Contents. Introduction. Supporting Processes

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2 Contents Introduction Chapter 1 Ecosystem Perspective and Bivalve Molluscs Evolutionary History 3 Ecosystems 5 Structural Attributes 6 System Hierarchy 6 Food Webs 8 Networks 10 Biodiversity 12 Functional Attributes 12 System Types 12 Remaining Contents 13 References 15 Supporting Processes Chapter 2 Physical-Environmental Interactions Introduction 19 Temperatura 19 Salinity 23 Temperature-Salinity and Other Factor Combinations 25 Water Movement 26 Tides 30 Sediments 31 References 32 Chapter 3 Organismic Level Processes Introduction 35 Feeding 36 Filter (Suspensión) Feeding 36 The Bivalve Filter and Pump 36 Filtration and Partióle Quality 38 Dissolved Organic Matter (DOM) 41 Filtration and Particle Size 42 Filtration Models 43 Deposit Feeding 46 Microbial Stripping 47

3 Cellulase Activity and Detritus Feeding 48 Particle Selection 48 Proportion of Foods 48 Optimal Foraging for Food in the Sediments 49 Shipworms 49 Symbiotic Nutrition 50 Photoautotrophic Symbioses 51 Chemoautotrophic Symbioses 54 Production (P) 57 Production Due to Growth (Pg) 57 Growth measurement 58 Shell growth 58 Production Due to Reproduction (Pr) 60 Growth Models 61 Respiration (R) 63 Excretion 66 References 68 Chapter 4 Population Processes Introduction 75 Life Cycle 75 Statistical Measures of Populations 76 Density 76 Reproduction 78 Population Growth 79 Bivalves as Open or Metapopulations 82 Life and Fecundity Tables 82 Mortality 85 Abiotic Mortality 86 Predation 86 Competition Induced Mortality 94 Aggregated Distributions 94 Zonation 97 Diseases and Parasites 98 Population Energy Budgets 101 References 103 Ecosystem Processes Chapter 5 Grazing Introduction 113 Grazing Theory as Applied to Bivalves 113 The Officer Conceptual Model for Bivalve Grazing 114 Methods Used to Estímate System Grazing 117 Upstream-Downstream Observations 117

4 Free Flow Designs 117 Laboratory Flumes 118 Field Flumes 120 Tunnels 121 Ecosystem Grazing Estimates 123 Askó, Sweden 123 Bay of Brest, France 123 Chesapeake Bay, U.S.A 124 Delaware Bay, U.S.A 124 Killary Harbour, Ireland 125 Laholm Bay, Sweden 125 Lowes Cove, Maine, U.S.A 126 Bay of Marennes-Oléron, France 126 Narragansett Bay, Rhode Island, U.S.A 126 North Inlet, South Carolina, U.S.A 127 Oosterschelde Estuary, The Netherlands 127 Ria de Arosa, Spain 128 San Francisco Bay, California, U.S.A 128 Eastern Wadden Sea, Sylt, Germany 130 Western Wadden Sea, The Netherlands 131 Ecosystem Comparisons 132 References 136 Chapter 6 System Metabolism and Nutrient Cycling System Metabolism 143 Methods for Measuring System Metabolism 143 Chamber Measurements 144 Free-Flow Measurements 145 Tunnel Studies 146 Nutrient Cycling 146 Theoretical Background 147 Nutrient Cycles 149 Carbón 149 Nitrogen 156 Phosphorus 164 Silicon 165 System Nutrient Turnover 165 Coupling Metabolism and Nutrients 167 Conclusions 167 References 168 Chapter 7 Ecosystem Experiments: Models, Experimental Ecosystems, and Field Manipulations Introduction 173 Models 175 System Simulation Models 176

5 Bivalve-Carrying Capacity of Marennes-Oléron 176 SMOES: Simulation Model Oosterschelde Ecosystem 181 Model Formation 181 Model Analysis 186 System Analytical Models 187 Oyster Reef Model 187 Marsh Fertilization and Bivalve Production 191 Chesapeake Bay Ecosystem Model and Oysters 192 Experimental Ecosystems 196 Merl, Rhode Island U.S.A 197 Roskilde Fjord, Denmark 198 Oresund and Skagerrak, Sweden 200 Oosterschelde, The Netherlands 200 Mesocosms and Bivalves: Conclusions 202 Ecosystem Field Experiments 202 Incidental Ecosystem Scale Experiments 203 Designed Ecosystem Scale Experiments 205 References 207 Chapter 8 Bivalves as Components of Ecosystem Health Introduction 213 Bivalves as Monitors 214 Bivalves as Integrators 215 Bivalve Responses 218 Shell Movements 218 Mortality 219 Shell and Tissue Growth 220 Bivalve Scope for Growth (SFG) 221 System Measures 223 Conclusions 224 References Synthesis and Conclusions Chapter 9 Conclusions Keystone Species 229 Ecosystem Engineers 230 Ecosystem Level Roles of Bivalves 230 Trophic 230 Nutrient Cycling 231 Structural 231 Monitors and Indicators 232 Ecosystem Issues 232 Organismic Level 232

6 Population Level 233 Ecosystem Level 233 Synthesis 234 References 234 Index