Principles of Terrestrial Ecosystem Ecology PDF
Preview Principles of Terrestrial Ecosystem Ecology
Principles of Terrestrial Ecosystem Ecology F. Stuart Chapin III Pamela A. Matson Harold A. Mooney Springer PTEPR 10/9/2002 7:23 PM Page i Principles of Terrestrial Ecosystem Ecology PTEPR 10/9/2002 7:23 PM Page ii Springer New York Berlin Heidelberg Barcelona Hong Kong London Milan Paris Singapore Tokyo PTEPR 10/9/2002 7:23 PM Page iii F. Stuart Chapin III Pamela A. Matson Harold A. Mooney Principles of Terrestrial Ecosystem Ecology Illustrated by Melissa C. Chapin With 199 Illustrations 1 3 PTEPR 10/9/2002 7:23 PM Page iv F. Stuart Chapin III Pamela A. Matson Institute of Arctic Biology Department of Geological and University of Alaska Environmental Science Fairbanks, AK 99775 School of Earth Sciences USA Green 355 PTEPR 10/9/2002 7:23 PM Page v Preface Human activities are affecting the global environment in myriad ways, with numerous direct and indirect effects on ecosystems. The climate and atmospheric composition of Earth are changing rapidly. Humans have directly modified half of the ice-free terres- trial surface and use 40% of terrestrial production. Our actions are causing the sixth major extinction event in the history of life on Earth and are radically modifying the interactions among forests, fields, streams, and oceans. This book was written to provide a con- ceptual basis for understanding terrestrial ecosystem processes and their sensitivity to environmental and biotic changes. We believe that an understanding of how ecosystems operate and change must underlie our analysis of both the consequences and the mitigation of human-caused changes. This book is intended to introduce the science of ecosystem ecology to advanced undergraduate students, beginning graduate students, and practicing scientists from a wide array of disciplines. We also provide access to some of the rapidly expanding literature in the many disciplines that contribute to ecosystem understanding. The first part of the book provides the context for understand- ing ecosystem ecology. We introduce the science of ecosystem ecology and place it in the context of other components of the Earth System—the atmosphere, ocean, climate and geological systems. We show how these components affect ecosystem processes and contribute to the global variation in terrestrial ecosystem structure and processes. In the second part of the book, we consider the mechanisms by which terrestrial ecosystems func- tion and focus on the flow of water and energy and the cycling of carbon and nutrients. We then compare and contrast these cycles between terrestrial and aquatic ecosystems. We also consider the important role that organisms have on ecosystem processes through trophic interactions (feeding relationships), environmen- tal effects, and disturbance. The third part of the book addresses temporal and spatial patterns in ecosystem processes. We finish by considering the integrated effects of these processes at the global scale and their consequences for sustainable use by human soci- v PTEPR 10/9/2002 7:23 PM Page vi vi Preface eties. Powerpoint lecture notes developed by one of the authors are available online (www.faculty.uaf.edu/fffsc/) as supplementary material. Many people have contributed to the development of this book. We particularly thank our families, whose patience has made the book possible, and our students from whom we have learned many of the important ideas that are presented. In addition, we thank the following individuals for their constructively critical review of chapters in this book: Kevin Arrigo, Teri Balser, Perry Barboza, Jason Beringer, Kim Bonine, Rich Boone, Syndonia Bret-Harte, John Bryant, Inde Burke, Zoe Cardon, Oliver Chadwick, Scott Chambers, Melissa Chapin, Kathy Cottingham, Joe Craine, Wolf- gang Cramer, Steve Davis, Sandra Diaz, Bill Dietrich, Rob Dunbar, Jim Ehleringer, Howie Epstein, Werner Eugster, Valerie Eviner, Scott Fendorf, Jon Foley, David Foster, Tom Gower, Peter Groff- man, Paul Grogan, Diego Gurvich, Bill Heal, Sarah Hobbie, Dave Hooper, Shuijin Hu, Pilar Huante, Bruce Hungate, Jill Johnstone, Jay Jones, Jürg Luterbacher, Frank Kelliher, Jennifer King, Dave Kline, Christian Körner, Hans Lambers, Amanda Lynch, Michelle Mack, Steve MacLean, Joe McFadden, Dave McGuire, Sam McNaughton, Knute Nadelhoffer, Jason Neff, Mark Oswood, Bob Paine, Bill Parton, Natalia Perez, Steward Pickett, Stephen Parder, Mary Power, Jim Randerson, Bill Reeburgh, Peter Reich, Jim Reynolds, Roger Ruess, Steve Running, Scott Rupp, Dave Schimel, Josh Schimel, Bill Schlesinger, Guthrie Schrengohst, Ted Schuur, Stephen Parder Mark Serreze, Gus Shaver, Nigel Tapper, Monica Turner, Dave Valentine, Peter Vitousek, Lars Walker, and Katey Walter.We particularly thank Phil Camil,Valerie Eviner, Jon Foley, and Paul Grogan for comments on the entire book; Mark Chapin, Patrick Endres, and Rose Meier for comments on illustrations; Phil Camil for comments on educational approaches; and Jon Foley and Nick Olejniczak for providing global maps. F. Stuart Chapin III Pamela A. Matson Harold A. Mooney PTEPR 10/9/2002 7:23 PM Page vii Contents Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v Part I Context Chapter 1 The Ecosystem Concept Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Overview of Ecosystem Ecology . . . . . . . . . . . . . . . . . . . 3 History of Ecosystem Ecology . . . . . . . . . . . . . . . . . . . . . 7 Ecosystem Structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Controls over Ecosystem Processes . . . . . . . . . . . . . . . . . 11 Human-Caused Changes in Earth’s Ecosystems . . . . . . . 13 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Review Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Additional Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Chapter 2 Earth’s Climate System Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Earth’s Energy Budget . . . . . . . . . . . . . . . . . . . . . . . . . . 18 The Atmospheric System . . . . . . . . . . . . . . . . . . . . . . . . . 21 Atmospheric Composition and Chemistry . . . . . . . . . . 21 Atmospheric Structure . . . . . . . . . . . . . . . . . . . . . . . . . 