PREFACE ........................................................ xv
ACKNOWLEDGMENTS .............................................. xvii
1 Why Study Continental Aquatic Systems? ....................... 1
Human Use of Water: Pressures on a Key Resource .............. 5
What Is the Value of Water Quality? .......................... 8
Advanced: Methods for Assigning Values to Ecosystem Goods
and Services ................................................ 12
Climate Change and Water Resources .......................... 14
Politics, Science, and Water ................................ 16
Summary ..................................................... 17
Questions for Thought ....................................... 18
2 Properties of Water ......................................... 19
Chemical and Physical Properties ............................ 20
Advanced: The Nature of Water ............................... 26
Relationships among Water Viscosity, Inertia, and Physical
Parameters .................................................. 27
Movement of Water ........................................... 31
Advanced: Equations Describing Properties of Moving Water ... 37
Forces That Move Water ...................................... 41
Summary ..................................................... 43
Questions for Thought ....................................... 44
3 Movement of Light, Heat, and Chemicals in Water ............. 45
Diffusion of Chemicals in Water ............................. 46
Light in Water .............................................. 52
Heat Balance in Water ....................................... 61
Summary ..................................................... 63
Questions for Thought ....................................... 63
4 Hydrologie Cycle and Physiography of Groundwater Habitats ... 65
Habitats and the Hydrologie Cycle ........................... 66
Advanced: Prediction of Amount and Variability of Runoff
with Global Climate Change .................................. 68
Movement of Water through Soil and Aquifers ................. 70
Groundwater Habitats ........................................ 76
Interaction of Groundwaters with Surface Waters ............. 80
Summary ..................................................... 81
Questions for Thought ....................................... 82
5 Hydrology and Physiography of Wetland Habitats .............. 83
Introduction and Definition of Wetland Habitats ............. 84
Wetland Conservation and Mitigation ......................... 88
Wetland Types ............................................... 89
Wetland Hydrology .......................................... 100
Wetlands and Global Change ................................. 103
Summary .................................................... 104
Questions for Thought ...................................... 105
6 Physiography of Flowing Water .............................. 107
Characterization of Streams ................................ 108
Stream Flow and Geology .................................... 116
Advanced: Classification of River and Stream Types as
a Restoration and Management Tool .......................... 126
Movement of Materials by Rivers and Streams ................ 127
Advanced: Characterizing the Movement of Dissolved
Materials in Rivers and Streams ............................ 135
Summary .................................................... 136
Questions for Thought ...................................... 137
7 Lakes and Reservoirs: Physiography ......................... 139
Formation: Geological Processes ............................ 140
Lake Habitats and Morphometry .............................. 151
Unique Properties of Reservoirs ............................ 155
Stratification ............................................. 156
Water Movement and Currents in Lakes ....................... 162
Summary .................................................... 165
Questions for Thought ...................................... 166
8 Types of Aquatic Organisms ................................. 167
The Species Concept ........................................ 168
Advanced: Molecular Methods for Assessing Microbial
Diversity in Natural Environments .......................... 172
Major Taxonomic Groups ..................................... 174
Classification of Organisms by Function, Habitats, and
Interactions ............................................... 176
Organisms Found in Freshwaters ............................. 180
Summary .................................................... 182
Questions for Thought ...................................... 182
9 Microbes and Plants ........................................ 185
Viruses .................................................... 187
Archaea .................................................... 190
Bacteria ................................................... 191
Cyanobacteria (Bluegreen Algae or Cyanophytes) .......... 195
Protoctista ................................................ 198
Eukaryotic Algae ........................................ 199
Rhodophyceae, the Red Algae ............................. 199
Chrysophyceae, the Golden Algae ......................... 199
Bacillariophyceae, the Diatoms .......................... 202
Dinophyceae, the Dinoflagellates ........................ 202
Euglenophyceae .......................................... 205
Chlorophyceae and Charophyceae, Green Algae and
Relatives ............................................... 205
Additional Algal Groups ................................. 208
Protozoa ................................................ 208
Fungi ...................................................... 210
Aquatic Fungi .............................................. 211
Aquatic Lichens ............................................ 212
Plantae .................................................... 212
Nonvascular Plants ......................................... 213
Vascular Plants ............................................ 213
Summary .................................................... 219
Questions for Thought ...................................... 219
10 Multicellular Animals ...................................... 221
Invertebrates .............................................. 222
Phylum Porifera ......................................... 222
Phylum Cnidaria ......................................... 223
Phyla Platyhelrninthes and Nemertea ..................... 225
Phylum Gastrotricba ..................................... 227
Phylum Rotifera ......................................... 227
Phylum Nematoda ......................................... 229
Phylum Nematomorpha ..................................... 229
Phylum Mollusca ......................................... 230
Phylum Annelida ......................................... 235
Bryozoans: Phyla Entoprocta and Ectoprocta .............. 236
Phylum Taidigrada ....................................... 238
Phylum Arthropoda ....................................... 238
Phylum Chordata, Subphylum Vertebrata ...................... 252
Fishes .................................................. 252
Tetrapods ............................................... 255
