Principles of Terrestrial Ecosystem Ecology

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1 Principles of Terrestrial Ecosystem Ecology

3 F. Stuart Chapin, III Pamela A. Matson Peter M.Vitousek Principles of Terrestrial Ecosystem Ecology Second Edition Illustrated by Melissa C. Chapin

4 F. Stuart Chapin, III University of Alaska Fairbanks Institute of Arctic Biology Department of Biology & Wildlife Fairbanks, AK, USA Pamela A. Matson School of Earth Sciences Stanford University Stanford, CA, USA Peter M. Vitousek Department of Biological Sciences Stanford University Stanford, CA, USA ISBN e-isbn DOI / Springer New York Dordrecht Heidelberg London Library of Congress Control Number: Springer Science+Business Media, LLC 2011 All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer Science+Business Media, LLC, 233 Spring Street, New York, NY 10013, 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. Cover illustrations: Temperate forest in the eastern U.S. (North Carolina), showing a complex multi-layered canopy with sunflecks common in all canopy layers. Cover Photograph courtesy of Norm Christensen Printed on acid-free paper Springer is part of Springer Science+Business Media (

5 Preface Human activities are affecting the global environment in many 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 radically modify the interactions among forests, fields, streams, and oceans. This book is 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 ecosystem dynamics must underlie our analysis of both the consequences and the mitigation of human-induced changes. This book is intended to introduce the science of terrestrial ecosystem ecology to advanced undergraduate students, beginning graduate students, and practicing scientists from a wide array of disciplines. We define terrestrial ecosystem ecology to include freshwater ecosystems and their terrestrial matrix. We also include a description of marine ecosystems to provide a broader context for understanding terrestrial ecosystems and as a basis for Earth-System analysis. We provide access to some of the rapidly expanding literature in the many disciplines that contribute to ecosystem understanding. This second edition incorporates new material that accounts for both the substantial scientific advances in ecosystem ecology during the past decade, as well as the evolution of our own understanding. The first section of this 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 section 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 consider the important role of organisms in ecosystem processes through trophic interactions (feeding relationships), environmental effects, and disturbance. The third section 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 societies. Powerpoint lecture notes that include the illustrations in v

6 vi Preface this book are available on the web ( 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, our students from whom we have learned many of the important ideas that are presented, and Hal Mooney who was a co-author of the first edition. In addition, we thank the following individuals for their constructively critical review of chapters in this book: Richard Bardgett, Dan Binkley, Dave Bowling, Pep Canadell, Mimi Chapin, Doug Cost, Joe Craine, Wolfgang Cramer, Eric Davidson, Sandra Díaz, Jim Elser, Eugenie Euskirchen, Valerie Eviner, Noah Fierer, Jacques Finlay, Doug Frank, Mark Harmon, Sarah Hobbie, Dave Hooper, Bob Howarth, Ivan Janssens, Julia Jones, Bill Lauenroth, Joe McFadden, Dave McGuire, Sam McNaughton, Russ Monson, Deb Peters, Mary Power, Steve Running, Josh Schimel, Ted Schuur, Tim Seastedt, Mark Serreze, Phil Sollins, Bob Sterner, Kevin Trenberth, Dave Turner, Monica Turner, Diana Wall, John Walsh. We also thank Julio Betancourt, Scott Chambers, Norm Christensen, Greg Cortopassi, Steve Davis, Sandra Díaz, Jack Dykinga, Jim Elser, Jim Estes, Peter Franks, Mark Harmon, Al Levno, Mike Kenner, Alan Knapp, Aaryn Olsson, Roger Ruess, Dave Schindler, and David Tongway for the use of their photographs. We particularly thank Joe Craine and Dana Nossov for their constructive comments on the entire book. Fairbanks, AK, USA Stanford, CA, USA Stanford, CA, USA F. Stuart Chapin, III Pamela A. Matson Peter M. Vitousek

