Principles of Terrestrial Ecosystem Ecology

Transcription

1 Principles of Terrestrial Ecosystem Ecology

2 Springer New York Berlin Heidelberg Barcelona Hong Kong London Milan Paris Singapore Tokyo

3 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

4 F. Stuart Chapin III Institute of Arctic Biology University of Alaska Fairbanks, AK USA Harold A. Mooney Department of Biological Sciences Herrin Hall, MC 5020 Stanford University Stanford, CA USA Pamela A. Matson Department of Geological and Environmental Science School of Earth Sciences Green 355 Stanford University Stanford, CA USA Cover illustration: Waterfall and forests on Valean Poas in Costa Rica. Photograph by Peter Vitousek. Library of Congress Cataloging-in-Publication Data Chapin, F. Stuart (Francis Stuart), III. Principles of terrestrial ecosystem ecology / F. Stuart Chapin III, Pamela A. Matson, Harold A. Mooney. p. cm. Includes bibliographical references (p. ) ISBN (hc :alk. paper) ISBN (sc :alk. paper) 1. Ecology. 2. Biogeochemical cycles. 3. Biological systems. I. Matson, P. A. (Pamela A.) II. Mooney, Harold A. III. Title. QH541.C dc ISBN (hardcover) Printed on acid-free paper. ISBN (softcover) 2002 Springer-Verlag New York, Inc. All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer-Verlag New York, Inc., 175 Fifth Avenue, New York, NY 10010, USA), except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden. The use in this publication of trade names, trademarks, service marks, and similar terms, even if they are not identified as such, is not to be taken as an expression of opinion as to whether or not they are subject to proprietary rights. Printed in the United States of America SPIN (hardcover) SPIN (softcover) Springer-Verlag New York Berlin Heidelberg A member of BertelsmannSpringer Science+Business Media GmbH

5 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 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 soci- v

6 vi Preface eties. Powerpoint lecture notes developed by one of the authors are available online ( 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

7 Contents Preface v Part I Context Chapter 1 The Ecosystem Concept Introduction Overview of Ecosystem Ecology History of Ecosystem Ecology Ecosystem Structure Controls over Ecosystem Processes Human-Caused Changes in Earth s Ecosystems Summary Review Questions Additional Reading Chapter 2 Earth s Climate System Introduction Earth s Energy Budget The Atmospheric System Atmospheric Composition and Chemistry Atmospheric Structure Atmospheric Circulation The Oceans Ocean Structure Ocean Circulation Landform Effects on Climate Vegetation Influences on Climate Temporal Variability in Climate Long-Term Changes Interannual Climate Variability Seasonal and Daily Variations vii

8 viii Contents Relationship of Climate to Ecosystem Distribution and Structure Summary Review Questions Additional Reading Chapter 3 Geology and Soils Introduction Controls over Soil Formation Parent Material Climate Topography Time Potential Biota Human Activities Controls over Soil Loss Development of Soil Profiles Additions to Soils Soil Transformations Soil Transfers Losses from Soils Soil Horizons and Soil Classification Soil Properties and Ecosystem Functioning Summary Review Questions Additional Reading Part II Mechanisms Chapter 4 Terrestrial Water and Energy Balance Introduction Surface Energy Balance Solar Radiation Budget Ecosystem Radiation Budget Energy Partitioning Seasonal Energy Exchange Water Inputs to Ecosystems Water Movements Within Ecosystems Basic Principles of Water Movement Water Movement from the Canopy to the Soil Water Movement Within the Soil Water Movement from Soil to Roots Water Movement Through Plants Water Losses from Ecosystems Evaporation from Wet Canopies Evapotranspiration from Dry Canopies

9 Contents ix Changes in Storage Runoff Summary Review Questions Additional Reading Chapter 5 Carbon Input to Terrestrial Ecosystems Introduction Overview Photosynthetic Pathways C 3 Photosynthesis C 4 Photosynthesis Crassulacean Acid Metabolism Photosynthesis Net Photosynthesis by Individual Leaves Basic Principle of Environmental Control Light Limitation CO 2 Limitation Nitrogen Limitation and Photosynthetic Capacity Water Limitation Temperature Effects Pollutants Gross Primary Production Canopy Processes Satellite Estimates of GPP Controls over GPP Summary Review Questions Additional Reading Chapter 6 Terrestrial Production Processes Introduction Overview Plant Respiration Physiological Basis of Respiration Net Primary Production What Is NPP? Physiological Controls over NPP Environmental Controls over NPP Allocation Allocation of NPP Allocation Response to Multiple Resources Diurnal and Seasonal Cycles of Allocation Tissue Turnover Global Distribution of Biomass and NPP Biome Differences in Biomass Biome Differences in NPP Net Ecosystem Production

