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<p>The European Union Water Framework Directive (WFD), with initiatives to manage surface water resources, has increased the need for fast and inexpensive methods for monitoring conditions in streams. The objective of this thesis is to assess the potential of near-infrared spectroscopy (NIRS) of epilithic material to become such method. NIRS, which is a technique that is commonly used in industry for process monitoring and quality control of products, registers the chemical properties of organic material on a molecular level. Epilithic material, i.e. the layer of dead and alive material that covers stone surfaces in streams, is continuously influenced by the stream water that flows over it, and it has the potential to integrate chemical and biological conditions over time. The temporal integration is a significant factor, since conditions in streams can change within hours or days. The thesis consists of two published papers. In the first paper a new sampler for epilithic material, the Stone Brusher, was described and the performance evaluated. The Stone Brusher is designed to take qualitative or semi-quantitative samples of epilithic material from stones at 7–50 cm water depth. The epilithic material is dislodged from the stone surface with a rotating brush enclosed in a chamber, and the material is drawn up directly into the sample bottle with an air-cylinder. The operator takes a sample quickly and without putting hands into the water. The sampler is made of plastic, stainless steel and aluminium and weighs 3.1 kg. It is designed to meet the demand for standardized sampling for research and environmental monitoring and to improve working conditions for sampling personnel. The equipment allows sampling from surfaces of bedrock and large stones that cannot be lifted from the bottom. Using data of near-infrared spectroscopy and diatom analyses, this new sampler was evaluated in comparison to the toothbrush method, a primitive method which is the current standard in EU. The results indicate that the Stone Brusher reduces sampling variability compared with the toothbrush method.</p><p>In the second paper, the Stone Brusher was used to collect epilithic material from 65 sites (42 uncontaminated and 23 contaminated) from streams in the widespread mining area called the Skellefte-district in Västerbotten, northern Sweden, in order to test the hypothesis that impact on the epilithic material caused by emissions from mining and mining-related industries can be detected using NIRS. The epilithic material was filtered onto glass fibre filters, measured by NIRS, and the results were modelled using Principal Component Analysis (PCA). The NIRS approach was evaluated by comparing it with the results of chemical and diatom analyses of the same samples. Based on PCA, the NIRS data distinguished contaminated from uncontaminated sites and performed slightly better than chemical analyses and clearly better than diatom analyses. Of the streams designated a priori as contaminated, 74 % were identified as contaminated by NIRS, 65 % by chemical analysis and 26 % by diatom analysis. Unlike chemical analyses of water or of epilithic material samples, NIRS data reflect biological impacts in the streams. Given that, and the simplicity of NIRS-analyses, further studies to assess the use of NIRS of epilithic material are justified. NIRS has the potential to become a fast method for screening in regions where large numbers of streams occur to find impacted streams or as a routine method for temporal monitoring in selected streams for early detection of environmental impact, similar to process monitoring in industry.</p>

The European Union Water Framework Directive (WFD), with initiatives to manage surface water resources, has increased the need for fast and inexpensive methods for monitoring conditions in streams. The objective of this thesis is to assess the potential of near-infrared spectroscopy (NIRS) of epilithic material to become such method. NIRS, which is a technique that is commonly used in industry for process monitoring and quality control of products, registers the chemical properties of organic material on a molecular level. Epilithic material, i.e. the layer of dead and alive material that covers stone surfaces in streams, is continuously influenced by the stream water that flows over it, and it has the potential to integrate chemical and biological conditions over time. The temporal integration is a significant factor, since conditions in streams can change within hours or days. The thesis consists of two published papers. In the first paper a new sampler