Biodiversity and Ecosystem Function

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1 Ernst-Detlef Schulze Harold A. Mooney (Eds.) Biodiversity and Ecosystem Function With 116 Figures, 7 Color Plates and 22 Tables Springer-Verlag Berlin Heidelberg New York London Paris Tokyo Hong Kong Barcelona Budapest

2 Contents Section A: Ecosystem Function 1 Biological Diversity and Terrestrial Ecosystem Biogeochemistry P. M. Vitousek and D. U. Hooper Introduction Semantics Biological Diversity and Biogeochemistry Experimental Tests Biogeographic Patterns Other Potential Effects of Plant Diversity on Biogeochemistry Conclusions 11 References 12 2 Biodiversity and Ecosystem Function in Agricultural Systems M. J. Swift and J. M. Anderson Introduction Characteristics of Agricultural Ecosystems Diversity and Complexity > Classification in Relation to Diversity and Complexity Sustainability : Productive Attributes of Low Number Multiple Cropping Systems Biodiversity and the Function of the Decomposer Subsystem Biodiversity in Relation to Function '.: Decomposer Diversity and Function in Agricultural Systems Interactions Between Plants and the Soil Biota Biodiversity and the Function of the Herbivore Subsystem Conclusions A Hypothesis of the Importance of Plant Diversity in Ecosystem Regulation 33

3 XIV Contents The Importance of Increasing Plant Species Number The Importance of Plant Species Composition Assessment of Long-Term Trends 38 References 38 3 Biodiversity and Interactions Within Pelagic Nutrient Cycling and Productivity C.E.W. Steinberg and W. Geller Introduction: Explanations to the Paradox of the Plankton Further Determinants of Biodiversity Plasticity and Cell Shape Turbulence Selection and Succession Descriptive Model of Plankton Succession Microbial Loop: Structure and Function Structure Structural Diversity Indices Ataxonomic Approach to Assess Ecosystem Stability. VJ Conclusions 60 References 61 Section B: Functional Groups 4 Functional Groups of Microorganisms O. Meyer Introduction Free-Living Components of the Soil Microbiota Metabolic Types of Bacteria The Role of Microorganisms in the Decomposition of Organic Material Cellulose Lignin Proteins, Peptides, and Amino Acids Pectin The Role of Microorganisms in the Biogeochemical Cycle of Nitrogen Nitrification Denitrification N 2 Fixation i The Role of Microorganisms in the Biogeochemical Cycle'of Sulfur The Oxidation of Reduced Sulfur Compounds Desulfurication 88

4 Contents XV 4.7 Conclusions 90 References Plant Traits and Adaptive Strategies: Their Role in Ecosystem Function O.T. Solbrig Introduction Schemes to Classify Plants on the Basis of Their Ecological Traits Single-Character Functional Classification of Vascular Plants Attempts to Classify Species Based on Their Overall Ecological Adaptability, Adaptive Strategies Optimization Plant Adaptive Strategies Why Optimality Criteria Are Not Always Sufficient Definition of Ecosystem Functional Properties The Meaning of Adaptive Strategy in a Complex, Nonlinear World Conclusions: The Importance of Diversity in a Nonequilibrium Situation 110 References Ill 6 Scaling from Species to Vegetation: The Usefulness of. Functional Groups Ch. Korner Introduction: What Are Functional Groups and Why Use Them? Selecting Functional Groups Narrow or Wide Grouping: The Dilemma of ; Experimental Safety and Ecological Applicability Grouping of Plant Species with Respect to Their Structural, Physiological, and Life Strategy Characteristics Life-Forms and Structures: The Morphotype Dry Matter Partitioning: Investment Type The Physiotype...: The Physiomorphotype Life Strategies The Spatial Definition of Functional Groups within Plant Communities Ecosystems: The Largest Functional Group Integration of Contrasting Levels of Complexity: A Compromise 131

