Arctic Ecology

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, 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 xiiiList of Contributors xv1 What Is the Arctic? 1Kjell Danell1.1 Setting the Scene 11.2 In Which Ways Is the Arctic Different? 21.3 How Was the Arctic Discovered? 31.4 How Large Is the Arctic? 41.5 What Is in the Arctic? 41.5.1 Arctic Haze and Ice Fog 41.5.2 Aurora Borealis 51.6 Climate and Weather 51.7 Ice and Snow 61.8 Permafrost, Polygons, Pingos, and Palsas 61.9 Animals, Plants, and Fungi 71.10 Arctic Ecosystems 91.10.1 Terrestrial Ecosystems 101.10.2 Freshwater Ecosystems 121.10.3 Marine Ecosystems 141.10.4 Humans 151.11 Which Natural Resources and Ecosystem Services does the Arctic Offer? 171.12 Biotic Changes in the Arctic 19References 222 Arctic Ecology – A Paleoenvironmental Perspective 23Michael Pisaric and John P. Smol2.1 Introduction 232.2 The Distant Past 252.2.1 Bones, DNA, and Megafauna 262.2.2 Beringian Biota 272.2.3 Ancient DNA 292.3 Rings and Things: Examining Paleoenvironmental and Paleoclimatic Change Using Dendrochronology 302.3.1 Dendrochronology in Action: Examples from the Field 322.4 Lake Sediments: Continuous Archives of Environmental Change 332.5 Paleolimnology and Arctic Climate Change 372.5.1 Subfossil Pollen, Stomata, and Macrofossils for Tracking Vegetation Change 382.5.2 Charcoal and Past Wildfires 392.5.3 Using Past Assemblage Changes in Lake Biota to Reconstruct Past Climatic Trends 392.5.4 Using Paleolimnology to Study the Source and Fate of Contaminants 432.5.5 Linking Paleolimnology and Archeology: Tracking the Limnological Effects of Early Peoples in the Arctic 482.6 Concluding Remarks 49References 503 Climate Change in the Arctic 57Edward Hanna, Joseph E. Nolan, James E. Overland, and Richard J. Hall3.1 Introduction to Arctic Climates – Datasets Available for Analyzing Climate Change 573.2 Atmospheric Aspects of Arctic Climate Change: Arctic Amplification and Global Warming, Changes in Air Temperature and Precipitation, and Changes in Atmospheric Circulation 593.3 Oceanic Aspects of Arctic Climate Change, Including Surface and Deep Ocean Circulation Changes 613.4 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 653.5 Feedbacks in the Arctic Climate System and Global Impacts – the Ice/Albedo Feedback and Ice Insulation Feedbacks – the “Warm Arctic, Cold Continents” Hypothesis 713.6 Concluding Remarks 73References 754 Arctic Permafrost and Ecosystem Functioning 81Torben R. Christensen4.1 Permafrost and Ecosystems in the Arctic 814.2 Permafrost Shapes the Landscape 834.2.1 Permafrost Specific Landforms and Their Importance for Ecosystems 834.2.2 Permafrost Specific Landforms and Effects of a Changing Climate 864.3 The Biology of Permafrost 874.3.1 Microbes 874.3.2 Vegetation 884.4 Ecosystem Function – Carbon Cycling in Permafrost Environments 914.4.1 General Carbon Cycling 914.4.2 Methane Emissions 924.5 Concluding Remarks 94References 955 Arctic Tundra 103John Hobbie, Gaius Shaver, Toke Thomas Høye, and Joseph Bowden5.1 Distribution and Description of Arctic Tundra 1035.2 Tundra Organisms: A Typical Food Web 1055.3 Flora and Fauna: Diversity and Communities 1065.4 Primary Production and Organic Matter Stocks in the Low and High Arctic 1105.5 Primary Production and Organic Matter Stocks 1125.6 Adaptations to the Arctic Tundra 1125.6.1 Plant Adaptations 1125.6.2 Microbial Activity and Soil Carbon and Nitrogen 1155.6.3 Invertebrates: Diversity, Freeze-Tolerance, and Freeze-Avoidance 1165.6.4 Vertebrates of the Tundra: Wintertime Survival Strategies 1175.7 Reproductive Strategies 1185.8 Populations and Communities of the Tundra 1205.8.1 Diversity and Interactions: The Case of Beringia 1205.8.2 Development of Arctic Tundra Food Webs: Complexity, Insects 1215.8.3 Belowground Arctic Food Web: Bacteria, Mycorrhizal Fungi, Nitrogen, and Carbon Cycling 1225.9 Tundra Ecosystem Analysis 1235.9.1 Why Nutrient Limitation? 