Disentangling multiple chemical and non-chemical stressors in a lotic ecosystem using a longitudinal approach.

Meeting ecological and water quality standards in lotic ecosystems is often failed due to multiple stressors. However, disentangling stressor effects and identifying relevant stressor-effect-relationships in complex environmental settings remain major challenges. By combining state-of-the-art methods from ecotoxicology and aquatic ecosystem analysis, we aimed here to disentangle the effects of multiple chemical and non-chemical stressors along a longitudinal land use gradient in a third-order river in Germany. We distinguished and evaluated four dominant stressor categories along this gradient: (1) Hydromorphological alterations: Flow diversity and substrate diversity correlated with the EU-Water Framework Directive based indicators for the quality element macroinvertebrates, which deteriorated at the transition from near-natural reference sites to urban sites. (2) Elevated nutrient levels and eutrophication: Low to moderate nutrient concentrations together with complete canopy cover at the reference sites correlated with low densities of benthic algae (biofilms). We found no more systematic relation of algal density with nutrient concentrations at the downstream sites, suggesting that limiting concentrations are exceeded already at moderate nutrient concentrations and reduced shading by riparian vegetation. (3) Elevated organic matter levels: Wastewater treatment plants (WWTP) and stormwater drainage systems were the primary sources of bioavailable dissolved organic carbon. Consequently, planktonic bacterial production and especially extracellular enzyme activity increased downstream of those effluents showing local peaks. (4) Micropollutants and toxicity-related stress: WWTPs were the predominant source of toxic stress, resulting in a rapid increase of the toxicity for invertebrates and algae with only one order of magnitude below the acute toxic levels. This toxicity correlates negatively with the contribution of invertebrate species being sensitive towards pesticides (SPEARpesticides index), probably contributing to the loss of biodiversity recorded in response to WWTP effluents. Our longitudinal approach highlights the potential of coordinated community efforts in supplementing established monitoring methods to tackle the complex phenomenon of multiple stress.

Longitudinal patterns and response lengths of algae in riverine ecosystems: A model analysis emphasising benthic-pelagic interactions.

In riverine ecosystems primary production is principally possible in two habitats: in the benthic layer by sessile algae and in the surface water by planktonic algae being transported downstream. The relevance of these two habitats generally changes along the rivers' continuum. However, analyses of the interaction of algae in these two habitats and their controlling factors in riverine ecosystems are, so far, very rare. We use a simplified advection-diffusion model system combined with ecological process kinetics to analyse the interaction of benthic and planktonic algae and nutrients along idealised streams and rivers at regional to large scales. Because many of the underlying processes affecting algal dynamics are influenced by depth, we focus particularly on the impact of river depth on this interaction. At constant environmental conditions all state variables approach stable spatial equilibria along the river, independent of the boundary conditions at the upstream end. Because our model is very robust against changes of turbulent diffusion and stream velocity, these spatial equilibria can be analysed by a simplified ordinary differential equation (ode) version of our model. This model variant reveals that at shallower river depths, phytoplankton can exist only when it is subsidised by detaching benthic algae, and in turn, at deeper river depths, benthic algae can exist only in low biomasses which are subsidised by sinking planktonic algae. We generalise the spatial dynamics of the model system using different conditions at the upstream end of the model, which mimic various natural or anthropogenic factors (pristine source, dam, inflow of a waste water treatment plant, and dilution from e.g. a tributary) and analyse how these scenarios influence different aspects of the longitudinal spatial dynamics of the full spatial model: the relation of spatial equilibrium to spatial maximum, the distance to the spatial maximum, and the response length. Generally, our results imply that shallow systems recover within significantly shorter distances from spatially distinct disturbances when compared to deep systems, independent of the type of disturbance.

Can nutrient pathways and biotic interactions control eutrophication in riverine ecosystems? Evidence from a model driven mesocosm experiment.

Ecological theory predicts that the relative importance of benthic to planktonic primary production usually changes along the rivers' continuum from a predomination of benthic algae in lower stream orders to a predomination of planktonic algae at higher orders. Underlying mechanisms driving the interaction between algae in these habitats, its controlling factors and consequences for riverine ecosystems are, however, only partly understood. We present a mechanistic analysis of the governing ecological processes using a simplified, numerical model and examine how abiotic factors and biotic interactions influence benthic and planktonic algae by changing resource competition. We compare the outcome of the model with the results of a factorial mesocosm experiment mimicking the parameter spaces of the model. The results show a remarkable similarity with regard to the temporal development of benthic and pelagic algal biomass and shifting dominance patterns. In particular we analyse the effects of the pathways of nutrient supply (upwelling from the hyporheic zone, direct supply to the surface water, or via both pathways) and grazing in a gradient of river depths. Our results show that detachment of benthic algae, sinking of planktonic algae and the pathway of nutrient supply are key processes determining the respective algal biomass distributions particularly in shallow and intermediate deep systems. Increasing nutrient supply increases algal biomasses, but does not change the general pattern of the interactions. Decreasing light supply decreases the dominance of planktonic algae, but increases dissolved nutrients. At intermediate to high grazing rates algal biomass can be controlled by grazers, but however, at high grazing rates, dissolved nutrients accumulate in the surface water. Our results indicate that nutrient pathways, resource competition and internal control by grazing need to be considered explicitly for the understanding and explanation of eutrophication phenomena in riverine ecosystems. As a consequence, ecologically effective eutrophication management of running water systems has to go beyond the control of nutrient emissions or the achievement of limiting threshold values in the receiving waters, but requires the consideration of the nutrient pathways (surface water versus groundwater) and the shifting biological controls from lower to higher order stream ecosystems.