5 XVI Contents 6.8 A Promising Tool: Using Functional Groups in Controlled Ecosystems Conclusions 136 References 137 Section C: Species Interaction 7 Evolution of Functional Groups in Basidiomycetes (Fungi) F. Oberwinkler Introduction What Are Fungi? Yeasts and Dimorphic Fungi Functional Fungal Groups Evolution of Fungal Parasites of Plants Evolution in Diverse Wood-Decaying Fungi Saprobic Fungi Evolution in Symbiontic Basidiomycetes Basidiolichens Mycorrhizae Diversity and Coevolutionary Trends in Septobasidiales Conclusions 162 References The Role of Parasites in Plant Populations and Communities J. J. Burdon Introduction The Diversity and Specialization of Parasites and Their Effects on the Fitness of the Host Plant Parasitic Plants" Fungal and Viral Pathogens The Hidden Effects of Parasite Attack - Changes in the Genetic Structure of Plant Populations 17Q 8.4 Parasite Attack as a Determinant of Ecosystem Structure Lessons from Exotic Pathogens and Severely Disturbed Natural Systems Evidence from Natural Parasite-Host Associations Conclusions 177 References 177

6 Contents XVII 9 Plant-Microbe Mutualisms and Community Structure D.J. Read Introduction Plant-Microbe Mutualisms in Grassland Communities Plant-Microbe Mutualisms in Savanna and Tropical Forest Communities Plant-Microbe Mutualisms in Boreal and Temperate Forest Communities Plant-Microbe Mutualisms in Heathland and Related Wetland Ecosystems The Role of Mutualisms in Successional Processes Conclusions 202 References The Evolution of Interactions and Diversity in Plant- Insect Systems: The Urophora-Eurytoma Food Web in Galls on Palearctic Cardueae H. Zwolfer and J. Arnold-Rinehart Introduction The Urophora Food Web General Ecological Characteristics of the Urophora- Eurytoma System Structure and Evolution of the Urophora Gall The Effect of the Gall Size on the Two Eurytoma spp Resource Exploitation, Interactions, and Evolution The Evolution of Diversity at the Herbivore Level of Plant-Insect Systems Host P'lants as Underexploited Resources Exploitation Strategies in the Urophora-Eurytoma System Interaction Patterns at the Second and Third Trophic Level: Evolutionary Adjustments in Food Webs \ Conclusions 226 References 227 Appendix 231 Section D: Community Interactions 11 Keystone Species W. J. Bond Introduction History of the Concept The Different Kinds of Keystone Species 238

7 XVIII Contents Keystone Predators Keystone Herbivores Keystone Pathogens Keystone Competitors Keystone Mutualists Earth-movers System Processes Abiotic Processes Summary of Types of Keystone Species Identifying Keystone Species Towards a General Protocol Which Keystone Species Are Vulnerable? Conclusions 249 References Redundancy in Ecosystems J. H. Lawton and V. K. Brown Introduction Evidence from the Fossil Record Patterns of Energy Flow, Biomass and the Structure of Food Webs Productivity and Biomass Food Webs Theoretical Models of Ecosystem Stability and Resilience Species Deletion Stability Possible Modelling Approaches Observations and Experiments on Real Systems Species Richness and Population Fluctuations Keystone Species Manipulation Experiments: General Considerations Manipulation Experiments: Examples ; Conclusions 266 References How Many Species Are Required for a Functional Ecosystem? F. I. Woodward Introduction : Ecosystems Species Diversity and Ecosystem Properties Introduction Species Enumerations and Ecosystem Functions The Inequality of Species in Ecosystem Function Species Diversity and Ecosystem Stability Species Numbers and Dynamics: Year-to-Year Averaging 279

8 Contents XIX Species Numbers and Dynamics: Species Feedbacks Species Diversity and Ecosystem Dynamics Introduction Experiments Modelling Conclusions 287 References Rare and Common Plants in Ecosystems, with Special Reference to the South-west Australian Flora J.S. Pate and S.D. Hopper Introduction Species Rareness or Commonness and Niche Specialization in Terms of Habitat and Nutritional Preference Fire as a Factor in Species Commonness and Rarity Strictly Serotinous Obligate Seeder Shrub or Tree Species Non-Serotinous or Partially Serotinous Obligate Seeder Shrub or Tree Species Obligate Seeder Species with Soil-Based Seed Reserves Resprouter Species of High Recruitment Potential Long-Lived, Clonally Reproducing Resprouter Species of Strictly Limited Recruitment Potential Fire Ephemerals Geophytes The Significance of Morphological and Physiological Variation to Commonness or Rareness of Species Evaluation of Commonness and Rareness in Related Taxonomic Groupings The Importance of Biotic Factors in Species Commonness or Rareness ; Genetic Correlates of Commonness and Rarity Conclusions : References Community Diversity and Succession: The Roles of Competition, Dispersal, and Habitat Modification D. Tilman Introduction Succession Environmental Constraints , Interspecific Trade-offs * Successional Theories Successional Dynamics and the Existing Species Pool Biotic Diversity 337

