on sea ice and land-coast interactions in the Arctic and Southern Oceans as well as the Baltic Sea. He also edited a related book: Sea Ice, 3rd Edition (2017), which is also published by Wiley-Blackwell.
Preface xiii
List of Contributors xv
1 What Is the Arctic? 1Kjell Danell
1.1 Setting the Scene 1
1.2 In Which Ways Is the Arctic Different? 2
1.3 How Was the Arctic Discovered? 3
1.4 How Large Is the Arctic? 4
1.5 What Is in the Arctic? 4
1.5.1 Arctic Haze and Ice Fog 4
1.5.2 Aurora Borealis 5
1.6 Climate and Weather 5
1.7 Ice and Snow 6
1.8 Permafrost, Polygons, Pingos, and Palsas 6
1.9 Animals, Plants, and Fungi 7
1.10 Arctic Ecosystems 9
1.10.1 Terrestrial Ecosystems 10
1.10.2 Freshwater Ecosystems 12
1.10.3 Marine Ecosystems 14
1.10.4 Humans 15
1.11 Which Natural Resources and Ecosystem Services does the Arctic Offer? 17
1.12 Biotic Changes in the Arctic 19
References 22
2 Arctic Ecology - A Paleoenvironmental Perspective 23Michael Pisaric and John P. Smol
2.1 Introduction 23
2.2 The Distant Past 25
2.2.1 Bones, DNA, and Megafauna 26
2.2.2 Beringian Biota 27
2.2.3 Ancient DNA 29
2.3 Rings and Things: Examining Paleoenvironmental and Paleoclimatic Change Using Dendrochronology 30
2.3.1 Dendrochronology in Action: Examples from the Field 32
2.4 Lake Sediments: Continuous Archives of Environmental Change 33
2.5 Paleolimnology and Arctic Climate Change 37
2.5.1 Subfossil Pollen, Stomata, and Macrofossils for Tracking Vegetation Change 38
2.5.2 Charcoal and Past Wildfires 39
2.5.3 Using Past Assemblage Changes in Lake Biota to Reconstruct Past Climatic Trends 39
2.5.4 Using Paleolimnology to Study the Source and Fate of Contaminants 43
2.5.5 Linking Paleolimnology and Archeology: Tracking the Limnological Effects of Early Peoples in the Arctic 48
2.6 Concluding Remarks 49
References 50
3 Climate Change in the Arctic 57Edward Hanna, Joseph E. Nolan, James E. Overland, and Richard J. Hall
3.1 Introduction to Arctic Climates - Datasets Available for Analyzing Climate Change 57
3.2 Atmospheric Aspects of Arctic Climate Change: Arctic Amplification and Global Warming, Changes in Air Temperature and Precipitation, and Changes in Atmospheric Circulation 59
3.3 Oceanic Aspects of Arctic Climate Change, Including Surface and Deep Ocean Circulation Changes 61
3.4 Climate Change Impacts on Arctic Sea Ice and Greenland Ice Sheet - The Unprecedented Recent Decline in Late Summer Sea-Ice Cover and Record Greenland Ice Sheet Surface Melt and Mass Loss 65
3.5 Feedbacks in the Arctic Climate System and Global Impacts - the Ice/Albedo Feedback and Ice Insulation Feedbacks - the "Warm Arctic, Cold Continents" Hypothesis 71
3.6 Concluding Remarks 73
References 75
4 Arctic Permafrost and Ecosystem Functioning 81Torben R. Christensen
4.1 Permafrost and Ecosystems in the Arctic 81
4.2 Permafrost Shapes the Landscape 83
4.2.1 Permafrost Specific Landforms and Their Importance for Ecosystems 83
4.2.2 Permafrost Specific Landforms and Effects of a Changing Climate 86
4.3 The Biology of Permafrost 87
4.3.1 Microbes 87
4.3.2 Vegetation 88
4.4 Ecosystem Function - Carbon Cycling in Permafrost Environments 91
4.4.1 General Carbon Cycling 91
4.4.2 Methane Emissions 92
4.5 Concluding Remarks 94
References 95
5 Arctic Tundra 103John Hobbie, Gaius Shaver, Toke Thomas Høye, and Joseph Bowden
5.1 Distribution and Description of Arctic Tundra 103
5.2 Tundra Organisms: A Typical Food Web 105
5.3 Flora and Fauna: Diversity and Communities 106
5.4 Primary Production and Organic Matter Stocks in the Low and High Arctic 110
5.5 Primary Production and Organic Matter Stocks 112
5.6 Adaptations to the Arctic Tundra 112
5.6.1 Plant Adaptations 112