1235.9.2 Nitrogen Budget: Pools of Nitrogen, Rates of Transport, and Transformations 1245.9.3 Carbon Budget: Pools, Gross Photosynthesis and Respiration, Accumulation and Feedbacks 1265.9.4 Insights from Manipulation Experiments: Control of Net Primary Production and Herbivory by Nutrients, Light, and Heat 1265.10 Expected Future Changes and Responses in Arctic Tundra 1285.10.1 Effects of Increased Shrubs 1285.10.2 Pest Outbreaks, Changes in Phenology and Species Interactions 129References 1306 Ecology of Arctic Glaciers 133Alexandre M. Anesio and Johanna Laybourn-Parry6.1 Introduction 1336.2 The Biodiversity and Food Webs of Glacial Habitats 1366.2.1 Ice Shelves 1366.2.2 Supraglacial or Cryolakes 1386.2.3 Cryoconite Material, Cryoconite Holes, and Wet Ice Surfaces 1396.2.4 Snow on the Surface of Ice 1416.2.5 Life Within Ice 1416.2.6 Life in Subglacial Environments 1436.3 Quantification of Microbial Processes in Glaciers and Export of Material to Adjacent Ecosystems 1446.3.1 Supraglacial Habitats: Ice Shelf Lakes, Ponds, Cryoconites 1446.3.2 Subglacial Habitats 1496.4 Anthropogenic Impacts 151References 1527 Ecology of Arctic Lakes and Ponds 159Erik Jeppesen, Kirsten S. Christoffersen, Milla Rautio, and Torben L. Lauridsen7.1 Introduction 1597.2 Physical and Chemical Characteristics of Arctic Lakes and Ponds 1607.3 Biological Communities and Production 1627.3.1 Phytoplankton and Phytobenthos 1627.3.2 Macrophytes 1647.3.3 Microbial Loop 1657.3.4 Zooplankton and Zoobenthos 1657.3.5 Fish 1687.3.6 Food Webs 1697.4 Global Climate Change and Arctic Lakes 171References 1748 Ecology of Arctic Streams and Rivers 181Alexander D. Huryn8.1 Introduction 1818.1.1 What Is an Arctic River? 1818.2 A Primer on Stream Ecology: General and Arctic Perspectives 1838.2.1 The Longitudinal Dimension 1848.2.2 The Vertical Dimension 2028.2.3 The Lateral Dimension 2078.3 Concluding Remarks 209References 2119 Ecology of Arctic Pelagic Communities 219Malin Daase, Jørgen Berge, Janne E. Søreide, and Stig Falk-Petersen9.1 Introduction 2199.2 The Arctic Marine Highways: The Transpolar Drift and the Interconnected Current Systems 2209.3 Members and Key Players of Arctic Pelagic Communities 2249.3.1 At the Base – Primary Producers and Microbial Communities 2249.3.2 In the Middle – Resident Consumers and Life Strategies of Arctic Zooplankton 2299.3.3 At the Top – Pelagic Predators 2399.4 A Lipid-Driven Food Chain 2419.5 Effects of Climate Change 2429.5.1 Timing 2439.5.2 Changes in Species Distribution 243References 24610 Ecology of Arctic Sea Ice 261C. J. Mundy and Klaus M. Meiners10.1 Introduction to Sea Ice 26110.2 Types of Habitats 26210.3 Food Webs and Carbon Flow 26410.4 Physical Environment 26810.5 Colonization of Sea Ice and Winter Survival 27010.6 Adaptations to and Relationships with Environmental Conditions 27210.6.1 Temperature and Salinity 27210.6.2 Space and Permeability 27510.6.3 Light 27610.7 Climate Change and the Ice-Associated Ecosystem 278References 27911 Ecology of Arctic Shallow Subtidal and Intertidal Benthos 289Paul E. Renaud, Jan Marcin Węsławski, and Kathleen Conlan11.1 Introduction 28911.2 The Physical Environment 29011.2.1 Temperature 29011.2.2 Light 29011.2.3 