LIMNOLOGY. Inland Water Ecosystems. JACOB KALFF McGill University. Prentice Hall. Upper Saddle River, New Jersey 07458

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1 LIMNOLOGY Inland Water Ecosystems JACOB KALFF McGill University Prentice Hall Prentice Hall Upper Saddle River, New Jersey 07458

2 Contents CHAPTER 1 Inland Waters and Their Catchments: An Introduction and Setting Introduction The Setting Organization of the Text 10 Acknowledgments 12 CHAPTER 2 The Development of Limnology Limnology and Its Roots Limnology Between World War I and World War II The Development of Ideas: Europe The Development of Ideas: North America Limnology after World War II Scales and Patterns: A Conceptual Exploration 2 7 CHAPTER 3 Water: A Unique and Important Substance Introduction Characteristics of Water 35 CHAPTER 4 Water Resources, Water Pollution, and Lakes Introduction Water Resources Lakes, Rivers, Wetlands, and Their Global Distribution A Look at "Typical" Lakes and Streams 48 CHAPTER 5 Hydrology and Climate Introduction Water Movement in Catchments 5.3 Humans and the Hydrologic Cycle 5.4 Global Patterns in Precipitation and Runoff Runoff and the Presence of Waterbodies Water Inputs anc1 Outputs The Aral Sea 68 The Caspian Sea 70 CHAPTER 6 Origin and Age of Lakes Introduction Glacial Lakes Tectonic Lakes Coastal Lakes '" 64 Riverine Lakes Volcanic Lakes Solution or Karst: Lakes 82 Manmade Lakes or Reservoirs 82 CHAPTER 7 Lake and Catchment Morphometry Introduction The Bathymetric Map Lake Surface Area Lake Depth VII

3 viii 7.5 Lake Shape Underwater and Catchment Slopes 91 CHAPTER 8 Rivers and the Export of Materials from Drainage Basins and the Atmosphere Introduction Flowing Water Systems Rivers and Their Ecotones Rivers, Their Banks, and Human Activity Drainage-Basin Export of Nitrogen and Phosphorus Atmospheric Deposition of Nutrients Nutrient Export, Catchment Size, Lake Morphometry, and the Biota: A Conceptualization Organic Carbon Export from Drainage Basins 117 CHAPTER 9 Aquatic Systems and their Catchments Catchment Size" Catchment Form Catchment Soils and Vegetation Water Residence Time Nutrient Concentrations, Trophic State, andwrt Retention of Dissolved and Particulate Materials by Lakes and Reservoirs Sediment Loading to Aquatic Systems 133 CHAPTER 10 Light Introduction Detectors Light Above and Below the Water Surface Absorption, Transmission, and Scattering of Light in Water Ultraviolet Radiation and Its Effects Light Attenuation Light Attenuation and Photosynthesis Light Attenuation and Lake Stratification The Secchi Disc and Its Utility Limitations of the Secchi Disc Light and Primary Production Underwater Vision 153 CHAPTER 11 Temperature Cycles, Lake Stratification, and Heat Budgets Introduction Types of Stratification and Mixing Morphometry and Stratification Seasonal Temperature Cycles and Stratification Stability of Stratification Stability of Temperate vs Tropical Lakes Thermocline Depth Thermocline Shape Meromictic Lakes Development of Meromixis Heat Budgets Climatic Change and Aquatic Systems 174 CHAPTER 12 Water Movements Introduction Laminar vs Turbulent Flow Surface Gravity Waves Turbulent Flow and Measures of Stability Coefficient of Vertical Eddy Coefficient of Horizontal Eddy Diffusion 187

4 IX 12.7 Horizontal Currents Long-term Surface Current Patterns Langmuir Currents Standing Surface Waves Internal or Thermocline Seiches Internal Seiches, Hypolimnetic Currents, and Sediment Resuspension Turbulent Mixing and the Biota 199 CHAPTER 13 Salinity and Major Ion Composition of Lakes and Rivers Introduction Salinity and Its Origins Total Salinity and Its Determination Major Ion Composition Human Activity, Climate, and Ion Composition Saline Lakes and Their Distribution Ionic Composition of Inland Saline Lakes and Wetlands" The Salinity Spectrum and the Biota 214 CHAPTER 14 Inorganic Carbon and ph Introduction 218, 14.2 Carbon Dioxide in Water ph and Its Range in Aquatic Systems Alkalinity of Inland Waters ph, Extreme Environmental Conditions, and Species Richness Carbonates: Precipitation and Solubilization 222 CHAPTER 15 Dissolved Oxygen Introduction Solubility of Oxygen in Water Sources and Sinks of Oxygen Photosynthesis, Respiration, and DOC Dissolved Oxygen Consumption and Lake Productivity Oxygen Depletion in Ice-covered Waters Dissolved Oxygen and the Biota 236 CHAPTER 16 Oxidation-Reduction Potential 16.1 Introduction Redox Reactions and Nutrient Cycling 239 CHAPTER 17 Phosphorus Concentrations and Cycling Introduction The Classical Model of Phosphorus Cycling The Modern Model and Aerobic Phosphorus Release The MassVBalance Equation and Phosphorus Cycling ' Sediment Phosphorus Release and Phytoplankton Production Phosphorus Control, Internal Loading, and Lake Management The Empirical Modeling of Phosphorus The Dynamic Modeling of Phosphorus 265 CHAPTER 18 Nitrogen Cycling Introduction: The Atmosphere, the Land, and the Water Nitrogen Transformation Processes Nitrification Denitrification 275