7 Contents Part I Context 1 The Ecosystem Concept... 3 Introduction... 3 A Focal Issue... 3 Overview of Ecosystem Ecology... 4 History of Ecosystem Ecology... 8 Ecosystem Structure and Functioning Ecosystem Processes Ecosystem Structure and Constraints Controls over Ecosystem Processes Human-Induced Ecosystem Change Human Impacts on Ecosystems Resilience and Threshold Changes Degradation in Ecosystem Services Summary Review Questions Additional Reading Earth s Climate System Introduction A Focal Issue Earth s Energy Budget The Atmospheric System Atmospheric Composition and Chemistry Atmospheric Structure Atmospheric Circulation The Ocean Ocean Structure Ocean Circulation Landform Effects on Climate Vegetation Influences on Climate Temporal Variability in Climate Long-Term Changes Anthropogenic Climate Change Interannual Climate Variability vii

8 viii Contents Seasonal and Daily Variation Storms and Weather Relationship of Climate to Ecosystem Distribution and Structure Summary Review Questions Additional Reading Geology, Soils, and Sediments Introduction A Focal Issue 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 Soil Physical Properties Soil Chemical Properties Summary Review Questions Additional Reading Part II Mechanisms 4 Water and Energy Balance Introduction A Focal Issue Surface Energy Balance Radiation Budget Partitioning of Absorbed Radiation Overview of Ecosystem Water Budgets Water Inputs to Ecosystems Water Movements Within Ecosystems Water Movement from the Canopy to the Soil Water Storage and Movement in the Soil Water Movement from Soil to Roots Water Movement Through Plants

9 Contents ix Water Losses from Ecosystems Evaporation from Wet Canopies Evapotranspiration from Dry Canopies Changes in Storage Runoff Summary Review Questions Additional Reading Carbon Inputs to Ecosystems Introduction A Focal Issue Overview of Carbon Inputs to Ecosystems Biochemistry of Photosynthesis Pelagic Photosynthesis Light Limitation CO 2 Supply Nutrient Limitation Pelagic GPP Living on the Edge: Streams and Shorelines Terrestrial Photosynthesis Photosynthetic Structure of Terrestrial Ecosystems C 4 Photosynthesis Crassulacean Acid Metabolism CO 2 Limitation Light Limitation Nitrogen Limitation and Photosynthetic Capacity Water Limitation Temperature Effects Pollutants Terrestrial GPP Canopy Processes Leaf Area Length of the Photosynthetic Season Satellite-Based Estimates of GPP Summary Review Questions Additional Reading Plant Carbon Budgets Introduction A Focal Issue Plant Respiration What Is NPP? Marine NPP Lake NPP Stream and River NPP

10 x Contents Terrestrial NPP Physiological Controls Over NPP Environmental and Species 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 Summary Review Questions Additional Reading Decomposition and Ecosystem Carbon Budgets Introduction A Focal Issue Overview of Decomposition and Ecosystem Carbon Balance Leaching of Litter Litter Fragmentation Chemical Alteration Fungi Bacteria and Archaea Soil Animals Temporal and Spatial Heterogeneity of Decomposition Temporal Pattern Vertical Distribution Factors Controlling Decomposition Litter Quality Rhizosphere Stimulation of Decomposition Microbial Community Composition and Enzymatic Capacity The Environment Soil Organic Matter Peat Accumulation and Trace Gas Emissions Heterotrophic Respiration Net Ecosystem Production (NEP) Net Ecosystem Carbon Balance Gaseous Carbon Fluxes Particulate Carbon Fluxes Dissolved Carbon Fluxes Stream Carbon Fluxes Stream Decomposition Stream Carbon Budgets Lake Carbon Fluxes Ocean Carbon Fluxes

11 Contents xi Carbon Exchange at the Global Scale Summary Review Questions Additional Reading Plant Nutrient Use Introduction A Focal Issue Overview Ocean Ecosystems Lake Ecosystems Rivers and Streams Terrestrial Ecosystems Nutrient Movement to the Root Diffusion Mass Flow Root Interception Nutrient Absorption Nutrient Supply Development of Root Length Mycorrhizae Nitrogen Fixation Root Absorption 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 Nutrient Cycling Introduction A Focal Point Overview of Nutrient Cycling Marine Nutrient Cycling Large-Scale Nutrient Cycles Estuaries Coastal Currents Lake Nutrient Cycling Stream Nutrient Cycling Nitrogen Inputs to Terrestrial Ecosystems Biological Nitrogen Fixation Nitrogen Deposition Internal Cycling of Nitrogen Overview of Mineralization Production and Fate of Dissolved Organic Nitrogen