Summary .................................................... 255
Questions for Thought ...................................... 257
11 Evolution of Organisms and Biodiversity of Freshwaters ..... 259
Measures of Diversity ...................................... 261
Temporal and Spatial Factors Influencing Evolution of
Freshwater Organisms ....................................... 263
Short-Term Factors Influencing Local Distribution of
Species .................................................... 270
Genetics and Populations of Species ........................ 275
Advanced: Populations and Their Spatial Distribution ....... 276
The Controversy Over Endemism and Microbial Species ..... 277
Invasions of Nonnative Species ............................. 278
Extinction ................................................. 282
What Is the Value of Freshwater Species Diversity? ......... 287
Summary .................................................... 288
Questions for Thought ...................................... 288
12 Aquatic Chemistry and Factors Controlling Nutrient
Cycling: Redox and O2 ...................................... 289
Chemicals in Freshwaters ................................... 290
Redox Potential, Potential Energy, and Chemical
Transformations ............................................ 296
Oxygen: Forms and Transformations .......................... 301
Photosynthesis ............................................. 303
Respiration ................................................ 310
Controls of Distribution of Dissolved Oxygen in the
Environment ................................................ 312
Summary .................................................... 319
Questions for Thought ...................................... 320
13 Carbon ..................................................... 323
Forms of Carbon ............................................ 324
Inorganic Carbon ........................................ 324
Organic Carbon .......................................... 328
Transformations of Carbon .................................. 330
Advanced: Breakdown Rates of Organic Carbon ............. 331
Oxidation of Organic Carbon with Inorganic Electron
Acceptors Other Than O2 ................................. 333
Fermentation............................................. 334
Methanoьophy ............................................ 336
Metbanogenesis, Acetogenesis, and Disproportionation
of Acetate .............................................. 337
A Conceptual Introduction to Nutrient Cycling .............. 337
The Carbon Cycle ........................................... 340
Advanced: Hydrogen and Carbon Cycling ...................... 341
Summary .................................................... 342
Questions for Thought ...................................... 342
14 Nitrogen, Sulfur, Phosphorus, and Other Nutrients .......... 345
Nitrogen ................................................... 347
Nitrogen Forms .......................................... 347
Nitrogen Fluxes ......................................... 348
Advanced: Alternative Pathways of Anoxic N Cycling and
Nitrous Oxide Production by Freshwater Environments ..... 354
Nitrogen Cycle .......................................... 355
Sulfur ..................................................... 357
Sulfur Forms ............................................ 357
Fundamental Sulfur Transformations ...................... 357
Advanced: More Complex Sulfur Transformations ........... 359
Sulfur Cycle ............................................... 360
Phosphorus .............................................. 360
Phosphorus Forms ....................................... 361
Phosphorus Transformations .............................. 361
Silicon, Iron, and Other Trace Nutrient Cycles ............. 363
Silicon ................................................. 363
Iron .................................................... 364
Cycling of Other Elements ................................. 368
Gradients of Redox and Nutrient Cycles and Interactions
among the Cycles ........................................... 369
Summary .................................................... 372
Questions for Thought ...................................... 373
15 Unusual or Extreme Habitats ................................ 375
Adaptations to Extremes .................................... 377
Saline Lakes ............................................... 380
Advanced: Chemistry of Saline Lakes and Salt Production .... 382
Hot Springs ................................................ 383
Cold Habitats .............................................. 386
Temporary Waters and Small Pools ........................... 389
Ultraoligotrophic Habitats ................................ 393
Hypereutrophic Habitats .................................... 393
Deep Subsurface Habitats ................................... 394
The Water Surface Layer .................................... 396
Summary .................................................... 398
Questions for Thought ...................................... 398
16 Responses to Stress, Toxic Chemicals, and Other
Pollutants in Aquatic Ecosystems ........................... 399
Basic Toxicology ........................................... 403
Bioassessment .............................................. 406
Organic Pollutants ......................................... 408
Pharmaceuticals and Endocrine Disruptors ................... 415
Acid Precipitation ......................................... 417
Sources and Geography of Acid Precipitation ............. 417
Biological Effects of Acidification ..................... 418
Metals and Other Inorganic Pollutants ...................... 425
Salt Pollution ............................................. 430
Suspended Solids ........................................... 431
Thermal Pollution .......................................... 432
Human-Induced Increases in UV Radiation .................... 434
Summary .................................................... 435
Questions for Thought ...................................... 436
17 Nutrient Use and Remineralization .......................... 437
Use of Nutrients ........................................... 438
Nutrient Limitation and Relative Availability ........... 446
Relative Availability of Nutrients ...................... 446
Nutrient Limitation ..................................... 449
The Paradox of the Plankton and Nutrient Limitation ........ 453
Resource Ratios and Stoichiometry of Primary Producers ..... 454
Nutrient Remineralization .................................. 454
What Short-Term Processes Control the Levels of Dissolved
Inorganic Nutrients Such as Ammonium and Phosphate? ..... 456
Processes Leading to Remineralization ................... 458
Advanced: Remineralization as a Source of Nutrient
Pulses in Lentic Systems ................................ 461
Stoichiometry of Heterotrophs, Their Food, and Nutrient
Remineralization ........................................... 462
Summary .................................................... 466
Questions for Thought ...................................... 467
18 Trophic State and Eutrophication ........................... 469