Biogeochemical patterns in a river network along a land use gradient.

The Bode catchment (Germany) shows strong land use gradients from forested parts of the National Park (23% of total land cover) to agricultural (70%) and urbanised areas (7%). It is part of the Terrestrial Environmental Observatories of the German Helmholtz association. We performed a biogeochemical analysis of the entire river network. Surface water was sampled at 21 headwaters and at ten downstream sites, before (in early spring) and during the growing season (in late summer). Many parameters showed lower concentrations in headwaters than in downstream reaches, among them nutrients (ammonium, nitrate and phosphorus), dissolved copper and seston dry mass. Nitrate and phosphorus concentrations were positively related to the proportion of agricultural area within the catchment. Punctual anthropogenic loads affected some parameters such as chloride and arsenic. Chlorophyll a concentration and total phosphorus in surface waters were positively related. The concentration of dissolved organic carbon (DOC) was higher in summer than in spring, whereas the molecular size of DOC was lower in summer. The specific UV absorption at 254 nm, indicating the content of humic substances, was higher in headwaters than in downstream reaches and was positively related to the proportion of forest within the catchment. CO2 oversaturation of the water was higher downstream compared with headwaters and was higher in summer than in spring. It was correlated negatively with oxygen saturation and positively with DOC concentration but negatively with DOC quality (molecular size and humic content). A principle component analysis clearly separated the effects of site (44%) and season (15%), demonstrating the strong effect of land use on biogeochemical parameters.

Physical determinants of phytoplankton production, algal stoichiometry, and vertical nutrient fluxes.

Most phytoplankters face opposing vertical gradients in light versus nutrient supplies but have limited capacities for vertical habitat choice. We therefore explored a dynamical model of negatively buoyant algae inhabiting a one-dimensional water column to ask how water column depth and turbulence constrain total (areal) phytoplankton biomass. We show that the population persistence boundaries in water column depth-turbulence space are set by sinking losses and light limitation but that nutrients are most limiting to total biomass in water columns that are neither too shallow or too weakly mixed (where sinking losses prevail) nor too deep and turbulent (where light limitation prevails). In shallow waters, the most strongly limiting process is nutrient influx to the bottom of the water column (e.g., from sediments). In deep waters, the most strongly limiting process is turbulent upward transport of nutrients to the photic zone. Consequently, the highest total biomasses are attained in turbulent waters at intermediate water column depths and in deep waters at intermediate turbulences. These patterns are insensitive to the assumption of fixed versus flexible algal carbon-to-nutrient stoichiometry, and they arise irrespective of whether the water column is a surface layer above a deep water compartment or has direct contact with sediments.

Transient dynamics of pelagic producer-grazer systems in a gradient of nutrients and mixing depths.

Phytoplankton-grazer dynamics are often characterized by long transients relative to the length of the growing season. Using a phytoplankton-grazer model parameterized for Daphnia pulex with either flexible or fixed algal carbon:nutrient stoichiometry, we explored how nutrient and light supply (the latter by varying depth of the mixed water column) affect the transient dynamics of the system starting from low densities. The system goes through an initial oscillation across nearly the entire light-nutrient supply space. With flexible (but not with fixed) algal stoichiometry, duration of the initial algal peak, timing and duration of the subsequent grazer peak, and timing of the algal minimum are consistently accelerated by nutrient enrichment but decelerated by light enrichment (decreasing mixing depth) over the range of intermediate to shallow mixing depths. These contrasting effects of nutrient vs. light enrichment are consequences of their opposing influences on food quality (algal nutrient content): algal productivity and food quality are positively related along a nutrient gradient but inversely related along a light gradient. Light enrichment therefore slows down grazer growth relative to algal growth, decelerating oscillatory dynamics; nutrient enrichment has opposite effects. We manipulated nutrient supply and mixing depth in a field enclosure experiment. The experimental results were qualitatively much more consistent with the flexible than with the fixed stoichiometry model. Nutrient enrichment increased Daphnia peak biomass, decreased algal minimum biomass, decreased the seston C:P ratio, and accelerated transient oscillatory dynamics. Light enrichment (decreasing mixing depth) produced the opposite patterns, except that Daphnia peak biomass increased monotonously with light enrichment, too. Thus, while the model predicts the possibility of the "paradox of energy enrichment" (a decrease in grazer biomass with light enrichment) at high light and low nutrient supply, this phenomenon did not occur in our experiment.