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The Ecology of Plants
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Table of Contents

1. The Science of Plant Ecology Ecology as a Science The Genesis of Scientific Knowledge Objectivity, Subjectivity, Choice, and Chance in Scientific Research Experiments: The Heart of Research Testing Theories Specific Results versus General Understanding Science and Other Ways of Knowing, Revisited Scale and Heterogeneity The Structure and History of Plant Ecology Questions for Further Study Additional Readings PART I. THE INDIVIDUAL AND ITS ENVIRONMENT 2. Photosynthesis and the Light Environment The Process of Photosynthesis Photosynthetic Rates Limitations Caused by Light Levels Limitations on Carbon Uptake Variation in Photosynthetic Rates Within and Between Habitats The Three Photosynthetic Pathways C3 Photosynthesis C4 Photosynthesis Box 2A. Photorespiration Box 2B. Stable Isotopes and Photosynthesis Crassulacean Acid Metabolism (CAM Photosynthesis) Evolution of the Three Photosynthetic Pathways Phylogeny of the Photosynthetic Pathways Photosynthesis through Evolutionary Time Growth Form, Phenology, and Distribution of C3, C4, and CAM Plants Growth Forms and Habitats Phenology Geographic Distributions Adaptations to the Light Environment Sun and Shade Leaves Species' Adaptations to High-Light and Low-Light Habitats Box 2C Leaf Iridescence and Structural Coloration Do Sun and Shade Adaptations Exist Within Species? Day Length: Responses and Adaptations Summary Questions for Further Study Additional Readings 3. Water Relations and Energy Balance Adapting to Life on Land Water Potential The Soil-Plant-Atmosphere Continuum Box 3A. Measuring Photosynthesis, Transpiration, and Water Potential Transpiration and the Control of Water Loss Strategies for Coping with Different Water Availability Conditions Water Use Efficiency Whole-Plant Adaptations to Low Water Availability Physiological Adaptations Anatomical and Morphological Adaptations The Energy Balance of Leaves Radiant Energy Box 3B. Why the Sky Is Blue and the Setting Sun Is Red Conduction and Convection Latent Heat Exchange Putting It All Together: Leaf and Whole-Plant Temperature Adaptations to Extreme Temperature Regimes Summary Questions for Further Study Additional Readings 4. Soils, Mineral Nutrition, and Belowground Interactions Soil Composition and Structure Soil Texture Soil pH Horizons and Profiles Origins and Classification Organic Matter and the Role of Organisms Water Movement within Soils Plant Mineral Nutrition The Stoichiometry of Nutrients Nitrogen in Plants and Soils Biological Nitrogen Fixation Box 4A. Symbioses and Mutualisms Phosphorus in Soils Nutrient Use Efficiency Leaf Life Span and Evergreen versus Deciduous Leaves Mycorrhizae Major Groups of Mycorrhizae The Role of Mycorrhizae in Plant Phosphorus Nutrition Other Functions of Mycorrhizae Orchids and Their Mycorrhizal Associations Mutualism or Parasitism? Effects of Mycorrhizae on Plant Interactions Summary Questions for Further Study Additional Readings PART II. POPULATIONS AND EVOLUTION 5. Population Structure, Growth, and Decline Some Issues in the Study of Plant Population Growth Population Structure Some Population Structure Issues Specific to Plants Sources of Population Structure Studying Population Growth and Decline Life Cycle Graphs Box 5A. Life Table Calculations Box 5B. Borrowing the Mark-Recapture Method from Animal Ecology Box 5C. Constructing Matrix Models Matrix Models Box 5D. Demography of an Endangered Cactus Box 5E. Multiplying a Population Vector by a Matrix Analyzing Matrix Models But Real Plants Live in Variable Environments Lifetime Reproduction: The Net Reproductive Rate Reproductive Value: The Contribution of Each Stage to Population Growth Box 5F. Reproductive Value Box 5G. How Do Changes in the Transition Probabilities Affect the Population Growth Rate? Sensitivity and Elasticity Life Table Response Experiments Age and Stage, Revisited Other Approaches to Modeling Plant Demography Demographic Studies of Long-Lived Plants Random Variation in Population Growth and Decline Causes of Random Variation Long-Term Growth Rates Studying Variable Population Growth Summary Questions for Further Study Additional Readings 6. Evolutionary Processes and Outcomes Natural Selection Variation and Natural Selection The Factors Necessary for Natural Selection Heritability Resemblance among Relatives Partitioning