5 18.5 Nitrogen Fixation: Rates and Process Nitrogen Fixation Rates: Plankton vs Littoral Zone Forms and Quantities of Nitrogen in Inland Waters 280 CHAPTER 19 Iron, Manganese, and Sulfur Introduction Iron Cycling Iron and Sulfur Iron and Organic Matter The Manganese Cycle Iron, Manganese, and Trace Metals 289 CHAPTER 20 Particle Sedimentation and Sediments Introduction Origin and Distribution of Sediments Sedimentation and Sediment Traps Sinking Velocities and Sedimentation Rates The Sediment Record Dating Sediments Profundal Sediment Characteristics 307 CHAPTER 21 The Phytoplankton Introduction Species Composition and Phylogenetic Generalities Phytoplankton Size and Activity: Small Cells vs Large Cells Seasonal Biomass Cycles: A Conceptual Model The Composition of Phytoplankton Cells 327 -' ' 21.6 Algal Sedimentation and Buoyancy Control Parasitism and Disease Photosynthesis, Light, and Temperature Photosynthesis, Respiration, and Growth Primary Production in Nature Production: Biomass (P:B) Ratios and Specific Growth Rates in Nature Limiting Nutrients and Eutrophication Nitrogen vs Phosphorus Empirical Nutrient-Phytoplankton Relationships The Maximum Phytoplankton Biomass 346 CHAPTER 22 The Bacteria Introduction From Past to Present Bacterial Size, Form, and Metabolism Abundance, Biomass, and Distribution Heterotrophic Bacterial Abundance and Environmental Factors Resource Limitation vs Grazing Control of Bacterial Abundance Heterotrophic Bacteria: Production, Losses, and System Contribution Viruses The Microbial Food Web Photosynthetic Bacteria Heterotrophic Sediment Bacteria 373 CHAPTER 23 Zooplankton Introduction Zooplankton Sampling Protozoa, Rotifers, and Crustaceans Species Richness and Its Prediction 3 84

6 XI 23.5 Seasonal Cycles Long-term Variation in Zooplankton Abundance Top-down Control of Zooplankton Biomanipulation and Lake Management Chaoborus: The Phantom Midge Zooplankton Feeding Nutrient Cycling and Zooplankton Resource Availability and Zooplankton Biomass Zooplankton Production Diel Migration and Cyclomorphosis 404 CHAPTER 24 Benthic Plants Introduction Wetlands and Their Utilization Macrophyte Distribution and Species Richness? Macrophyte Biomass and Its Determinants Submerged Macrophyte Distribution: Light and Lake Morphometry Submerged Macrophyte Distributions and Plant Nutrients Submerged Macrophyte Distribution and Dissolved Inorganic Carbon (DIC): A Physiological Exploration Plant Size, Community Structure, and Function Attached Algae Eutrophication and Benthic Plants Lake Management and Macrophytes 43 0 CHAPTER 25 Zoobenthos Introduction Taxonomic Distribution, Species Richness, and Abundance Life-History Aspects Lake Morphometry, Substrate Characteristics, and the Zoobenthos Resource and Predation Control The Zebra Mussel: A Keystone Species The Zoobenthos and Energy Flow in Lakes 448 CHAPTER 26 Fish and Water Birds Introduction Fish Species and Species Richness Life-History Attributes and Population Dynamics of Age-0 Fish Fish Growth: Determinants and Measurement Fisheries and Fisheries Management The Biology of a Temperate Zone Fish and a Tropical Fish: Perch and Tilapia Predicting Fish Biomass, Production, - and Yield Aquaculture and Water Quality Water Birds 475 CHAPTER 27 Acidification of Waterways Introduction Sources and Distributions Acid-Sensitive Waters Characteristics of Acid-Sensitive Waters and Catchments Catchments and Lake Acidification: Wet and Dry Deposition Neutralization and Buffering Processes in Catchments Buffering Capacity of Lakes, Rivers, and Wetlands Aluminum and Other Toxic Metals Effects of Acidification on the Aquatic Biota Modeling the Acidification Process i 493

7 Xll Lake Management: Recovering from. Acidification The Future 497 CHAPTER 28 Contaminants Introduction Toxic Substances Sources of Contaminants The Fate of Contaminants The Sediment Record Physical and Chemical Characteristics of Contaminants and Their Distribution in Nature Toxicity and Its Prediction Bioaccumulation and Biomagnification Mercury and the Mercury Cycle Toxic Chemicals, Environmental Health, and Lake Management Natural Lakes and Reservoirs 524 The River-Lake-Reservoir Continuum 529 Water Residence Time and Plankton Growth Rates 530 Reservoir Zonation: A Conceptual View 531 Drawdowns 532 Reservoir Aging and the Trophic Upsurge 533 Large Reservoirs and Their Impacts APPENDIX 1 International Organization for Standardization of Country Codes APPENDIX 2 Conversion Factors for Selected Elements and Reported Species 538. CHAPTER 29 Reservoirs 523 BIBLIOGRAPHY Introduction 523 INDEX 573