5.6.2 Microbial Activity and Soil Carbon and Nitrogen 115
5.6.3 Invertebrates: Diversity, Freeze-Tolerance, and Freeze-Avoidance 116
5.6.4 Vertebrates of the Tundra: Wintertime Survival Strategies 117
5.7 Reproductive Strategies 118
5.8 Populations and Communities of the Tundra 120
5.8.1 Diversity and Interactions: The Case of Beringia 120
5.8.2 Development of Arctic Tundra Food Webs: Complexity, Insects 121
5.8.3 Belowground Arctic Food Web: Bacteria, Mycorrhizal Fungi, Nitrogen, and Carbon Cycling 122
5.9 Tundra Ecosystem Analysis 123
5.9.1 Why Nutrient Limitation? 123
5.9.2 Nitrogen Budget: Pools of Nitrogen, Rates of Transport, and Transformations 124
5.9.3 Carbon Budget: Pools, Gross Photosynthesis and Respiration, Accumulation and Feedbacks 126
5.9.4 Insights from Manipulation Experiments: Control of Net Primary Production and Herbivory by Nutrients, Light, and Heat 126
5.10 Expected Future Changes and Responses in Arctic Tundra 128
5.10.1 Effects of Increased Shrubs 128
5.10.2 Pest Outbreaks, Changes in Phenology and Species Interactions 129
References 130
6 Ecology of Arctic Glaciers 133Alexandre M. Anesio and Johanna Laybourn-Parry
6.1 Introduction 133
6.2 The Biodiversity and Food Webs of Glacial Habitats 136
6.2.1 Ice Shelves 136
6.2.2 Supraglacial or Cryolakes 138
6.2.3 Cryoconite Material, Cryoconite Holes, and Wet Ice Surfaces 139
6.2.4 Snow on the Surface of Ice 141
6.2.5 Life Within Ice 141
6.2.6 Life in Subglacial Environments 143
6.3 Quantification of Microbial Processes in Glaciers and Export of Material to Adjacent Ecosystems 144
6.3.1 Supraglacial Habitats: Ice Shelf Lakes, Ponds, Cryoconites 144
6.3.2 Subglacial Habitats 149
6.4 Anthropogenic Impacts 151
References 152
7 Ecology of Arctic Lakes and Ponds 159Erik Jeppesen, Kirsten S. Christoffersen, Milla Rautio, and Torben L. Lauridsen
7.1 Introduction 159
7.2 Physical and Chemical Characteristics of Arctic Lakes and Ponds 160
7.3 Biological Communities and Production 162
7.3.1 Phytoplankton and Phytobenthos 162
7.3.2 Macrophytes 164
7.3.3 Microbial Loop 165
7.3.4 Zooplankton and Zoobenthos 165
7.3.5 Fish 168
7.3.6 Food Webs 169
7.4 Global Climate Change and Arctic Lakes 171
References 174
8 Ecology of Arctic Streams and Rivers 181Alexander D. Huryn
8.1 Introduction 181
8.1.1 What Is an Arctic River? 181
8.2 A Primer on Stream Ecology: General and Arctic Perspectives 183
8.2.1 The Longitudinal Dimension 184
8.2.2 The Vertical Dimension 202
8.2.3 The Lateral Dimension 207
8.3 Concluding Remarks 209
References 211
9 Ecology of Arctic Pelagic Communities 219Malin Daase, Jørgen Berge, Janne E. Søreide, and Stig Falk-Petersen
9.1 Introduction 219
9.2 The Arctic Marine Highways: The Transpolar Drift and the Interconnected Current Systems 220
9.3 Members and Key Players of Arctic Pelagic Communities 224
9.3.1 At the Base - Primary Producers and Microbial Communities 224
9.3.2 In the Middle - Resident Consumers and Life Strategies of Arctic Zooplankton 229
9.3.3 At the Top - Pelagic Predators 239
9.4 A Lipid-Driven Food Chain 241
9.5 Effects of Climate Change 242
9.5.1 Timing 243
9.5.2 Changes in Species Distribution 243
References 246
10 Ecology of Arctic Sea Ice 261C. J. Mundy and Klaus M. Meiners
10.1 Introduction to Sea Ice 261
10.2 Types of Habitats 262
10.3 Food Webs and Carbon Flow 264
10.4 Physical Environment 268
10.5 Colonization of Sea Ice and Winter Survival 270
10.6 Adaptations to and Relationships with Environmental Conditions 272
10.6.1 Temperature and Salinity 272
10.6.2 Space and Permeability 275
10.6.3 Light 276
10.7 Climate Change and the Ice-Associated Ecosystem 278
References 279