for epilithic material, the Stone Brusher, was described and the performance evaluated. The Stone Brusher is designed to take qualitative or semi-quantitative samples of epilithic material from stones at 7–50 cm water depth. The epilithic material is dislodged from the stone surface with a rotating brush enclosed in a chamber, and the material is drawn up directly into the sample bottle with an air-cylinder. The operator takes a sample quickly and without putting hands into the water. The sampler is made of plastic, stainless steel and aluminium and weighs 3.1 kg. It is designed to meet the demand for standardized sampling for research and environmental monitoring and to improve working conditions for sampling personnel. The equipment allows sampling from surfaces of bedrock and large stones that cannot be lifted from the bottom. Using data of near-infrared spectroscopy and diatom analyses, this new sampler was evaluated in comparison to the toothbrush method, a primitive method which is the current standard in EU. The results indicate that the Stone Brusher reduces sampling variability compared with the toothbrush method. In the second paper, the Stone Brusher was used to collect epilithic material from 65 sites (42 uncontaminated and 23 contaminated) from streams in the widespread mining area called the Skellefte-district in Västerbotten, northern Sweden, in order to test the hypothesis that impact on the epilithic material caused by emissions from mining and mining-related industries can be detected using NIRS. The epilithic material was filtered onto glass fibre filters, measured by NIRS, and the results were modelled using Principal Component Analysis (PCA). The NIRS approach was evaluated by comparing it with the results of chemical and diatom analyses of the same samples. Based on PCA, the NIRS data distinguished contaminated from uncontaminated sites and performed slightly better than chemical analyses and clearly better than diatom analyses. Of the streams designated a priori as contaminated, 74 % were identified as contaminated by NIRS, 65 % by chemical analysis and 26 % by diatom analysis. Unlike chemical analyses of water or of epilithic material samples, NIRS data reflect biological impacts in the streams. Given that, and the simplicity of NIRS-analyses, further studies to assess the use of NIRS of epilithic material are justified. NIRS has the potential to become a fast method for screening in regions where large numbers of streams occur to find impacted streams or as a routine method for temporal monitoring in selected streams for early detection of environmental impact, similar to process monitoring in industry.

Intensive Landwirtschaft ist eine der allgegenwärtigsten anthropogenen Bedrohungen für aquatische Ökosysteme und beeinflusst sowohl die abiotischen als auch die biotischen Eigenschaften von Fließgewässerökosystemen. Diese resultieren aus der Vielzahl von Störungen, die mit Landnutzungsänderungen verbunden sind, wie z.B. Uferkahlschlag und dem damit verbundenen Verlust von allochthonem organischem Material und Beschattung, Nährstoff- und Pestizidkontamination, Feinsedimenteintrag durch Erosion und Oberflächenabfluss, sowie beeinträchtigte Hydromorphologie und damit verbundenem Verlust von vielfältigen Lebensräumen. Umfangreiche Literatur zu den Auswirkungen von Landwirtschaft auf einzelne Ökosystemkomponenten wie Makroinvertebraten, Biofilm oder Fischfauna existiert bereits, wobei der Schwerpunkt hier zumeist auf strukturellen Ansätzen oder einzelnen Ökosystemprozessen liegt. Allerdings haben nur wenige Studien umfassende Auswertungen angestellt, wie intensive Landwirtschaft die Funktion des gesamten aquatischen Ökosystems und die Wechselwirkungen zwischen den Ökosystemkomponenten beeinflusst. Um komplexe Effektmuster landwirtschaftlicher Stressoren auf Fließgewässerstruktur- und funktion sowie mechanistische Wirkzusammenhänge zwischen trophischen Ebenen besser zu verstehen wurden im Rahmen der vorliegenden Arbeit zwei Bäche die innerhalb landwirtschaftlicher Einzugsgebiete liegen und zwei bewaldete Referenzbäche im Harz, Sachsen-Anhalt, hinsichtlich ihrer physiko-chemischen Eigenschaften, der Menge und der Einträge von benthischem organischem Material, der Primärproduktion und organischen Kohlenstoffkreisläufen sowie der Sekundärproduktion von Makroinvertebraten und Fischen über einen Zeitraum von 12-17 Monate miteinander verglichen. Intensive Landwirtschaft hatte starke Auswirkungen auf die physiko-chemischen Eigenschaften der untersuchten Bachoberläufe. Wir ermittelten spezifische Leitfähigkeitswerte in landwirtschaftlich genutzten Bächen, welche die Konzentration