9 XX Contents Spatial Heterogeneity Local Recruitment Limitation Succession and Biodiversity Constraints, Trade-offs, and the Conservation of Biodiversity Conclusions 341 References 341 Section : Ecosystem Integrity 16 Biodiversity and the Balance of Nature S. L. Pimm What Biodiversity is Good for A History of Ecological Stability Controversy The Stability of Populations Resilience: The Example of Pest Outbreaks Year-to-Year Variability in Densities The Persistence of Communities Extinction Invasions Resistance to Change Conclusions " 358 'References Biodiversity and Function of Grazing Ecosystems S. J. McNaughton Introduction Intellectual Origins Conceptual Development An Individual Remark Theory and Empiricism Conceptual Definitions How to Test Tests Diversity and Productivity Diversity and Stability Stability of Species Composition to Drought and Grazing: Yellowstone Grazing Ecosystem Conclusions: Biodiversity and Ecosystem Function Biodiversity, Productivity, and Stability Biodiversity, System Perpetuation, and Global Change 380 References 382

10 Contents XXI 18 Resource Supply and Disturbance as-controls over Present and Future Plant Diversity S.E. Hobbie, D.B. Jensen, and F.S. Chapin, III Introduction Future Resource and Disturbance Regimes Plant Genetic Diversity Patterns of Genetic Diversity Land-Use Changes and Habitat Fragmentation Climatic Effects Resource Availability Plant Species Diversity Regional Patterns Latitudinal Patterns Paleoecological Patterns Future Changes Diversity of Plant Functional Groups General Considerations Control by Resourcesand Disturbance Types of Functional Groups Climatic Predictors Future Diversity Landscape Diversity Consequences of Changing Biodiversity Conclusions 404 References Ecosystem Stability, Competition, and Nutrient Cycling F. Berendse Introduction Stability of Model Ecosystems, Competition and the Loss of Diversity : Stabilizing Consequences of Competitive Interactions Effects of Organisms on Their Physical Environment Features Affecting Plant Fitness Under Different Nutrient Supply Conditions Consequences of the Different Effects of Plant Species on the Nutrient Cycle Conclusions 427 References t Modelling Biodiversity: Latitudinal Gradient of Forest Species Diversity Y. Iwasa, K. Sato, M. Kakita, and T. Kubo Introduction 433

11 XXII Contents 20.2 Hypotheses Explaining the Variation of Species Diversity Specialization of Resource Use Mode of Disturbance Smaller Opportunity for Competition Productivity Specific Herbivores and Pathogens Evolutionary/Ecological History Tree-byrTree Replacement: Finite Population Models Spatial Scale of Disturbance and Dispersal Inhibited Regeneration Temporal Fluctuation of Regeneration Ability Species Packing to Temporal Niches: Infinite Population Models Model Species Diversity Versus the Length of the Unfavorable Season Species Diversity Versus Niche Width Phenology of Coexisting Species Conclusions 447 References Functional Aspects of Landscape Diversity: A Bavarian Example E.-D. Schulze and P. Gerstberger Introduction Geology and Vegetation Land Use in Northeast Bavaria Hedgerows Grasslands Forests Conclusions 464 References 465 Section F: Industrial Analogy and Policy 22 Biodiversity Issues in Computing: A Study of Networked Computer Viruses C. Partridge and C. Malmstrom Introduction Stable Distributed Computer Systems Computer Viruses Duff's Virus The Morris Virus 472

12 Contents XXIII 22.4 Diversity and the Spread of a Networked Virus A Simple Mathematical Model Functional Diversity Species Diversity Conclusions,479 References Biodiversity and Policy Decisions L.F. Pitelka Introduction Conserving Biodiversity Global Climate Change Ecological Research and Policy Decisions Providing Policy-Relevant Research Results Conclusions 492 References 492 Conclusion 24 Ecosystem Function of Biodiversity: A Summary E.-D. Schulze and H.A. Mooney Introduction What Is an Ecosystem? The Regulation of Ecosystem Processes Are There Functional Groups? Determinants of Species Numbers Ecosystem Integrity Effects of Global Change on Land Use and Climate Conclusions 507 References ;, 509 Species Index 511 Subject Index 521

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