11 Ecology of Arctic Shallow Subtidal and Intertidal Benthos 289Paul E. Renaud, Jan Marcin Weslawski, and Kathleen Conlan
11.1 Introduction 289
11.2 The Physical Environment 290
11.2.1 Temperature 290
11.2.2 Light 290
11.2.3 Waves 291
11.2.4 Ice Cover 291
11.2.5 Freshwater Discharge and Salinity 293
11.3 Biomes 294
11.3.1 Origins and Distribution of Sediments 294
11.3.2 Soft-Sediment Communities 295
11.3.3 Hard Substrate 297
11.3.4 Vegetated Substrate 300
11.4 Disturbance Regimes and Succession 302
11.4.1 Ice Scour 302
11.4.2 Strudel Scour 304
11.4.3 Natural Gas Seepage and Petroleum Extraction 305
11.4.4 Large-Mammal Feeding Pits 305
11.4.5 Recolonization of Arctic Benthos 306
11.4.6 Human Impacts 307
11.5 Trophic Interactions 308
11.5.1 Feeding Strategies in Arctic Shallow Benthos 308
11.5.2 Food Sources for Benthic Fauna 309
11.5.3 Benthos as a Food for Top Predators in the Arctic 310
11.5.4 Carbon and Nutrient Cycling 314
11.6 Reproduction in Coastal Benthos 315
11.7 Effects of Global Climate Change on Shallow Arctic Benthos 316
11.7.1 Reduced Ice Cover 319
11.7.2 Documented Effects on Natural Systems in the Intertidal Zone 319
11.7.3 Coastal Erosion 319
References 320
12 Ecology of Arctic Shelf and Deep Ocean Benthos 325Monika Kedra and Jacqueline M. Grebmeier
12.1 Introduction 325
12.2 The Physical Environment 326
12.2.1 Light 327
12.2.2 Temperature 327
12.2.3 Sea Ice 328
12.2.4 Sediment Characteristics 328
12.3 Biodiversity, Community Structure, and Functioning of Shelf and Deep Sea Benthos 330
12.3.1 Benthic Definitions 330
12.3.2 Brief Overview of Major Taxa in Benthic Communities 331
12.3.3 Biodiversity 335
12.3.4 Functional Diversity 337
12.3.5 Arctic Commercial Benthic Species 338
12.4 Productivity and Food Webs of Shelf and Deep Sea Benthos 339
12.4.1 Primary Production and Food Sources 339
12.4.2 Pelagic-Benthic Coupling 341
12.4.3 Benthic Community Structure and Food Webs 343
12.4.4 Benthic Communities as a Food Source for Benthic-Feeding Upper Trophic Levels 346
12.5 Impact of Global Climate Change on Shelf and Deep Sea Benthic Communities 348
References 349
13 Fat, Furry, Flexible, and Functionally Important: Characteristics of Mammals Living in the Arctic 357Niels M. Schmidt, Olivier Gilg, Jon Aars, and Rolf A. Ims
13.1 Introduction 357
13.2 The Mammal Assemblage in the Arctic Today 358
13.2.1 Terrestrial Mammals 358
13.2.2 Herbivores 358
13.2.3 Predators 359
13.2.4 Marine Mammals 360
13.3 Arctic Mammals and Adaptations to Life in the Arctic 366
13.3.1 Fur, Fat and Extremities 367
13.3.2 Behavioral Adaptations to Life in the Arctic 368
13.4 The Role of Mammals in Arctic Ecosystems 369
13.4.1 Removal of Plant Material by Terrestrial Herbivores 371
13.4.2 Transport of Nutrients and Seeds by Arctic Mammals 372
13.4.3 Mammal Predator-Prey Dynamics 374
13.5 The Future for Arctic Mammals in a Changing Climate 374
13.6 Concluding Remarks 377
References 377
14 Ecology of Arctic Birds 385Anthony D. Fox
14.1 Introduction: The Bird Species and Their Feeding Ecology 385
14.2 Traveling to Breed 386
14.3 Long Distance Migrations 388
14.4 Reproduction 389
14.5 Survival 394
14.6 Population Change 396
14.7 Climate Change 397
14.8 Endangered Species 399
14.9 Concluding Remarks 402
References 403
15 Arctic Ecology, Indigenous Peoples and Environmental Governance 409Mark Nuttall
15.1 Introduction 409
15.2 The Impacts of Social and Environmental Change 410
15.3 Traditional Ecological Knowledge and Wildlife Management 413
15.4 Arctic Ecology and Community-Based Monitoring 415
15.5 Indigenous Peoples and Environmental Policy: The Case of the Inuit Circumpolar Council 418
15.6 Concluding Remarks 419
References 420
Index 423
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