Phenotypic Variation Box 6A. A Simple Genetic System and the Resemblance of Relatives Genotype-Environment Interactions Gene-Environment Covariation Patterns of Adaptation Heavy-Metal Tolerance Adaptive Plasticity Levels of Selection Other Evolutionary Processes Processes that Increase Variation Processes that Decrease Variation Variation among Populations Ecotypes Speciation Adaptation and Speciation through Hybridization Summary Questions for Further Study Additional Readings 7. Growth and Reproduction of Individuals Plant Growth Ecology of Growth Plant Architecture and Light Interception Growth of Clonal Plants Plant Reproduction Vegetative Reproduction Seeds Produced Asexually Sexual Life Cycles of Plants Pollination Ecology Wind Pollination Attracting Animal Visitors: Visual Displays Attracting Animal Visitors: Floral Odors and Acoustic Guides Limiting Unwanted Visits Pollination Syndromes Box 7A. Specialized Plants and Pollinators Aquatic Plants and Pollination Box 7B. Some Complex Plant-Pollinator Interactions Who Mates with Whom? Plant Gender Box 7C. Pollination Experiments Competition for Pollinators and among Pollen Grains Pollen Dispersal and Its Consequences Assortative Mating Frequency-Dependent Selection Factors that Shape Plant Mating Systems Applications of Pollination and Mating System Ecology The Ecology of Fruits and Seeds Seed Dispersal Patterns Seed Banks Summary Questions for Further Study Additional Readings 8. Plant Life Histories Size and Number of Seeds Life History Strategies Life Span r- and K-selection Grime's Triangular Model Demographic Life History Theory Reproductive Allocation Difficulties in Measuring Trade-Offs Variation among Years Consequences of Variable Environments Seed Germination Masting Phenology: Within-Year Schedules of Growth and Reproduction Vegetative Phenology Reproductive Phenology: Abiotic Factors Reproductive Phenology: Biotic Factors Summary Questions for Further Study Additional Readings PART III. COMMUNITIES AND THEIR CAUSES 9. Community Properties What Is a Community? The History of a Controversy Box 9A. Communities, Taxa, Guilds, and Functional Groups A Modern Perspective on the Issues in Contention Are Communities Real? Box 9B. A Deeper Look at Some Definitions: Abiotic Factors and Emergent Properties Describing Communities Species Richness Diversity, Evenness, and Dominance Sampling Methods and Parameters for Describing Community Composition Physiognomy Long-Term Studies Summary Box 9C. The Long-Term Ecological Research Network Questions for Further Study Additional Readings 10. Competition and Other Interactions among Plants Competition at the Level of Individuals Seedlings: Density, Size, Inequality, and Timing of Emergence Seedlings: Density and Mortality Mechanisms of Competition for Resources Size and Resource Competition Experimental Methods for Studying Competition Greenhouse and Garden Experiments Box 10A. How Competition Is Measured, and Why That Matters Field Experiments From Interspecific Competition to Allelopathy to Facilitation Trade-offs and Strategies Competitive Hierarchies Allelopathy Facilitation Modeling Competition and Coexistence Equilibrium Models Nonequilibrium Approaches to Modeling Competition Effects of Competition on Species Coexistence and Community Composition Competition along Environmental Gradients Conceptual Models of Competition in Habitats with Differing Productivities Experimental Evidence Evidence from Research Syntheses Resolution of Differing Results Summary Questions for Further Study Additional Readings 11. Herbivory and Plant-Pathogen Interactions Herbivory at the Level of Individuals Herbivory and Plant Populations Herbivory and Spatial Distribution of Plants Granivory Biological Control Effects of Herbivory at the Community Level Consequences of Herbivore Behavior Apparent Competition Introduced and Domesticated Herbivores Effects of Native Herbivores Generality Plant Defenses against Herbivory Physical Defenses Plant Secondary Chemistry Constitutive versus Induced Defenses Evolutionary Consequences of Plant-Herbivore Interactions Parasitic Plants Pathogens Effects of Disease on Individual Plants BOX 11A. Effects of Plant Disease on Humans: Potato Blight and the Irish Potato Famine Physiological and Evolutionary Responses to Pathogens Effects of Pathogens at the Population and Community Level More Complex Interactions Summary Questions for Further Study Additional Readings 