Waves 29111.2.4 Ice Cover 29111.2.5 Freshwater Discharge and Salinity 29311.3 Biomes 29411.3.1 Origins and Distribution of Sediments 29411.3.2 Soft-Sediment Communities 29511.3.3 Hard Substrate 29711.3.4 Vegetated Substrate 30011.4 Disturbance Regimes and Succession 30211.4.1 Ice Scour 30211.4.2 Strudel Scour 30411.4.3 Natural Gas Seepage and Petroleum Extraction 30511.4.4 Large-Mammal Feeding Pits 30511.4.5 Recolonization of Arctic Benthos 30611.4.6 Human Impacts 30711.5 Trophic Interactions 30811.5.1 Feeding Strategies in Arctic Shallow Benthos 30811.5.2 Food Sources for Benthic Fauna 30911.5.3 Benthos as a Food for Top Predators in the Arctic 31011.5.4 Carbon and Nutrient Cycling 31411.6 Reproduction in Coastal Benthos 31511.7 Effects of Global Climate Change on Shallow Arctic Benthos 31611.7.1 Reduced Ice Cover 31911.7.2 Documented Effects on Natural Systems in the Intertidal Zone 31911.7.3 Coastal Erosion 319References 32012 Ecology of Arctic Shelf and Deep Ocean Benthos 325Monika Kędra and Jacqueline M. Grebmeier12.1 Introduction 32512.2 The Physical Environment 32612.2.1 Light 32712.2.2 Temperature 32712.2.3 Sea Ice 32812.2.4 Sediment Characteristics 32812.3 Biodiversity, Community Structure, and Functioning of Shelf and Deep Sea Benthos 33012.3.1 Benthic Definitions 33012.3.2 Brief Overview of Major Taxa in Benthic Communities 33112.3.3 Biodiversity 33512.3.4 Functional Diversity 33712.3.5 Arctic Commercial Benthic Species 33812.4 Productivity and Food Webs of Shelf and Deep Sea Benthos 33912.4.1 Primary Production and Food Sources 33912.4.2 Pelagic–Benthic Coupling 34112.4.3 Benthic Community Structure and Food Webs 34312.4.4 Benthic Communities as a Food Source for Benthic-Feeding Upper Trophic Levels 34612.5 Impact of Global Climate Change on Shelf and Deep Sea Benthic Communities 348References 34913 Fat, Furry, Flexible, and Functionally Important: Characteristics of Mammals Living in the Arctic 357Niels M. Schmidt, Olivier Gilg, Jon Aars, and Rolf A. Ims13.1 Introduction 35713.2 The Mammal Assemblage in the Arctic Today 35813.2.1 Terrestrial Mammals 35813.2.2 Herbivores 35813.2.3 Predators 35913.2.4 Marine Mammals 36013.3 Arctic Mammals and Adaptations to Life in the Arctic 36613.3.1 Fur, Fat and Extremities 36713.3.2 Behavioral Adaptations to Life in the Arctic 36813.4 The Role of Mammals in Arctic Ecosystems 36913.4.1 Removal of Plant Material by Terrestrial Herbivores 37113.4.2 Transport of Nutrients and Seeds by Arctic Mammals 37213.4.3 Mammal Predator–Prey Dynamics 37413.5 The Future for Arctic Mammals in a Changing Climate 37413.6 Concluding Remarks 377References 37714 Ecology of Arctic Birds 385Anthony D. Fox14.1 Introduction: The Bird Species and Their Feeding Ecology 38514.2 Traveling to Breed 38614.3 Long Distance Migrations 38814.4 Reproduction 38914.5 Survival 39414.6 Population Change 39614.7 Climate Change 39714.8 Endangered Species 39914.9 Concluding Remarks 402References 40315 Arctic Ecology, Indigenous Peoples and Environmental Governance 409Mark Nuttall15.1 Introduction 40915.2 The Impacts of Social and Environmental Change 41015.3 Traditional Ecological Knowledge and Wildlife Management 41315.4 Arctic Ecology and Community-Based Monitoring 41515.5 Indigenous Peoples and Environmental Policy: The Case of the Inuit Circumpolar Council 41815.6 Concluding Remarks 419References 420Index 423

"An extremely useful source of data and information for individuals with research projects in arctic ecology, paleoecology, and climate change."—John T. Andrews, Arctic, Antarctic, and Alpine Research