gelöster Ionen in bewaldeten Referenzbächen um eine Größenordnung übertrafen. Die Konzentrationen von gelöstem anorganischem Stickstoff und löslichem reaktivem Phosphor waren in den landwirtschaftlichen Bächen im Durchschnitt viermal höher und der pH-Wert war im Durchschnitt eine Einheit niedriger als in den bewaldeten Bächen. Die Wassertemperatur war in landwirtschaftlich genutzten Bächen im Durchschnitt 3°C wärmer und zeigte höhere tageszeitliche Schwankungen. Der Eintrag von partikulärem organischem Material (POM) in die landwirtschaftlichen Bäche war ca. 30-mal niedriger als der Eintrag in die Referenzbäche, und die Menge an benthischem organischem Material (BOM) war in landwirtschaftlichen Bächen signifikant niedriger als in den Referenzbächen. Landwirtschaftliche Fließgewässer wiesen signifikant höhere Mengen an feinem benthischem organischem Material (FBOM) auf, hatten aber geringere Bestände an grobem benthischem organischem Material (CBOM) als die Referenzgewässer. Während in bewaldeten Fließgewässern die zeitliche Dynamik des BOM einem saisonalen Muster folgte, waren die Schwankungen in den landwirtschaftlichen Fließgewässern überwiegend stochastisch und durch anthropogene Aktivitäten, wie das Mähen der Uferrandstreifen, beeinflusst. Auch die Retention von POM war in landwirtschaftlich genutzten Bächen im Vergleich zu bewaldeten Bächen deutlich geringer, was auf eine geringere morphologische Komplexität und die größere hydrologische Variabilität, insbesondere auf die Abflussganglinien, zurückzuführen ist. Innerhalb des Experimentes zur Aufnahme von gelöstem organischem Kohlenstoff (DOC) war die Biofilmqualität in den landwirtschaftlichen Bächen ebenfalls höher. Interessanterweise zeigte jedoch der landwirtschaftliche Bach Hassel einen höheren Grad an Heterotrophie als der bewaldete Bach Wormsgraben. Da die biofilmspezifische DOC-Aufnahme im bewaldeten Bach nur 4 % der Gesamtaufnahme betrug, kann davon ausgegangen werden, dass die mikrobiell vermittelte Verarbeitung von Nährstoffen in der hyporheischen Zone den größten Anteil an der DOC-Aufnahme ausmachte. Vermutlich begünstigte die höhere Verweildauer und der bessere Austausch zwischen Oberflächenwasser und der hyporheischen Zone im bewaldeten Bach Wormsgraben die heterotrophe Nährstoffumsetzung im interstitiellen Porenraum. Die Bewertung der Makroinvertebraten-Gemeinschaft ergab, dass der Artenreichtum in landwirtschaftlichen Bächen im Vergleich zu bewaldeten Bächen um die Hälfte reduziert war. Die Biomasse, Dichte und Sekundärproduktion der Makroinvertebraten (Makroinvertebratensekundärproduktion, MSP) nahm jedoch entlang eines Gradienten der Ressourcenqualität und -quantität zu, mit höchster Produktivität im landwirtschaftlichen Bach Sauerbach, gefolgt vom landwirtschaftlichen Bach Getel und den bewaldeten Bächen Ochsenbach und Wormsgraben. Der Anstieg der MSP war hauptsächlich auf Zunahmen innerhalb der funktionellen Ernährungsgruppen der Sedimentfresser und Weidegänger zurückzuführen. Höhere MSP, Biomasse und Dichte korrelierten positiv mit niedrigeren N:P- und C:P-Verhältnissen des Biofilms und der Ressourcenmenge (höhere Biofilm-Zuwachsrate und Menge an krautiger Vegetation, Chlorophyll a-, TDN- (Summe an gelöstem Stickstoff), SRP- (gelöster reaktiver Phosphor) und POC- (Partikulärer organischer Kohlenstoff) Konzentrationen von Biofilm und Seston). Hohe Interaktionsstärken zwischen Zerkleinerern und grobpartikulärem organischem Material (CPOM) in landwirtschaftlichen Bächen deuteten auf eine Ressourcenlimitierung durch allochthones organisches Material hin. Da die Interaktionsstärken zwischen Weidegängern und Biofilm sowie zwischen feinpartikulärem organischem Material (FPOM) und Sammlern in landwirtschaftlichen Bächen deutlich geringer waren als in bewaldeten Bächen ist anzunehmen, dass die Gruppe der Zerkleinerer diese Limitation durch hohe funktionelle Plastizität und damit alternative Ressourcennutzung ausgleichen konnten. Die Analyse der Fischsekundärproduktion und Interaktionsstärken zwischen Fischen und ihrer Makroinvertebraten-Beute konnte zusätzlich zeigen, dass die landwirtschaftliche Flächennutzung zu einer starken Verschiebung des Fischartenspektrums führen und die Top-down Kontrolle der MSP durch Fische dadurch deutlich zurückgehen kann. Diese Ergebnisse zeigen, dass der Verlust der Ufervegetation und die übermäßige Nährstoffbelastung ökologische Nischen homogenisieren und hochproduktive Generalisten mit hoher Nahrungsplastizität begünstigten, die flexibel zwischen ubiquitären und stochastisch verfügbaren Nahrungsressourcen wechseln können. Insgesamt konnten wir mit dieser Studie darlegen, durch welche Mechanismen