陆地生态系统生态学原理(Principles of Terrestrial Ecosystem Ecology).pdf

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Cover

Principles of Terrestrial Ecosystem Ecology, 2nd Edition

ISBN 9781441995032

Preface

Contents

Part I: Context

1: The Ecosystem Concept

Introduction

A Focal Issue

Overview of Ecosystem Ecology

History of Ecosystem Ecology

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

2: 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

Seasonal and Daily Variation

Storms and Weather

Relationship of Climate to Ecosystem Distribution and Structure

Summary

Review Questions

Additional Reading

3: 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

Box 4.1 The Energetics of Water Movement

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

Roots

Box 4.2 Tracing Water Flow Through Ecosystems

Stems

Leaves

Water Losses from Ecosystems

Evaporation from Wet Canopies

Evapotranspiration from Dry Canopies

Changes in Storage

Runoff

Summary

Review Questions

Additional Reading

5: 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

Box 5.1 Carbon Isotopes

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

6: Plant Carbon Budgets

Introduction

A Focal Issue

Plant Respiration

What Is NPP?

Marine NPP

Lake NPP

Stream and River NPP

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

7: 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

Box 7.1 Isotopes and Soil Carbon Turnover

Factors Controlling Decomposition

Litter Quality

Rhizosphere Stimulation of Decomposition

Microbial Community Composition and Enzymatic Capacity

The Environment

Moisture

Temperature

Soil Organic Matter

Soil Properties

Soil Disturbance

Humus Formation

Peat Accumulation and Trace Gas Emissions

Heterotrophic Respiration

Net Ecosystem Production (NEP)

Box 7.2 Measuring Carbon Fluxes of Ecosystems and Regions

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

Carbon Exchange at the Global Scale

Summary

Review Questions

Additional Reading

8: 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

9: 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

Groups of Nitrogen Fixers

Causes of Variation in 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

Atmospheric Roles of Nitrogen Gases

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

10: 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

11: Species Effects on Ecosystem Processes

Introduction

A Focal Issue

Overview of Species Effects on Ecosystem Processes

Effect Functional Types

Species Effects on Biogeochemistry

Nutrient Supply

Nutrient Turnover

Species Effects on Biophysical Processes

Species Effects on Trophic Interactions

Species Effects on Disturbance Regime

Response Functional Types

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

Sources of Resilience

Limits to Resilience

Thresholds and Regime Shifts

Box 12.1 Resilience and Regime Shifts

Disturbance

Conceptual Framework

Impact of a Disturbance Event

Recovery and Renewal after Disturbance

Disturbance Regime

Succession

Ecosystem Structure and Composition

Primary Succession

Secondary Succession

Water and Energy Exchange

Carbon Balance

Primary Succession

Secondary Succession

Nutrient Cycling

Primary Succession

Secondary Succession

Trophic Dynamics

Temporal Scaling of Ecological Processes

Summary

Review Questions

Additional Reading

13: 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

Human Land-Use Change and Landscape Heterogeneity

Extensification

Intensification

Extrapolation to Larger Scales

Box 13.1 Spatial Scaling Through Ecological Modeling

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 2

Marine Sinks for CO 2

Box 14.1 Partitioning of Carbon Uptake Between the Land and Ocean

Terrestrial Sinks for CO 2

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

15: Managing and Sustaining Ecosystems

Introduction

A Focal Issue

Sustaining Social–Ecological Systems

Box 15.1 Social-Ecological Interactions and the Flooding of New Orleans

Sustainability

Ecological Dimensions of Sustainability

Box 15.2 Assessing Tradeoffs Among Ecosystem Services: Hydropower Versus Conservation in New Zealand

Box 15.3 Water Purification for New York City

Conceptual Framework for Ecosystem Management

Sustaining Soil Resources

Sustaining Biodiversity

Sustaining Variability and Resilience

Applying Ecosystem Principles to Management

Forest Management

Fisheries Management

Ecosystem Renewal

Box 15.4 Everglades Restoration Study

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

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

Pamela A. Matson School of Earth Sciences Stanford University Stanford, CA, USA matson@stanford.edu F. Stuart Chapin, III University of Alaska Fairbanks Institute of Arctic Biology Department of Biology & Wildlife Fairbanks, AK, USA terry.chapin@alaska.edu Peter M. Vitousek Department of Biological Sciences Stanford University Stanford, CA, USA vitousek@stanford.edu ISBN 978-1-4419-9503-2 DOI 10.1007/978-1-4419-9504-9 Springer New York Dordrecht Heidelberg London e-ISBN 978-1-4419-9504-9 Library of Congress Control Number: 2011935993 © 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 (www.springer.com)

Preface Human activities are affecting the global environment in many ways, with numerous direct and indirect effects on ecosystems. The climate and atmo- spheric composition of Earth are changing rapidly. Humans have directly modified half of the ice-free terrestrial surface and use 40% of terrestrial pro- duction. 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 environ- mental and biotic changes. We believe that an understanding of ecosystem dynamics must underlie our analysis of both the consequences and the mitiga- tion 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 terres- trial 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 sub- stantial 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 eco- system 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 distur- bance. 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

vi Preface this book are available on the web (http://terrychapin.org/) as supplementary material. Many people have contributed to the development of this book. We particu- larly 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 addi- tion, 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

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 ........................................................................ 2 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 ................................................. 11 11 12 13 17 17 20 21 21 22 22 23 23 23 23 26 26 28 30 35 35 36 38 40 41 41 44 45 vii

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