Part I Research Programme and Study Area
1. General Concept of the Research Programme and Methodology
of Investigations ............................................ 3
Otto Fränzle, Ludger Kappen, Hans-Peter Blume, Klaus
Dierssen, Ulrich Irmler, Winfrid Kluge, Uwe Schleuß,
and Joachim Schrautzer
1.1. Introduction ............................................ 3
1.2. History and General Concept of the Research Programme ... 3
1.2.1. Conceptual Background and Organizational
Framework ........................................ 3
1.2.2. Selection of Representative Study Areas and
Sites ............................................ 5
1.2.3. General and Specific Research Objectives
of the Bönhoved Project ......................... 7
1.2.4. Plan of Research ................................ 10
1.2.5. Organizational Structure of the Project ......... 11
1.3. Methodology of Investigations .......................... 13
1.3.1. Instruments and Methods of Meteorological
and Hydrological Observations ................... 15
1.3.2. Soil Surveys and Analyses ....................... 18
1.3.3. Biocoenotic Investigations ...................... 19
1.3.4. Element Fluxes in Air, Water and the Soil-
Vegetation Complex .............................. 22
1.3.5. Ecological Modelling ................................. 24
2. Ecological Setting of the Study Area ........................ 29
Hans-Peter Blume, Otto Fränzle, Georg Hörmann, Ulrich
Irmler, Winfried Kluge, Uwe Schleuß, and Joachim
Schrautzer
2.1. Introduction ........................................... 29
2.2. Geological Setting ..................................... 29
2.2.1. Rocks and Relief Features ....................... 29
2.2.2. Climate ......................................... 30
2.2.3. Soil Pattern .................................... 32
2.2.4. Drainage Systems ................................ 33
2.2.5. Vegetation ...................................... 40
2.2.6. Development and Socio-Economic Structure
of the Cultural Landscape ....................... 42
2.3. Site and Community Characteristics in Catenary
Analysis ............................................... 44
2.3.1. Beech Forest .................................... 45
2.3.2. Mixed Forest .................................... 48
2.3.3. Alder Carrs ..................................... 50
2.3.4. Agroecosystems .................................. 51
2.3.5. Grassland without Groundwater Contact ........... 53
2.3.6. Grassland with Groundwater Contact .............. 54
2.3.7. Hedgerows ....................................... 55
2.3.8. Reed Swamps ..................................... 56
Part II Structure and Function of Ecosystems in a Complex
Landscape
3. Ecophysiological Key Processes in Agricultural and Forest
Ecosystems .................................................. 61
Oliver Dilly, Christiane Eschenbach, Werner L. Kutsch,
Ludger Kappen, and Jean Charles Munch
3.1. Introduction ........................................... 61
3.2. Control of Carbon Input ................................ 62
3.2.1. Leaf Gas Exchange ............................... 62
3.2.2. Upscaling to Canopy Level ....................... 69
3.3. Partitioning of Carbon ................................. 69
3.4. Respiration as Carbon Loss from the System ............. 71
3.5. Decomposition and Mineralization of Organic С .......... 73
3.6. Structure and Activity of Soil Microbiota .............. 73
3.7. Discussion ............................................. 78
3.8. Conclusions ............................................ 80
4. Carbon and Energy Balances of Different Ecosystems
and Ecosystem Complexes of the Bornhöved Lake District ...... 83
Werner L. Kutsch, Georg Hörmann, and Ludger Kappen
4.1. Introduction ........................................... 83
4.2. Carbon and Energy Fluxes and Balances at the
Ecosystem Scale ........................................ 83
4.2.1. Abiotic Ecosystem Energy Balances ............... 83
4.2.2. Carbon Fixation, Primary Production and
Biological Energy Consumption ................... 85
4.2.3. Soil Carbon Balances ............................ 85
4.2.4. Net Ecosystem Carbon Balances ................... 90
4.2.5. Extended Energy Balances at the Patch Scale ..... 90
4.3. Farm Gate Balances ..................................... 92