12. Disturbance and Succession Theories of the Mechanisms of Succession Disturbance Gaps Fire Wind Water Animals Earthquakes and Volcanoes Disease Humans Colonization Determining the Nature of Succession Interaction between Methodology and Understanding Mechanisms Responsible for Successional Change The Predictability of Succession Community Restoration Primary Succession Climax Revisited Summary Questions for Further Study Additional Readings 13. Local Abundance, Diversity, and Rarity Dominance Are Dominant Species Competitively Superior? Abundance Curves Rarity and Commonness The Nature of Rarity Patterns of Rarity and Commonness Causes of Rarity and Commonness Invasive Species and Community Susceptibility to Invasion Why Do Some Species Become Invasive? What Makes a Community Susceptible to Invasion? Abundance and Community Structure Productivity and Diversity Niche Differentiation, Environmental Heterogeneity, and Diversity Gaps, Disturbance, and Diversity Effects of Increasing Diversity Testing the Effects of Diversity on Ecosystems Diversity and Stability Regional Processes Summary Questions for Further Study Additional Readings PART IV. ECOSYSTEMS AND LANDSCAPES 14. Ecosystem Processes Biogeochemical Cycles: Quantifying Pools and Fluxes The Global Water Cycle Carbon in Ecosystems Productivity Methods for Estimating Productivity Decomposition and Soil Food Webs Carbon Storage Models of Ecosystem Carbon Cycles Nitrogen and the Nitrogen Cycle at Ecosystem and Global Levels Nitrogen Fixation Other Sources of Nitrogen Input to Living Organisms Nitrogen Mineralization Denitrification and Leaching of Nitrogen Decomposition Rates and Nitrogen Immobilization Plant Uptake of Nitrogen Phosphorus in Terrestrial Ecosystems Ecosystem Nutrient Cycling and Plant Diversity Ecosystem Processes for Some Other Elements Sulfur Calcium Box 14A. Serpentine Soils Summary Questions for Further Study Additional Readings 15. Communities in Landscapes Comparing Communities Non-numerical Techniques Univariate Techniques Multivariate Techniques Landscape Patterns Ordination: Describing Patterns Determining Causes of Patterns Types of Data Classification Box 15A. Differentiating Vegetation Based on Spectral Quality Views on Continuous versus Discrete Landscapes Landscape Diversity Differentiation Diversity Pattern Diversity Summary Questions for Further Study Additional Readings 16. Landscape Ecology Spatial Patterns Six Types of Species-Area Curves Defining Patches Quantifying Patch Characteristics and Interrelationships The Effects of Spatial Patterns on Ecological Processes Scale Definitions and Concepts Process and Scale Spatial and Ecological Scale Quantifying Aspects of Spatial Pattern and Scale Toward a Theoretical Basis for Landscape Patterns: Island Biogeography Theory Metapopulation Theory Box 16A. Metapopulation Models Metapopulation Patterns Species-Time-Area Relationships Landscape Ecology and Conservation Reserve Design Fragmentation Edges, Connectivity, and Nestedness Summary Questions for Further Study Additional Readings PART V. GLOBAL PATTERNS AND PROCESSES 17. Climate and Physiognomy Climate and Weather Temperature Short-Term Variation in Radiation and Temperature Long-Term Cycles Precipitation Global Patterns Box 17A. The Coriolis Effect Continental-Scale Patterns Seasonal Variation in Precipitation The El Nino Southern Oscillation Predictability and Long-Term Change Plant Physiognomy across the Globe Forests Tree Line Grasslands and Woodlands Shrublands and Deserts Summary Questions for Further Study Additional Readings 18. Biomes Categorizing Vegetation Converging Biomes and Convergent Evolution Moist Tropical Forests Tropical Rainforest Tropical Montane Forest Seasonal Tropical Forests and Woodlands Tropical Deciduous Forest Thorn Forest Tropical Woodland Temperate Deciduous Forest Other Temperate Forests and Woodlands Temperate Rainforest Temperate Evergreen Forest Temperate Woodland Taiga Temperate Shrubland Grasslands Temperate Grassland Tropical Savanna Deserts Hot Desert Cold Desert Alpine and Arctic Vegetation Alpine Grassland and Shrubland Tundra Summary Questions for Study and Thought Additional Readings 19. Regional and Global Diversity Large-Scale Patterns of Species Richness General Factors Affecting Diversity Levels of Explanation Null Models The Importance of Available Energy Contributions of ?, ?, and ? Diversity Diversity along Ecological Gradients Productivity and Scale Diversity