| | TABLE OF CONTENTS: List of Tables xi
List of Figures xiii
Preface xix
Chapter One: Introduction 1
Chapter Two: The Hawaiian Islands as a Model Ecosystem 6
Model Systems 6
Microcosms and Well-studied Systems 8
A Brief Natural History 9
The Formation of the Hawaiian Islands 9
Determinants of Climate 15
Isolation 19
Evolution, Conservation, and Culture 20
Evolution and Speciation 20
Conservation Biology 22
Cultural Evolution 22
Chapter Three: Gradients in Environmental Factors, Gradients in Ecosystems 24
The State Factor Framework 24
Environmental Gradients as Model Systems 26
Temperature 27
Precipitation 29
The Mauna Loa Matrix 30
A Substrate Age Gradient across the Hawaiian Islands 31
Age Control 35
Climate History 35
Basic Features of the Gradient 39
Chapter Four: Patterns and Processes in Long-term Ecosystem Development 42
A Theory for Nutrient Dynamics during Ecosystem Development 42
Biogeochemical Processes on the Substrate Age Gradient 45
Soil P Pools 45
C and N Pools 45
Available Nutrients 46
Foliar Nutrients 49
Forest Productivity 51
Efficiencies of Resource Use 53
Decomposition and Nutrient Regeneration 59
Soil Organic Matter Turnover 66
Plant-Soil-Microbial Feedbacks 66
Chapter Five: Experimental Studies of Nutrient Limitation and the Regulation of Nutrient Cycling 70
Fertilization Experiments 71
Nutrient Limitation 74
Nutrient Availability and Plant-Soil-Microbial Feedback 78
Tissue Nutrient Concentrations 78
Productivity 78
Resource Efficiencies 79
Decomposition 84
Nutrient Regeneration 87
Controls of Plant-Soil-Microbial Feedback 87
Chapter Six: Nutrient Inputs to Hawaiian Ecosystems: Pathways, Rates, and Controls 92
Inputs of Elements 92
Weathering 93
Concepts and Definitions 93
Approaches 94
Element Inputs via Weathering 97
Atmospheric Inputs 98
Background 98
Deposition Measurements 100
Inputs of Water 101
Nitrogen Inputs 101
Influence of an Active Volcano 102
Inputs of Other Elements 103
Long-Distance Dust Transport 105
Background 105
Methods 105
Element Inputs 106
The Fate of Dust 107
Biological N Fixation 109
Background 109
Approach 109
Rates of Fixation 110
Other Inputs 110
Combined Inputs by All Known Pathways 111
Strontium Isotopes: A Direct Test of Input Pathways 112
Chloride and Sulfate 114
Mobile Cations 114
Silicon and Aluminum 116
Nitrogen and Phosphorus 117
Chapter Seven: Nutrient Outputs: Pathways, Controls, and Input-Output Budgets 121
Output Pathways 122
Leaching 122
N-Containing Trace Gases 124
Erosion 126
Other Pathways of Loss 126
Rates and Controls of N and P Losses 128
Input-Output Budgets 133
Budget Calculations 134
Using These Element Budgets 142
Chapter Eight: Issues and Opportunities 143
Interactions of Time Scales 143
An Exploratory Model 143
Supply versus Demand 144
Plant-Soil-Microbial Feedbacks 146
Sources and Sinks 148
Inputs and Outputs 150
Interactions across Scales 152
The Regulation of Nutrient Inputs and Outputs 154
Demand-Independent Pathways of Element Loss 155
Implications of Demand-Independent Nutrient Losses 159
Stoichiometry and Flexibility 160
Within-System Element Cycling 162
Inputs and Outputs 169
Biological N Fixation 173
Differences in Populations, Species, and Diversity 177
Biological Differences and Ecosystem Functioning 177
Diversity and Ecosystem Functioning 184
Three Final Points 188
References 191
Index 219
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