4.4. Energy Fluxes at the Landscape Scale ................... 95
4.5. Conclusions ............................................ 98
5. Water Relations at Different Scales ........................ 101
Georg Hörmann, Matthias Herbst, and Christiane
Eschenbach
5.1. Introduction .......................................... 101
5.2. Water Relations at the Plant Leaf Scale ............... 101
5.2.1. Diurnal Course of Leaf Water Relations ......... 102
5.2.2. Dependence of Leaf Conductance and
Transpiration on Irradiance and Saturation
Deficit ........................................ 105
5.2.3. Interrelations between Leaf Water Potential,
Leaf Conductance and Transpiration ............. 106
5.2.4. Acclimation of Leaves to Environmental
Conditions ..................................... 107
5.2.5. Measurement and Parameterization of
Interception and Soil Evaporation .............. 107
5.3. Water Turnover at the Stand Level ..................... 109
5.3.1. Evaporation and Advective Energy Supply ........ 112
5.3.2. Comparison of Water Vapour Flux Measurements
with Different Modelling Approaches ............ 114
5.4. Conclusions ........................................... 117
6. Site-Related Biocoenotic Dynamics .......................... 119
Ulrich Irmler, Oliver Dilly, Joachim Schrautzer, and
Klaus Dierssen
6.1. Introduction .......................................... 119
6.2. Vertical Distribution Patterns of Fauna and
Microbiota in Beech and Alder Stands .................. 120
6.2.1. Vertical Stratification in the Soil Layer ...... 120
6.2.2. Vertical Stratification of Animals in the
Vegetation Cover ............................... 120
6.3. Spatial Patterns of Microbiota, Vegetation and Fauna
in Beech Forest ....................................... 124
6.3.1. Microbiota and Decomposition ................... 124
6.3.2. Vegetation ..................................... 125
6.3.3. Fauna .......................................... 128
6.4. Seasonal Dynamics ..................................... 131
6.4.1. Seasonal Changes in the Animal Community ....... 131
6.4.2. Seasonal Change of Habitats .................... 134
6.4.3. Seasonal Changes of Food Resources ............. 135
6.5. Long-Term Dynamics .................................... 138
6.5.1. Effects of Climatic Change ..................... 138
6.5.2. Influence of Temperature on the Generation
Cycle .......................................... 140
6.5.3. Succession from Grassland to Alder Carrs ....... 140
6.5.4. Changes during Decomposition Processes ......... 142
6.6. Conclusions ........................................... 145
7. Biocoenotic Interactions between Different Ecotopes ........ 147
Ulrich Irmler, Franz Hölker, Hans-Werner Pfeiffer,
Walter Nellen, and Hauke Reuter
7.1. Introduction .......................................... 147
7.2. Interactions between Different Ecosystems ............. 148
7.2.1. Interactions between Terrestrial Ecosystems .... 148
7.2.2. Interactions between Terrestrial and Aquatic
Ecosystems ..................................... 153
7.2.3. Interactions within Aquatic Systems ............ 154
7.3. Modelling Species Interactions between Habitats ....... 161
7.3.1. Modelling Interactions between Terrestrial
Habitats ....................................... 161
7.3.2. Modelling Interactions between Aquatic
Habitats ....................................... 164
7.4. Conclusions ........................................... 167
8. Element Fluxes in Atmosphere, Vegetation and Soil .......... 169
Otto Fränzle and Claus-Georg Schimming
8.1. Introduction .......................................... 169
8.2. Atmospheric Deposition and Leaching Processes
of the Vegetation Cover ............................... 169
8.2.1. Medium-Scale Deposition Patterns of the
Study Area ..................................... 169
8.2.2. Small-Scale Atmospheric Deposition, Canopy
Throughfall and Litterfall of Beech Stands ..... 170
8.3. Element Cadasters and Nutrient Fluxes in Arenic
Umbrisols of Beech Stands and Eutri-Cambic
Arenosols of Arable Land .............................. 176
8.3.1. Element Cadasters of Soils ..................... 176
8.3.2. Bacterial Populations and Degradation
of Soil Organic Matter ......................... 177
8.3.3. Annual Course of Element Concentrations in