10 x Contents Ecosystem Carbon Storage Leaching Lateral Transfers Disturbance Controls over Net Ecosystem Production Net Ecosystem Exchange Global Patterns of NEE Summary Review Questions Additional Reading Chapter 7 Terrestrial Decomposition Introduction Overview Leaching of Litter Litter Fragmentation Chemical Alteration Fungi Bacteria Soil Animals Temporal and Spatial Heterogeneity of Decomposition. 157 Temporal Pattern Spatial Pattern Factors Controlling Decomposition The Physical Environment Substrate Quality and Quantity Microbial Community Composition and Enzymatic Capacity Long-Term Storage of Soil Organic Matter Decomposition at the Ecosystem Scale Aerobic Heterotrophic Respiration Anaerobic Heterotrophic Respiration Summary Review Questions Additional Reading Chapter 8 Terrestrial Plant Nutrient Use Introduction Overview Nutrient Movement to the Root Diffusion Mass Flow Root Interception Nutrient Uptake Nutrient Supply Development of Root Length Mycorrhizae

11 Contents xi Root Uptake Properties Nutrient Use Nutrient Loss from Plants Senescence Leaching Loss from Plants Herbivory Other Avenues of Nutrient Loss from Plants Summary Review Questions Additional Reading Chapter 9 Terrestrial Nutrient Cycling Introduction Overview Nitrogen Inputs to Ecosystems Biological Nitrogen Fixation Nitrogen Deposition Internal Cycling of Nitrogen Overview of Mineralization Production and Fate of Dissolved Organic Nitrogen Production and Fate of Ammonium Production and Fate of Nitrate Temporal and Spatial Variability Pathways of Nitrogen Loss Gaseous Losses of Nitrogen Ecological Controls Solution Losses Erosional Losses Other Element Cycles Phosphorus Sulfur Essential Cations Nonessential Elements Interactions Among Element Cycles Summary Review Questions Additional Reading Chapter 10 Aquatic Carbon and Nutrient Cycling Introduction Ecosystem Properties Oceans Carbon and Light Availability Nutrient Availability Carbon and Nutrient Cycling Lakes Controls over NPP

12 xii Contents Carbon and Nutrient Cycling Streams and Rivers Carbon and Nutrient Cycling Summary Review Questions Additional Reading Chapter 11 Trophic Dynamics Introduction Overview Plant-Based Trophic Systems Controls over Energy Flow Through Ecosystems Ecological Efficiencies Food Chain Length and Trophic Cascades Seasonal Patterns Nutrient Transfers Detritus-Based Trophic Systems Integrated Food Webs Mixing of Plant-Based and Detritus-Based Food Chains Food Web Complexities Summary Review Questions Additional Reading Chapter 12 Community Effects on Ecosystem Processes Introduction Overview Species Effects on Ecosystem Processes Species Effects on Resources Species Effects on Climate Species Effects on Disturbance Regime Species Interactions and Ecosystem Processes Diversity Effects on Ecosystem Processes Summary Review Questions Additional Reading Part III Patterns Chapter 13 Temporal Dynamics Introduction Fluctuations in Ecosystem Processes Interannual Variability Long-Term Change

13 Contents xiii Disturbance Conceptual Framework Disturbance Properties Succession Ecosystem Structure and Composition Carbon Balance Nutrient Cycling Trophic Dynamics Water and Energy Exchange Temporal Scaling of Ecological Processes Summary Review Questions Additional Reading Chapter 14 Landscape Heterogeneity and Ecosystem Dynamics Introduction Concepts of Landscape Heterogeneity Causes of Spatial Heterogeneity State Factors and Interactive Controls Community Processes and Legacies Disturbance Interactions Among Sources of Heterogeneity Patch Interactions on the Landscape Topographic and Land-Water Interactions Atmospheric Transfers Movement of Plants and Animals on the Landscape Disturbance Spread Human Land Use Change and Landscape Heterogeneity Extensification Intensification Spatial Heterogeneity and Scaling Summary Review Questions Additional Reading Part IV Integration Chapter 15 Global Biogeochemical Cycles Introduction The Global Carbon Cycle Long-Term Change in Atmospheric CO Anthropogenic Changes in the Carbon Cycle Terrestrial Sinks for CO The Global Methane Budget

14 xiv Contents The Global Nitrogen Cycle Anthropogenic Changes in the Nitrogen Cycle The Global Phosphorus Cycle Anthropogenic Changes in the Phosphorus Cycle The Global Sulfur Cycle The Global Water Cycle Anthropogenic Changes in the Water Cycle Consequences of Changes in the Water Cycle Summary Review Questions Additional Reading Chapter 16 Managing and Sustaining Ecosystems Introduction Ecosystem Concepts in Management Natural Variability Resilience and Stability State Factors and Interactive Controls Application of Ecosystem Knowledge in Management Forest Management Fisheries Management Ecosystem Restoration Management for Endangered Species Integrative Approaches to Ecosystem Management Ecosystem Management Integrated Conservation and Development Projects Valuation of Ecosystem Goods and Services Summary Review Questions Additional Reading Abbreviations Glossary References Index