along Latitudinal Gradients An Array of Explanations The Role of ? Diversity Continental Differences Other Geographic Patterns Species Diversity and Patterns of Overlap Endemism, Centers of Diversification, and Isolation Relationships between Regional and Local Diversity Box 19A. The Fynbos and the Cape Region of Africa Noisy Data and Limits to Methodology Summary Questions for Further Study Additional Readings 20. Paleoecology The Paleozoic Era The Mesozoic Era The Dominance of Gymnosperms The Breakup of Pangaea and the Rise of the Angiosperms The Cretaceous-Tertiary (K-T) Boundary The Cenozoic Era Paleoecology Methods The Recent Past At the Glacial Maximum Glacial Retreat Climatic Fluctuations in the Recent Past Summary Questions for Further Study Additional Readings 21. Global Change: Humans and Plants Carbon and Plant-Atmosphere Interactions The Global Carbon Cycle Direct Effects of Increasing CO2 on Plants Anthropogenic Global Climate Change The Greenhouse Effect Global Climate Change: Evidence Global Climate Change: Predictions Box 21A. Modeling Climate Biotic Consequences of Climate Change Anthropogenic Effects on the Global Carbon Cycle Deforestation Fossil Fuel Combustion Box 21B. Daily Human Activities and CO2 Generation Acid Precipitation and Nitrogen Deposition Declining Global Biodiversity and Its Causes Habitat Fragmentation and Loss Other Threats to Rare and Common Species in a Range of Communities Invasive Species as Threats to Biodiversity Human Populations and Land Use Patterns A Ray of Hope? Summary Questions for Further Study Additional Readings Appendix: A Statistics Primer Glossary Photo Credits Literature Cited Index

About the Author

Jessica Gurevitch is Professor in the Department of Ecology and Evolution at the State University of New York at Stony Brook. She earned a B.S. in Biological Sciences (Ecology, Evolution, and Systematics) at Cornell University, and a Ph.D. in Ecology and Evolutionary Biology at the University of Arizona. Research projects currently underway include empirical, experimental and modeling approaches to studying pine demography, and experimental studies of plant invasions in forests. Samuel M. Scheiner is currently working for the U.S. federal government. Previously he was on the faculties of Arizona State University, Northern Illinois University, and the University of Arizona. He earned his B.A., M.S., and Ph.D., all in Biology, from the University of Chicago. Dr. Scheiner's research has been involved equally with plants and animals, the theoretical and the empirical, including population genetics, physiological ecology, population biology, and macroecology. Currently, his research centers on three issues: phenotypic plasticity, measuring natural selection, and both theoretical and empirical work on large-scale patterns of species diversity. Gordon A. Fox is Associate Professor of Biology at the University of South Florida. He earned a B.A. in History at the University of California, Berkeley and a Ph.D. in Ecology and Evolutionary Biology at the University of Arizona, where he researched the ecology, genetics, and evolution of annual plants in the Sonoran and Chihuahuan Deserts. His postdoctoral work at the University of California, Davis concentrated on theoretical population genetics. His current research interests include: population dynamics of plants in a stochastic world; the ecology of populations in fire-prone regions; the ecology and evolution of reproductive timing in plants; and applications of population ecology to conservation of endangered plants.

Reviews

"This is the only volume I have seen that contains a good treatment of evolutionary and physiological ecology, as well as more traditional topics such as competition and herbivory, vegetation types, and biomes. Overall, this useful book, beautifully laid out and well written, fills an important gap in the educational literature on plant ecology. It is up to date and quantitatively sophisticated, and it provides a nice balance between presentation of facts and concepts and narration of evocative examples. I plan to continue to use this volume in my course and recommend it highly."--Ingrid M. Parker, The Quarterly Review of Biology "Overall, The Ecology of Plants provides a useful and complete tool to teach plant ecology to undergraduate students. Therefore, I strongly recommend it."--Stephane Boudreau, Ecoscience

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