Principles of Terrestrial Ecosystem Ecology

455 Pages·2002·9.11 MB·English

by  Harold A. Mooney, F. Stuart Chapin III, Pamela A. Matson, M.C. Chapin

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