Field Eutri-Brunic Arenosols and Forest
Arenic Umbrisols ............................... 179
8.3.4. Long-Term Element Budgets of Forest Arenic
Umbrisols and Eutri-Brunic Arenosols under
Tillage ........................................ 187
8.4. Nutrient Fluxes in Alder Stands and Wetlands .......... 190
8.4.1. Elemental Concentrations in Alder Stands ....... 190
8.4.2. Nitrogen Budgets of Wetlands ................... 194
8.5. Stress, Strain and Metastability of Beech, Alder,
Pasture and Agro-Ecosystems ........................... 195
8.5.1. Elemental Imbalances as Strain Indicators ...... 196
8.5.2. Element Budgets and Strain Reactions
of Beech Stands ................................ 198
8.5.3. Strain Reactions of Alder Stands and
Pastures ....................................... 203
8.5.4. Agricultural Impact ............................ 204
8.6. Conclusions ........................................... 204
9. Transport Processes between Lake Belau and its Drainage
Basin ...................................................... 207
Winfrid Kluge and Otto Fränzle
9.1. Introduction .......................................... 207
9.2. A Path Concept as a System-Linking Methodological
Platform .............................................. 207
9.3. Exchange of Water between Lake Belau and its
Catchment ............................................. 209
9.3.1. Hydrological Structure of the Catchment ........ 209
9.3.2. Ecohydrological Structure of Lentic Ecotones ... 214
9.3.3. Water Distribution Matrices of Lentic
Ecotones ....................................... 218
9.4. Non-Point Inputs of Nitrogen .......................... 218
9.4.1. Bonding Forms and Concentrations ............... 218
9.4.2. Path-Based Balance of Nitrogen Inputs
from Contiguous Uplands ........................ 224
9.4.3. Influence of Lentic Ecotones on Non-Point
Inputs of Nitrogen ............................. 224
9.4.4. Inter-Scale Balances of Lateral Fluxes ......... 226
9.5. Non-Point Inputs of Phosphorus ........................ 231
9.5.1. Bonding Forms and Concentrations ............... 231
9.5.2. Atmospheric Input of Phosphorus into Lake
Belau .......................................... 234
9.5.3. Path-Related Phosphorus Inputs into Riparian
Ecotones ....................................... 234
9.5.4. Influence of Lentic Ecotones on Phosphorus
Transfer ....................................... 235
9.6. Comparative Evaluation of Nitrogen and Phosphorus
Fluxes ................................................ 237
9.7. Conclusions ........................................... 239
10.Lake Belau ................................................. 241
Otto Fränzle and Gerald Schernewski
10.1.Introduction .......................................... 241
10.2.Hydrographic Structure of the Lake Belau Drainage
Basin ................................................. 242
10.3.Energetic Setting of Lake Belau and its Drainage
Basin ................................................. 242
10.3.1.Short-Wave Net Radiation ....................... 242
10.3.2.Water Temperature, Wind and Stratification ..... 245
10.4.Dissolved and Particulate Nutrients and Trace
Elements in Water and Sediments ....................... 250
10.4.1.Macronutrient and Carbon Fluxes ................ 253
10.4.2.Micronutrients and Trace Elements .............. 260
10.5.Biocoenoses ........................................... 263
10.5.1.Reed Belts ..................................... 263
10.5.2.Aufwuchs-Associated Nematodes and
Oligochaetes ................................... 265
10.5.3.Molluscs ....................................... 266
10.5.4.Chironomids .................................... 266
10.5.5.Pelagic Phytoplankton and Benthic Algal
Assemblages .................................... 267
10.5.6.Zooplankton and the Microbial Loop ............. 270
10.5.7.Fish ........................................... 272
10.5.8.Species Diversity in the Light of Site
Conditions and Organismic Motility ............. 273
10.6.Conclusions ........................................... 275
11.Ecological Gradients as Causes and Effects of Ecosystem
Organization ............................................... 277
Felix Müller, Otto Fränzle, and Claus-Georg Schimming
11.1.Introduction .......................................... 277