aus hoch-diversen, nährstoffbegrenzten, bewaldeten Bachoberläufen, die stark von terrestrischen Laubeinträgen abhängig sind, durch landwirtschaftliche Flächennutzung homogene, nährstoffbelastete und taxonomisch vereinfachte Bäche, mit einem hohen Maß an Autochthonie, werden können. Die gleichzeitige Analyse struktureller und funktioneller Indikatoren zeigte, dass die Landwirtschaft strukturelle Messgrößen wie den Artenreichtum beeinflusst, ohne dass es zu größeren Veränderungen in der Funktion kommen muss (ähnliche MSP im landwirtschaftlich genutzten Bach Getel im Vergleich zu bewaldeten Referenzbächen), und dass die Variabilität in der Funktion (Retention von organischem Material, Laubabbau) nicht automatisch die Informationen widerspiegelt, die die üblicherweise bewerteten chemischen oder biologischen strukturellen Messgrößen liefern. Die Ergebnisse dieser Arbeit zeigen deutlich, dass eine Kombination mehrerer struktureller und funktioneller Indikatoren entlang der Nahrungsnetzhierarchie erforderlich ist, um mechanistisch zu verstehen, wie intensive Landwirtschaft die Ökosysteme von Fließgewässern beeinflusst. So erlaubten z.B. nur die Informationen über die Qualität und Quantität der Primärproduktion, in Kombination mit strukturellen und funktionellen Informationen über die Makroinvertebratengemeinschaft, ein mechanistisches Verständnis darüber, wie landwirtschaftliche Stressoren die Produktivität mehrerer trophischer Ebenen des Nahrungsnetzes von Fließgewässern beeinflussen und damit auch wie diese Auswirkungen gemildert werden könnten. Diese Ergebnisse verdeutlichen die dringende Notwendigkeit, effektive und breite Gewässerrandstreifen zu erhalten oder zu errichten, um Sedimenteinträge durch Erosion- und Oberflächenabfluss zu verringern, sowie Einträge von Nährstoffen und Pestiziden zurückzuhalten, um eine verbesserte Vereinbarkeit von Landwirtschaft und Naturschutz zu erwirken. Diese Maßnahmen hätten eine positive Auswirkung auf Lebensraum- und Ressourcenvielfalt und schützen damit sowohl terrestrische als auch aquatische Ökosysteme und die damit verbundenen essentiellen Ökosystemdienstleistungen dieser Systeme.:Table of contents Table of contents 1 List of tables 5 List of figures 6 List of abbreviations and acronyms 8 Abstract 11 Zusammenfassung 14 1. General introduction 18 1.1 Forested headwater streams - the significance of aquatic-terrestrial coupling 18 1.2 Effects of agricultural land use on forested headwater streams 22 1.3 Integrating measures of ecosystem functioning in stream ecosystem assessment 24 1.4 Aims and objectives 28 1.5 Thesis outline 30 2. Chapter I: Agricultural land use alters temporal dynamics and the composition of organic matter in temperate headwater streams 31 2.1 Introduction 31 2.2 Methods 33 2.2.1 Study sites 33 2.2.2 Sampling of POM inputs and standing crops 37 2.2.3 Organic matter sample processing 37 2.2.4 Physical and chemical stream characteristics 38 2.2.5 C spiraling metrics 39 2.2.6 Litter decomposition 39 2.2.7 Data analyses 41 2.3 Results 43 2.3.1 Organic matter inputs 43 2.3.2 Benthic organic matter 46 2.3.3 Organic matter retention 52 2.3.4 Environmental drivers of BOM dynamics 55 2.3.5 Leaf litter decomposition 55 2.4 Discussion 57 2.4.1 Effects of agricultural land use on POM inputs and BOM dynamics 57 2.4.2 Organic matter retention 60 2.4.3 Organic matter processing 61 2.4.4 Ecosystem-level implications of altered OM dynamics 62 2.5 Conclusions 63 3. Chapter II: Biofilm-specific uptake does not explain differences in whole-stream DOC tracer uptake between a forest and an agricultural stream 64 3.1 Introduction 64 3.2 Methods 66 3.2.1 Site description 66 3.2.2 Stable isotope tDOC labeling 68 3.2.3 Experimental design 68 3.2.4 Sampling and analysis 69 3.2.5 Statistical analysis 72 3.3 Results 74 3.3.1 Stream characteristics 74 3.3.2 DOC uptake 75 3.3.3 Benthic biofilm attributes 79 3.4 Discussion 84 3.4.1 Mechanisms linking hydromorphology, benthic-biofilm uptake and whole-stream uptake 84 3.4.2 Comparison of whole-stream tDOC uptake with other studies 86 3.4.3 Benthic biofilm attributes 87 3.5 Conclusions 89 4. Chapter III: Resource supply and organismal dominance are associated with high secondary production in temperate agricultural streams 90 4.1 Introduction 90 4.2 Methods 93 4.2.1 Study sites 93 4.2.2 Macroinvertebrate sampling and processing 97 4.2.3 Macroinvertebrate secondary production (MSP) 97 4.2.4 Environmental characteristics of streams 98 4.2.5 Benthic organic matter 99 4.2.6 Biofilm biomass accrual and stoichiometry 99 4.2.7 Fish biomass 100 4.2.8 Ingestion rates and interaction strength 100 4.2.9 Data analyses 101 4.3 Results 