F.Stuart Chapin III Pamela A.Matson Harold A.Mooney Principles of Terrestrial Ecosystem Ecology Illustrated by Melissa C.Chapin With 199 Illustrations Springer
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
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 ha ol the ice-free terres use 0%of terrestrial produc Our ac r ex th vent in the :f history of hte on fields trean tte toprovide their sensitivit to on ng and biotic cha We believe that an understanding of how ecosystems erate and change must t itoduce the of ye 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 rt sho osphere,ocean,cl systems and contribute to the ecosystem str th nd fo the fows by nd nd thst carbon and nutrients.We then co are and contrast these between terrestrial and aquatic e systems.We also consider th important role that organisms have on ecosystem 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-
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 terrestrial 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 conceptual 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 understanding 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 function 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), environmental 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 sociPreface v
vi Preface eties.Powerpoint lecture notes developed by one of the authors are available online(www.faculty.uafedu/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. Beringer. Bonine Cham ryan Rich Boone e h az.Bitha olf- im hmer.Ste Val Scott Fendorf.Jon Folev.David Fo man.Paul Grogan.Diego Gurvich.Bill Heal.Sarah Hobbie.Dave Hooper.Shuijin Hu.Pilar Huante.Bruce Hunga Jay jones,Jurg Luterbacher,Frank Kelliher,Jennifer King. Dave Kline christian korner Hans iambers amanda iynch michelle Mack,Steve MacLean,Joe McFadden,Dave McGuire,Sam MeNaughton,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 Parde Serreze,Gus Shaver,Ni ave alentine larly thank Phil C rogan ark Ch ck Endres,an d Ro cnes:a maps E Start Chapin Pamela A.Matson Harold A.Mooney
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, Wolfgang 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 Groffman, 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
Contents Preface........... Part I Context 。 2 of E Ecosystem Structure o Controls over Ecosystem Processes Human-Caused Changes in Earth's Ecosystems Summary 1 Review ouestions 11 Additional Reading............................. 17 Chapter2 Earth's Climate System 8811248 Ocean structur Landform Effects on Climate 31 Vegetation Influences on Climate Temporal Variability in Climate Long-Term Changes. Interannual Climate Variability Seasonal and Daily Variations 40
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
viii Contents Relationship of climate to ecosystem distribution and Structure 41 Summary Review ouestions Additional Reading 45 Chapter 3 Geology and Soils Introduction 。。。。。。。。。。。。 Soil Formation 48 pography Human activities Controls over Soil Loss 5 Development of Soil Profiles Additions to Soils Soil Transformations Soil Transfers Losses from Soils Soil Horizons and Soil Classification Soil Properties and Ecosystem Functioning 586656 Additional Reading PartⅡMechanisms Chanter 4 Terrestrial Water and Energy Balance Introduction Surface eneroy balance Solar Radiation Budget 73 Ecosystem Radiation Budget Energy Partitioning Seasonal Energy Exchange Water Inputs to Ecosystems Water Movements Within Ecosystems Basic Principles of Water Movement rom the Canopy to the Soil vement 777908 om S hrough Plants 05 89 Wet C vapotranspiration from Dry Ev porat nopies
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