Definition of Trophic State ................................ 471
Advanced: Determining Reference Nutrient Conditions in
Freshwater Environments .................................... 475
Why Do Algal Blooms Caused by Nutrient Pollution Matter
in Lakes? .................................................. 476
Natural and Cultural Processes of Eutrophication ........... 477
Relationships among Nutrients, Water Clarity, and
Phytoplankton: Managing Eutrophication in Lakes ............ 481
Advanced: Empirical Relationships Used to Predict Control
of Eutrophication .......................................... 485
Mitigating Lake Eutrophication ............................. 487
Control of Nutrient Sources ............................. 489
Treatment in the Lake ................................... 492
Macrophyte Removal ...................................... 493
Managing Eutrophication in Streams and Rivers .............. 495
Case Studies of Eutrophication in Lakes and Lotic Systems .. 496
Lake Washington ......................................... 496
Lake Trummen ............................................ 498
Lake Tahoe .............................................. 500
Lake Okeechobee ......................................... 501
The Clark Fork River .................................... 502
The Murray-Darling River ................................ 502
Managing Eutrophication in Wetlands ........................ 503
Wetlands as Nutrient Sinks ................................. 505
Summary ........ 506
Questions for Thought ...................................... 507
19 Behavior and Interactions among Microorganisms and
Invertebrates .............................................. 509
Behavior of Microorganisms ................................. 511
Motility ................................................ 511
Taxis ................................................... 512
Interaction Types in Communities ........................... 515
Predation and Parasitism Including the Microbial Loop ...... 516
Viruses ................................................. 519
Consumption of Small Planktonic Cells ................... 521
Scrapers ................................................ 524
Shredders ............................................... 525
Collector-Gatherers ..................................... 526
Filter Feeders .......................................... 527
Microbial Adaptations to Avoid Predation ................ 530
Parasitism .............................................. 531
Other Exploitative Interactions ......................... 534
Competition ................................................ 534
Mutualism: Facilitation and Syntrophy ...................... 539
Chemical Mediation of Microbial Interactions ............... 542
Summary .................................................... 543
Questions for Thought ...................................... 544
20 Predation and Food Webs .................................... 545
Herbivory .................................................. 546
Detritivory ................................................ 548
Omnivory ................................................... 550
Adaptation to Predation Pressure ........................... 553
Adaptations of Predators ................................... 557
Nonlethal Effects of Predation ............................. 560
Trophic Levels, Food Webs, and Food Chains ................. 561
The Trophic Cascade ........................................ 561
Summary .................................................... 568
Questions for Thought ...................................... 569
21 Nonpredatory Interspecific Interactions among Plants and
Animals in Freshwater Communities .......................... 571
Competition ................................................ 572
Mutualism and Facilitation ................................. 581
Other Species Interactions ................................. 583
Summary .................................................... 584
Questions for Thought ...................................... 585
22 Complex Community Interactions ............................. 587
Disturbance ................................................ 588
Succession ................................................. 592
Indirect Interactions ...................................... 598
Strong Interactors ......................................... 602
Theoretical Community Ecology and Aquatic Food Webs ........ 603
Thresholds and Alternative Stable States ................... 605
Invasion and Extinction Revisited .......................... 606
Summary .................................................... 608
Questions for Thought ...................................... 609
23 Fish Ecology and Fisheries ................................. 611
Biogeographical and Environmental Determinants of Fish
Assemblage Diversity ....................................... 612
Physiological Aspects Influencing Growth, Survival, and
Reproduction ............................................... 617
Population Dynamics of Fishes .............................. 621
Regulating Exploitation of Fish Stocks ..................... 627
Stocking Fish for Fisheries ................................ 629
Aquaculture ................................................ 630
Summary .................................................... 632
Questions for Thought ...................................... 633
24 Freshwater Ecosystems ...................................... 635
General Approaches to Ecosystems ........................... 636
Secondary Production ....................................... 638
Energy Fluxes and Nutrient Cycling ......................... 642
Nutrient Budgets ........................................... 643
Biodiversity and Ecosystem Function ........................ 646
Groundwater Ecosystems ..................................... 648
Streams and Rivers ......................................... 650
Lakes and Reservoirs ....................................... 658
Advanced: Reservoirs as Unique Ecosystems .................. 662
Wetlands ................................................... 663
Whole Ecosystem Experiments ................................ 668
Comparison of Freshwater Ecosystems ........................ 669
Summary .................................................... 672
Questions for Thought ...................................... 673
25 Conclusions ................................................ 675
Appendix: Experimental Design in Aquatic Ecology .............. 679
Natural Observations and Experiments ....................... 681
Multivariate Methods ....................................... 682
Simulation Modeling ........................................ 685
Manipulative Experiments ................................... 685
Whole System Manipulations ................................. 687
Summary .................................................... 688
GLOSSARY ...................................................... 691
REFERENCES .................................................... 719
TAXONOMIC INDEX ............................................... 787
SUBJECT INDEX ................................................. 795
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