104 4.3.1 Environmental characteristics of the study streams 104 4.3.2 Structural descriptors of the macroinvertebrate community 106 4.3.3 Macroinvertebrate secondary production (MSP) 107 4.3.4 Environmental drivers of MSP and biomass 109 4.3.5 Relationships between MSP and species richness and evenness 111 4.3.6 Consumer-resource interactions 111 4.4 Discussion 113 4.4.1 Environmental drivers of MSP and biomass 113 4.4.2 Biological mechanisms associated with MSP 116 4.4.3 Top-down vs. bottom-up control 118 4.5 Conclusion 120 5. General discussion 121 5.1 Environmental conditions 121 5.2 Allochthonous organic matter dynamics 122 5.3 Organic matter processing 124 5.4 Primary production 124 5.5 Drivers of macroinvertebrate secondary production 127 5.6 Fish community 128 6. General conclusion 132 7. Implications for the assessment of running water ecosystems 134 8. Outlook 137 References 139 Appendix 182 Chapter I 182 Chapter II 189 Supplemental methods 189 Chapter III 198 Acknowledgements 212 Curriculum vitae 215 List of publications 217 Publication output during thesis period 217 Further publications 218 Eidesstattliche Erklärung 219 / Intensive crop agriculture is one of the most ubiquitous and pervasive anthropogenic threats to aquatic ecosystems. Important agriculture-related pressures include riparian clearcutting and the associated loss of allochthonous organic matter inputs and shading, nutrient and pesticide contamination, fine sediment inputs due to erosion and run-off, increased surface runoff, flashier hydrographs as well as degraded geomorphology and habitat diversity. The multitude of stressors deriving from agricultural land use often produces concomitant effects on several groups of biological organism and associated functions. While effects on single ecosystem components such as macroinvertebrates, biofilm or fish fauna with primary focus on structural components or single ecosystem processes are well described, only few studies have compiled comprehensive data sets on how agriculture affects the functioning of the entire aquatic ecosystem and interactions among ecosystem components. Thus, this thesis aimed to fill this knowledge gap by comparing agricultural and forested reference streams regarding their physicochemical characteristics, organic matter standing stocks and inputs, primary production, organic carbon spiraling, DOC uptake rates as well as secondary production of macroinvertebrates and fish for a period of 12-17 months. Intensive agricultural land use had strong effects on the physico-chemistry of the studied headwater streams of the main study (two agricultural streams vs. two forested streams in the Harz mountains, Saxony-Anhalt). We ascertained specific conductivity values in agricultural streams that exceeded the ones measured in forested streams by an order of magnitude. Concentrations of dissolved inorganic nitrogen and soluble reactive phosphorus were on average four times higher and pH was on average one unit lower in agricultural streams than in forested streams. Water temperature was on average 3°C higher in agricultural streams and showed higher diurnal variations. Particulate organic matter (POM) inputs to forested streams were approx. 30-times higher than inputs into agricultural streams, and mean standing crops of total benthic organic matter (BOM) were significantly lower in agricultural streams than in forested streams. Agricultural streams had significantly higher standing crops of fine benthic organic matter (FBOM), but less coarse benthic organic matter (CBOM) than forested streams. While in forested streams temporal dynamics of organic matter standing stocks followed a seasonal pattern, variations were predominantly stochastic and influenced by anthropogenic activities such as stream margin mowing in agricultural streams. Also, the retention of POM was distinctly lower in agricultural compared to forested streams due to lower in-stream complexity and flashier hydrographs in agricultural streams indicating that agricultural streams are less efficient to retain organic matter, facilitating the loss of carbon to downstream sections of streams. Within the main study streams, biofilm accrual rates and chlorophyll a content were six and eight times higher, respectively, and biofilm N:P and C:P ratios were three times lower in agricultural than forested streams. In the dissolved organic carbon-uptake experiment (one agricultural stream vs. one forested stream), biofilm quality was similarly higher in the agricultural stream Hassel. Against expectations, the Hassel showed a higher level of heterotrophy than the forested stream Wormsgraben. However, the total