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In many lakes, the most conspicuous seasonal events are the phytoplankton spring bloom and the subsequent clear-water phase, a period of low-phytoplankton biomass that is frequently caused by mesozooplankton (Daphnia) grazing. In Central European lakes, the timing of the clear-water phase is linked to large-scale climatic forcing, with warmer winters being followed by an earlier onset of the clear-water phase. Mild winters may favour an early build-up of Daphnia populations, both directly through increased surface temperatures and indirectly by reducing light limitation and enhancing algal production, all being a consequence of earlier thermal stratification. We conducted a field experiment to disentangle the separate impacts of stratification depth (affecting light supply) and temperature on the magnitude and timing of successional events in the plankton. We followed the dynamics of the phytoplankton spring bloom, the clear-water phase and the spring peak in Daphnia abundance in response to our experimental manipulations. Deeper mixing delayed the timing of all spring seasonal events and reduced the magnitudes of the phytoplankton bloom and the subsequent Daphnia peak. Colder temperatures retarded the timing of the clear-water phase and the subsequent Daphnia peak, whereas the timing of the phytoplankton peak was unrelated to temperature. Most effects of mixing depth (light) and temperature manipulations were independent, effects of mixing depth being more prevalent than effects of temperature. Because mixing depth governs both the light climate and the temperature regime in the mixed surface layer, we propose that climate-driven changes in the timing and depth of water column stratification may have far-reaching consequences for plankton dynamics and should receive increased attention.
We independently manipulated mixing intensity (strong artificial mixing vs. background turbulence) and water‐column depth (2 m, 4 m, 8 m, and 12 m) in order to explore their separate and combined effects in a field enclosure experiment. To accentuate the vertical light gradient, enclosures had black walls, resulting in a euphotic depth of only 3.7 m. All enclosures were placed in a well‐mixed water bath to equalize temperature across treatments. Phytoplankton responded to an initial phosphorus pulse with a transient increase in biomass, which was highest in the shallowest, least light‐limited water columns where dissolved mineral phosphorus subsequently became strongly limiting. As a consequence, the depth‐averaged mineral phosphorus concentration increased and the seston carbon (C) : phosphorous (P) ratio decreased with increasing water‐column depth. Low turbulence enclosures became quickly dominated by motile taxa (flagellates) in the upper water column, whereas mixed enclosures became gradually dominated by pennate diatoms, which resulted in higher average sedimentation rates in the mixed enclosures over the 35‐d experimental period. Low turbulence enclosures showed pronounced vertical structure in water columns >4 m, where diversity was higher than in mixed enclosures, suggesting vertical niche partitioning. This interpretation is supported by a primary production assay, where phytoplankton originating from different water depths in low‐turbulence treatments had the relatively highest primary productivity when incubated at their respective depths of origin.
We employed the well-established Horton-Strahler, hierarchical, stream-order (ω) scheme to investigate scaling of nutrient loads (P and N) from ~845 wastewater treatment plants (WWTPs) distributed along the river network in urbanized Weser River, the largest national basin in Germany (~46K km2; ~8.4 million population). We estimated hydrologic and water quality impacts at the reach- and basin-scales, at two steady river discharge conditions (median flow, QR50; low-flow, QR90). Of the five WWTPs class-sizes (1 ≤ k ≤ 5), ~68% discharge to small low-order streams (ω < 3). We found large variations in capacity to dilute WWTP nutrient loads because of variability in (1) treated wastewater discharge (QU) within and among different class-sizes, and (2) river discharge (QR) within low-order streams (ω < 3) resulting from differences in drainage areas. For QR50, reach-scale water quality impairment assessed by nutrient concentration was likely at 136 (~16%) locations for P and 15 locations (~2%) for N. About 90% of these locations were lower-order streams (ω < 3). At QR50 and only with dilution, basin-scale cumulative nutrient loads from multiple upstream WWTPs increase impaired locations to 266 (~32% of total) for P. Considering in-stream uptake decreased P-impaired streams to 225 (~27%), suggesting the dominant role of dilution in the Weser River basin. Role of in-stream uptake diminished along the flow paths, while dilution in larger streams (4 ≤ ω ≤ 7) minimizes the impact of WWTP loads. Under QR90 conditions [(QR50/QR90) ~ 2.5], water quality impaired locations will likely double for the basin-scale analyses. Long-term water quality data suggested that diffuse sources are the primary contributors for water quality impairments in large streams. Our data-modeling synthesis approach is transferable to other urbanized river basins and extends understanding of point source impacts on water quality across spatial scales.
1. Understanding and explaining the structure of communities in response to environmental gradients is a central goal in ecology. Trait‐based approaches are promising but yet rarely applied to understand community dynamics in response to changing environmental conditions.
2. Here, we investigate seasonal succession patterns of functional traits in phytoplankton communities and how nutrient reductions (oligotrophication) alter these patterns. We used phytoplankton data from 40 years of observation from the Rappbode Reservoir (Germany), which underwent a strong shift in trophic conditions, and translated taxonomic composition into functional traits by assigning trait values compiled from the literature.