22 Atmospheric Circulation . . . . . . . . . . . . . . . . . . . . . . . 24 The Oceans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Ocean Structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Ocean Circulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Landform Effects on Climate . . . . . . . . . . . . . . . . . . . . . 31 Vegetation Influences on Climate . . . . . . . . . . . . . . . . . . 32 Temporal Variability in Climate . . . . . . . . . . . . . . . . . . . . 34 Long-Term Changes . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Interannual Climate Variability . . . . . . . . . . . . . . . . . . 38 Seasonal and Daily Variations . . . . . . . . . . . . . . . . . . . 40 vii PTEPR 10/9/2002 7:23 PM Page viii viii Contents Relationship of Climate to Ecosystem Distribution and Structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Review Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Additional Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 Chapter 3 Geology and Soils Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 Controls over Soil Formation . . . . . . . . . . . . . . . . . . . . . 46 Parent Material . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 Climate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Topography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 Potential Biota . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 Human Activities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 Controls over Soil Loss . . . . . . . . . . . . . . . . . . . . . . . . . . 50 Development of Soil Profiles . . . . . . . . . . . . . . . . . . . . . . 53 Additions to Soils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 Soil Transformations . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 Soil Transfers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 Losses from Soils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 Soil Horizons and Soil Classification . . . . . . . . . . . . . . . . 58 Soil Properties and Ecosystem Functioning . . . . . . . . . . 61 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 Review Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Additional Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Part II Mechanisms Chapter 4 Terrestrial Water and Energy Balance Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 Surface Energy Balance . . . . . . . . . . . . . . . . . . . . . . . . . . 73 Solar Radiation Budget . . . . . . . . . . . . . . . . . . . . . . . . 73 Ecosystem Radiation Budget . . . . . . . . . . . . . . . . . . . . 74 Energy Partitioning . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Seasonal Energy Exchange . . . . . . . . . . . . . . . . . . . . . 77 Water Inputs to Ecosystems . . . . . . . . . . . . . . . . . . . . . . 77 Water Movements Within Ecosystems . . . . . . . . . . . . . . . 78 Basic Principles of Water Movement . . . . . . . . . . . . . . 78 Water Movement from the Canopy to the Soil . . . . . . 79 Water Movement Within the Soil . . . . . . . . . . . . . . . . . 80 Water Movement from Soil to Roots . . . . . . . . . . . . . . 81 Water Movement Through Plants . . . . . . . . . . . . . . . . 83 Water Losses from Ecosystems . . . . . . . . . . . . . . . . . . . . 89 Evaporation from Wet Canopies . . . . . . . . . . . . . . . . . 89 Evapotranspiration from Dry Canopies . . . . . . . . . . . . 90 PTEPR 10/9/2002 7:23 PM Page ix Contents ix Changes in Storage . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 Runoff . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95 Review Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96 Additional Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96 Chapter 5 Carbon Input to Terrestrial Ecosystems Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 Photosynthetic Pathways . . . . . . . . . . . . . . . . . . . . . . . . . 98 C3 Photosynthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 C4 Photosynthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102 Crassulacean Acid Metabolism Photosynthesis . . . . . . 103 Net Photosynthesis by Individual Leaves . . . . . . . . . . . . 105 Basic Principle of Environmental Control . . . . . . . . . . 105 Light Limitation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105 CO2 Limitation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 Nitrogen Limitation and Photosynthetic Capacity . . . . 110 Water Limitation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 Temperature Effects . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 Pollutants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115 Gross Primary Production . . . . . . . . . . . . . . . . . . . . . . . . 115 Canopy Processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115 Satellite Estimates of GPP . . . . . . . . . . . . . . . . . . . . . . 117 Controls over GPP . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 Review Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 Additional Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122 Chapter 6 Terrestrial Production Processes Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123 Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123 Plant Respiration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125 Physiological Basis of Respiration . . . . . . . . . . . . . . . . 125 Net Primary Production . . . . . . . . . . . . . . . . . . . . . . . . . . 127 What Is NPP? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127 Physiological Controls over NPP . . . . . . . . . . . . . . . . . 128 Environmental Controls over NPP . . . . . . . . . . . . . . . 129 Allocation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132 Allocation of NPP . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132 Allocation Response to Multiple Resources . . . . . . . . 133 Diurnal and Seasonal Cycles of Allocation . . . . . . . . . 134 Tissue Turnover . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 Global Distribution of Biomass and NPP . . . . . . . . . . . . 137 Biome Differences in Biomass . . . . . . . . . . . . . . . . . . . 137 Biome Differences in NPP . . . . . . . . . . . . . . . . . . . . . . 138 Net Ecosystem Production . . . . . . . . . . . . . . . . . . . . . . . 140
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