and biofilm associated uptake of labeled leaf-leachate dissolved organic carbon (DOC) in the Hassel was six and two times higher than in the forested Wormsgraben, respectively. As the biofilm-specific uptake in the forested stream amounted to only 4% of the whole stream uptake, it seems that microbial-mediated processing of nutrients in the hyporheic zone is responsible for the largest proportion of DOC uptake. Presumably, the higher transient storage and the heterotrophic community resident in the interstitial pore space of the forested stream has a higher demand for labile DOC and hence, retained distinctly more DOC than in the agricultural stream, highlighting the importance of functional hyporheic zones for carbon spiraling. The macroinvertebrate community assessment revealed that species richness was reduced by half in agricultural compared to forested streams. Macroinvertebrate biomass, density, and secondary production (MSP), however, increased along a gradient of resource quality and quantity with highest productivity in the agricultural stream Sauerbach, followed by the agricultural stream Getel, and the forested streams Ochsenbach and Wormsgraben. The increase in MSP was mainly due to higher production within the functional feeding groups (FFGs) of gatherer/collectors and grazers. Higher MSP, biomass, and density correlated positively with lower biofilm N:P and C:P ratios and resource quantity (higher biofilm accrual rate and standing crops of riparian herbaceous vegetation, biofilm and seston chlorophyll a, total dissolved nitrogen, soluble reactive phosphorus, and particulate organic carbon concentrations). High interaction strengths between shredders and coarse particulate organic carbon (CPOM) in agricultural streams indicated a resource limitation of allochthonous organic matter, while this seemed to have no effect on macroinvertebrate productivity as interaction strengths between grazers and biofilm as well as between fine particulate organic carbon (FPOM) and gatherers were distinctly lower in agricultural than in forested streams. These findings show that the loss of riparian canopy and excess nutrient conditions homogenized ecological niches and favored highly productive non-insect generalist species with high feeding plasticity to switch flexibly between ubiquitous and stochastically available food resources. In addition, analyses of fish secondary production and interaction strengths between fish and their macroinvertebrate prey demonstrated that agriculture may cause a major shift in fish species community composition, resulting in a decline in fish top-down control on MSP. Overall, the presented results demonstrate that agriculture induces a shift from nutrient-limited forested headwaters dependent on detrital pathways and closely coupled to riparian subsidies with highly diverse macroinvertebrate communities to homogenous, nutrient contaminated and taxonomically simplified streams with high levels of autochthony. The simultaneous analysis of structural and functional indicators showed that agriculture affects structural measures of community composition such as species richness without major changes in process rates (similar level of macroinvertebrate secondary production in the agricultural stream Getel compared to forested streams) and that variability in function (organic matter retention, decomposition) does not automatically reflect the information provided by commonly assessed structural measures of communities. The results of this thesis clearly showed that a combination of multiple structural and functional indicators along the food web hierarchy is required to mechanistically understand how intensive agricultural land use affects stream ecosystems. For example, only the information on quality and quantity of primary production combined with structural and functional information on the macroinvertebrate community allowed to mechanistically understand how agricultural stressors affect the productivity of multiple trophic levels of the stream food web and thus, how these effects can be mitigated. Given that such mitigation measures would largely address excessive nutrient and sediment inputs, the loss of habitat heterogeneity and natural resource dynamics, the high relevance of riparian buffer zones known for their effective the retention of nutrients, erosional run-off and the provision of resource quantity and quality is evident. Consequently, there is an urgent need to preserve or restore effective riparian zones along stream networks aiding both terrestrial and aquatic ecosystems while integrating protection measures for other anthropogenic pressures such as