3. All studied traits (morphological, behavioural and physiological traits) responded to changing environmental conditions and showed consistent, reoccurring seasonal developments. The seasonal succession of phytoplankton communities was shaped by a trade‐off between small‐celled, fast‐growing species that are able to rapidly incorporate existing resources (r ‐strategists) and large‐celled species with more complex and efficient mechanisms to exploit scarce mineral nutrients or acquire previously unexploited nutrient pools (k ‐strategists). In summer, when nutrients were scarce, the k ‐strategy was prevailing (important traits: phosphate affinity, nitrogen fixation, motility and mixotrophy). During the rest of the year, nutrients and turbulence were high and r ‐strategists dominated (important traits: maximum growth rate and light affinity).
4. A comparison between eutrophic and oligotrophic years revealed that the main features of functional trait succession were largely preserved, but intra‐annual fluctuations from spring to summer were stronger during eutrophic years. Nutrient reductions mainly affected functional traits and biomass in spring, while in summer the functional community composition changed little.
5. Synthesis. This study provides for the first time a quantitatively supported functional template for trait‐based succession patterns in lakes under different nutrient conditions. By translating taxonomic composition into trait information, we demonstrate that the quantification of functional characteristics enables ecological interpretation of observed community dynamics and provides not only a testable template but also a powerful tool towards a more mechanistic understanding. The quantification of functional traits further improves the predictability of community shifts in response to changing environmental conditions and thus opens new perspectives for predictive limnology using lake ecosystem models.
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.
Die Charakterisierung und Quantifizierung von Phytoplankton-Gemeinschaften sind ein wesentlicher Aspekt der Gütebeurteilung von Standgewässern. In dieser Arbeit wird eine neue Herangehensweise hergeleitet und demonstriert, welche die klassische, taxonomisch-orientierte Methodik durch einen funktionalen Ansatz ergänzt, der auf den physiologischen und ökologischen Eigenschaften der Taxa (Traits) beruht. Kern des Ansatzes bildet der sogenannte Trait-Schlüssel, mit dem eine taxonomische Information in eine funktionale Information übersetzt werden kann. Die Aussagekraft und Potenziale des Verfahrens werden anhand von Langzeitdaten (50 Jahre) der Phytoplankton-Gemeinschaft der Rappbode-Talsperre exemplarisch demonstriert. Die Trait-basierte Analyse reduziert die Komplexität der Daten und ermöglicht eine quantitative Analyse der beobachteten Veränderungen auf der Basis von statistischen Methoden, die auf den klassischen taxonomischen Ansatz nicht anwendbar sind. Sowohl hinsichtlich der saisonalen Sukzession als auch der langfristigen Trends lassen die funktionalen Eigenschaften Rückschlüsse auf die maßgeblichen Umweltfaktoren zu. Für die wasserwirtschaftliche Praxis erweitert sich damit der Informationsgehalt und die Interpretierbarkeit der vorhandenen Daten. Diese Aufwertung der Ergebnisse kann ohne Mehraufwand erhalten werden. Von besonderem Interesse sind hierbei Kausalanalysen signifikanter Langzeittrends oder von sprunghaften Veränderungen in der Phytoplankton-Entwicklung, die hiermit in einen kausalen und systemanalytischen Zusammenhang gestellt werden können.
Transient dynamics of pelagic producer-grazer systems in a gradient of nutrients and mixing depths
(2008)
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.
Physical determinants of phytoplankton production, algal stoichiometry, and vertical nutrient fluxes
(2010)
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.
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.
In shallow aquatic systems, benthic and pelagic primary producers typically compete for light and nutrients along opposing vertical supply axes: pelagic algae shade the benthic habitat; conversely, benthic algae intercept the nutrient flux from the sediment to the pelagic habitat. We present a general framework for analyzing such spatially asymmetric resource competition across habitat boundaries using a mechanistic, dynamical model. We visualize the mechanisms determining the outcome of these cross‐habitat interactions using zero‐net‐growth isoclines, resource supply points, and resource consumption vectors. In extensive invasion analyses, we characterize the abiotic and competitive persistence boundaries of pelagic and benthic primary producers, which are set by environmental factors determining nutrient and light supply and are modified by resource use by the competitor in the respective other habitat.
We note several qualitative differences between cross‐habitat and “classical” within‐habitat resource competition. First, coexistence of cross‐habitat competitors is facilitated by, but does not require niche differentiation with respect to, the utilization of resources. Because each species has a competitive edge for the resource that is supplied from “its” side of the system, a competitor that is inferior in utilizing both resources can sometimes coexist with, or even exclude, a superior competitor. Second, increasing the external supply of one resource (the nutrient) may initially favor both competitors, until a breakpoint is reached where the benthic producer goes abruptly extinct. Finally, whether a given pair of cross‐habitat competitors coexist or shows alternative states may depend on the environment. Specifically, benthic and pelagic algae may coexist at low nutrient and light supply but produce alternative states at high nutrient and light supply. Alternative states are, in turn, promoted by any algal trait combination that increases the spatial asymmetry in resource consumption, i.e., leads to a higher nutrient consumption in the benthic habitat and/or a higher light consumption in the pelagic habitat.
In a first empirical application, we show that predictions from our model give a good fit to published data on benthic and pelagic primary production in temperate and arctic lakes spanning a broad range of nutrient environments.