11.2.Gradients in Ecosystems ............................... 278
11.2.1.Structural Gradients ........................... 278
11.2.2.Functional Gradients ........................... 283
11.3.Gradients as Elements of an Integrative Ecosystem
Theory ................................................ 292
11.4.Conclusions ........................................... 294
Part III From Research to Application
12.An Indicator-Based Characterization of the Bornhoved
Key Ecosystems ............................................. 297
Joachim Schrautzer, Felix Müller, Hans-Peter Blume,
Uwe Heinrich, Ernst-Walter Reiche, Uwe Schleuß,
and Klaus Dierssen
12.1.Introduction .......................................... 297
12.2.Methodology ........................................... 298
12.2.1.Ecosystem Classification ....................... 298
12.2.2.Ecosystem Structure and Diversity .............. 298
12.2.3.Water and Nutrient Budgets ..................... 299
12.2.4.Indication of Ecosystem Integrity .............. 303
12.3.Characterization of Ecosystem Types ................... 303
12.3.1.Successional Series on Histosols ............... 303
12.3.2.Successional Series on Mineral Soils ........... 305
12.4.Discussion ............................................ 305
12.4.1.Patterns of Plant-Species Richness ............. 305
12.4.2.Comparative Carbon Budgets of Ecosystems
and their Successional Phases .................. 308
12.4.3.Comparative Nitrogen Budgets of Ecosystems
and their Successional Phases .................. 310
12.4.4.Efficiency Measures ............................ 312
12.4.5.Integrative Characterization of Ecosystem
Evolution ...................................... 312
12.4.6.Relationships between Species Richness
and Ecosystem Functioning ...................... 313
12.4.7.Nutrient Balances of the Study Area ............ 316
12.5.Conclusions ........................................... 316
13.Ecosystem Research and Sustainable Land Use Management ..... 319
Jan Barkmann, Hans-Peter Blume, Ullrich Irmler,
Winfried Kluge, Werner L. Kutsch, Heinrich Reck,
Ernst-Walter Reiche, Michael Trepel, Wilhelm
Windhorst, and Klaus Dierssen
13.1.Introduction .......................................... 319
13.2.Ecosystem Research and Land Use Strategies ............ 319
13.2.1.Concepts of Sustainable Landscape Management ... 319
13.2.2.The Demand for an Ecosystem Approach in
National and International Regulations ......... 321
13.2.3.Ecosystem Integrity - Protection in the Face
of Unspecific Risks to the Human-Environment
Interaction .................................... 325
13.2.4.The Role of Biological Diversity - Losing
Insurance Benefits ............................. 327
13.3.Beyond Sectoral Planning - Outline of the
Precautionary Ecosystem Approach to Sustainable
Landscape Planning .................................... 329
13.3.1.From Sectoral Planning to an Ecosystem-
Oriented Approach .............................. 329
13.3.2.Digital Landscape Analysis and Modelling
as Tools for an Integrative Landscape
Management ..................................... 335
13.3.3.Modelling Biotic Interactions with
Individual-Based Models ........................ 336
13.3.4.Ecohydrological Modelling of Wetland Systems ... 336
13.3.5.Process-Oriented Modelling on the Landscape
Level .......................................... 338
13.3.6.Ecological Economics and Scenario-Guided
Adjustment of Control Systems .................. 342
14.Conclusions: Perspectives for Integrative Landscape
Planning, Management and Monitoring ........................ 345
Klaus Dierssen and Jan Barkmann
14.1.Introduction .......................................... 345
14.2.Definition of Goals and Valuation Procedures .......... 346
14.3.Appropriate Planning Procedures - Integrative
Analysis .............................................. 347
14.4.Realization of Planned Measures ....................... 348
14.5.Ecosystem-Oriented Monitoring and Feedback to
Planning and Management Processes ..................... 349
14.6.Transfer of Knowledge to the Community, to
Stakeholders and to Decision Makers ................... 350
References ................................................. 351
Index ...................................................... 385
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