habitat degradation and climate change.:Table of contents Table of contents 1 List of tables 5 List of figures 6 List of abbreviations and acronyms 8 Abstract 11 Zusammenfassung 14 1. General introduction 18 1.1 Forested headwater streams - the significance of aquatic-terrestrial coupling 18 1.2 Effects of agricultural land use on forested headwater streams 22 1.3 Integrating measures of ecosystem functioning in stream ecosystem assessment 24 1.4 Aims and objectives 28 1.5 Thesis outline 30 2. Chapter I: Agricultural land use alters temporal dynamics and the composition of organic matter in temperate headwater streams 31 2.1 Introduction 31 2.2 Methods 33 2.2.1 Study sites 33 2.2.2 Sampling of POM inputs and standing crops 37 2.2.3 Organic matter sample processing 37 2.2.4 Physical and chemical stream characteristics 38 2.2.5 C spiraling metrics 39 2.2.6 Litter decomposition 39 2.2.7 Data analyses 41 2.3 Results 43 2.3.1 Organic matter inputs 43 2.3.2 Benthic organic matter 46 2.3.3 Organic matter retention 52 2.3.4 Environmental drivers of BOM dynamics 55 2.3.5 Leaf litter decomposition 55 2.4 Discussion 57 2.4.1 Effects of agricultural land use on POM inputs and BOM dynamics 57 2.4.2 Organic matter retention 60 2.4.3 Organic matter processing 61 2.4.4 Ecosystem-level implications of altered OM dynamics 62 2.5 Conclusions 63 3. Chapter II: Biofilm-specific uptake does not explain differences in whole-stream DOC tracer uptake between a forest and an agricultural stream 64 3.1 Introduction 64 3.2 Methods 66 3.2.1 Site description 66 3.2.2 Stable isotope tDOC labeling 68 3.2.3 Experimental design 68 3.2.4 Sampling and analysis 69 3.2.5 Statistical analysis 72 3.3 Results 74 3.3.1 Stream characteristics 74 3.3.2 DOC uptake 75 3.3.3 Benthic biofilm attributes 79 3.4 Discussion 84 3.4.1 Mechanisms linking hydromorphology, benthic-biofilm uptake and whole-stream uptake 84 3.4.2 Comparison of whole-stream tDOC uptake with other studies 86 3.4.3 Benthic biofilm attributes 87 3.5 Conclusions 89 4. Chapter III: Resource supply and organismal dominance are associated with high secondary production in temperate agricultural streams 90 4.1 Introduction 90 4.2 Methods 93 4.2.1 Study sites 93 4.2.2 Macroinvertebrate sampling and processing 97 4.2.3 Macroinvertebrate secondary production (MSP) 97 4.2.4 Environmental characteristics of streams 98 4.2.5 Benthic organic matter 99 4.2.6 Biofilm biomass accrual and stoichiometry 99 4.2.7 Fish biomass 100 4.2.8 Ingestion rates and interaction strength 100 4.2.9 Data analyses 101 4.3 Results 104 4.3.1 Environmental characteristics of the study streams 104 4.3.2 Structural descriptors of the macroinvertebrate community 106 4.3.3 Macroinvertebrate secondary production (MSP) 107 4.3.4 Environmental drivers of MSP and biomass 109 4.3.5 Relationships between MSP and species richness and evenness 111 4.3.6 Consumer-resource interactions 111 4.4 Discussion 113 4.4.1 Environmental drivers of MSP and biomass 113 4.4.2 Biological mechanisms associated with MSP 116 4.4.3 Top-down vs. bottom-up control 118 4.5 Conclusion 120 5. General discussion 121 5.1 Environmental conditions 121 5.2 Allochthonous organic matter dynamics 122 5.3 Organic matter processing 124 5.4 Primary production 124 5.5 Drivers of macroinvertebrate secondary production 127 5.6 Fish community 128 6. General conclusion 132 7. Implications for the assessment of running water ecosystems 134 8. Outlook 137 References 139 Appendix 182 Chapter I 182 Chapter II 189 Supplemental methods 189 Chapter III 198 Acknowledgements 212 Curriculum vitae 215 List of publications 217 Publication output during thesis period 217 Further publications 218 Eidesstattliche Erklärung 219

<p>To further investigate the strength of the association and the relative advantages of the association to both organisms, several manipulation experiments were set up. A cage experiment set up in the shallow subtidal zone showed that the coralline survived equally well without the winkle and did therefore not require the winkle or its empty shell for survival. A second controlled laboratory aquarium experiment was designed under both fluorescent (rich in blue light) and incandescent light (rich in red light) to ascertain whether the coralline had a preference for O. sinensis over the similar O. tigrina. This experiment was inconclusive as no recruitment was obtained under either of the light regimes. A third laboratory experiment was designed to determine whether the extra coralline weight had any possible advantage to the winkle, particularly against predation from the rock lobster Jasus lalandii. Results suggested that there were no apparent