12 xii Contents Production and Fate of Ammonium Production and Fate of Nitrate Temporal and Spatial Variability Pathways of Nitrogen Loss Gaseous Losses of Nitrogen Solution Losses Erosional Losses Other Element Cycles Phosphorus Sulfur Essential Cations Micronutrients and Nonessential Elements Nitrogen and Phosphorus Cycling in Agricultural Systems Summary Review Questions Additional Reading Trophic Dynamics Introduction A Focal Issue Overview of Trophic Dynamics Controls Over Energy Flow through Ecosystems Bottom-Up Controls Top-Down Controls Trophic Effects on Nutrient Cycling Ecological Efficiencies Trophic Efficiency and Energy Flow Consumption Efficiency Assimilation Efficiency Production Efficiency Food Chain Length Seasonal and Interannual Patterns Nutrient Transfers Detritus-Based Trophic Systems Integrated Food Webs Summary Review Questions Additional Reading Species Effects on Ecosystem Processes Introduction A Focal Issue Overview of Species Effects on Ecosystem Processes Effect Functional Types Species Effects on Biogeochemistry Species Effects on Biophysical Processes Species Effects on Trophic Interactions Species Effects on Disturbance Regime Response Functional Types

13 Contents xiii Integrating the Effects of Traits on Ecosystems Functional Matrix of Multiple Traits Linkages Between Response and Effect Traits Diversity as Insurance Species Interactions and Ecosystem Processes Summary Review Questions Additional Reading Part III Patterns 12 Temporal Dynamics Introduction A Focal Issue Ecosystem Resilience and Change Alternative Stable States Resilience and Thresholds Disturbance Conceptual Framework Impact of a Disturbance Event Recovery and Renewal after Disturbance Disturbance Regime Succession Ecosystem Structure and Composition Water and Energy Exchange Carbon Balance Nutrient Cycling Trophic Dynamics Temporal Scaling of Ecological Processes Summary Review Questions Additional Reading Landscape Heterogeneity and Ecosystem Dynamics Introduction A Focal Issue Concepts of Landscape Heterogeneity Causes of Spatial Heterogeneity Detection and Analysis 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

14 xiv Contents Human Land-Use Change and Landscape Heterogeneity Extensification Intensification Extrapolation to Larger Scales Summary Review Questions Additional Reading Part IV Integration 14 Changes in the Earth System Introduction A Focal Issue Human Drivers of Change The Global Water Cycle Water Pools and Fluxes Anthropogenic Changes in the Water Cycle Consequences of Changes in the Water Cycle The Global Carbon Cycle Carbon Pools and Fluxes Changes in Atmospheric CO Marine Sinks for CO Terrestrial Sinks for CO CO 2 Effects on Climate The Global Methane Budget The Global Nitrogen Cycle Nitrogen Pools and Fluxes Anthropogenic Changes in the Nitrogen Cycle The Global Phosphorus Cycle Phosphorus Pools and Fluxes Anthropogenic Changes in the Phosphorus Cycle The Global Sulfur Cycle Summary Review Questions Additional Reading Managing and Sustaining Ecosystems Introduction A Focal Issue Sustaining Social Ecological Systems Sustainability Ecological Dimensions of Sustainability Conceptual Framework for Ecosystem Management Sustaining Soil Resources Sustaining Biodiversity Sustaining Variability and Resilience Applying Ecosystem Principles to Management Forest Management Fisheries Management

15 Contents xv Ecosystem Renewal Management for Endangered Species Socioeconomic Contexts of Ecosystem Management Meeting Human Needs and Wants Managing Flows of Ecosystem Services Addressing Political Realities Innovation and Adaptive Management Sustainable Development: Social Ecological Transformation Summary Review Questions Additional Reading Abbreviations Glossary References Index

Principles of Terrestrial Ecosystem Ecology

E Stuart Chapin III Pamela A. Matson Harold A. Mooney Principles of Terrestrial Ecosystem Ecology Illustrated by Melissa C. Chapin With 199 Illustrations Teehnische Un.fversitSt Darmstadt FACHBEREIGH 10