advantages to the winkle bearing the extra coralline load as adult O. sinensis bearing the coralline alga (3.7 &plusmn / 2.2 winkles 24hr-1) were equally prone to predation than those lacking the coralline (2.3 &plusmn / 1.9 winkles 24hr-1) (p = 0.184). Observations suggested instead that the convoluted nature of the coralline may indeed have promoted predation. We ultimately deduced that the high occurrence of the coralline on the shells of O. sinensis was probably due to the substantial overlap in the niches of the two organisms. This conclusion was supported by the high densities of juvenile O. sinensis combined with the high percent cover abundance of the coralline in intertidal rockpools. Understanding sexual reproduction in coralline algae as well as the life cycle of the winkle, ultimately provided insight into the postulated life cycle of this coralline-winkle association...</p>

<p>To further investigate the strength of the association and the relative advantages of the association to both organisms, several manipulation experiments were set up. A cage experiment set up in the shallow subtidal zone showed that the coralline survived equally well without the winkle and did therefore not require the winkle or its empty shell for survival. A second controlled laboratory aquarium experiment was designed under both fluorescent (rich in blue light) and incandescent light (rich in red light) to ascertain whether the coralline had a preference for O. sinensis over the similar O. tigrina. This experiment was inconclusive as no recruitment was obtained under either of the light regimes. A third laboratory experiment was designed to determine whether the extra coralline weight had any possible advantage to the winkle, particularly against predation from the rock lobster Jasus lalandii. Results suggested that there were no apparent advantages to the winkle bearing the extra coralline load as adult O. sinensis bearing the coralline alga (3.7 &plusmn / 2.2 winkles 24hr-1) were equally prone to predation than those lacking the coralline (2.3 &plusmn / 1.9 winkles 24hr-1) (p = 0.184). Observations suggested instead that the convoluted nature of the coralline may indeed have promoted predation. We ultimately deduced that the high occurrence of the coralline on the shells of O. sinensis was probably due to the substantial overlap in the niches of the two organisms. This conclusion was supported by the high densities of juvenile O. sinensis combined with the high percent cover abundance of the coralline in intertidal rockpools. Understanding sexual reproduction in coralline algae as well as the life cycle of the winkle, ultimately provided insight into the postulated life cycle of this coralline-winkle association...</p>

Magister Scientiae (Biodiversity and Conservation Biology) / To further investigate the strength of the association and the relative advantages of the association to both organisms, several manipulation experiments were set up. A cage experiment set up in the shallow subtidal zone showed that the coralline survived equally well without the winkle and did therefore not require the winkle or its empty shell for survival. A second controlled laboratory aquarium experiment was designed under both fluorescent (rich in blue light) and incandescent light (rich in red light) to ascertain whether the coralline had a preference for O. sinensis over the similar O. tigrina. This experiment was inconclusive as no recruitment was obtained under either of the light regimes. A third laboratory experiment was designed to determine whether the extra coralline weight had any possible advantage to the winkle, particularly against predation from the rock lobster Jasus lalandii. Results suggested that there were no apparent advantages to the winkle bearing the extra coralline load as adult O. sinensis bearing the coralline alga (3.7 &plusmn; 2.2 winkles 24hr-1) were equally prone to predation than those lacking the coralline (2.3 &plusmn; 1.9 winkles 24hr-1) (p = 0.184). Observations suggested instead that the convoluted nature of the coralline may indeed have promoted predation. We ultimately deduced that the high occurrence of the coralline on the shells of O. sinensis was probably due to the substantial overlap in the niches of the two organisms. This conclusion was supported by the high densities of juvenile O. sinensis combined with the high percent cover abundance of the coralline in intertidal rockpools. Understanding sexual reproduction in coralline algae as well as the life cycle of the winkle, ultimately provided insight into the postulated life cycle of this coralline-winkle association. / South Africa