Phosphorus stoichiometric homeostasis of submerged macrophytes and associations with interspecific interactions and community stability in Erhai Lake, China.

According to stoichiometric homeostasis theory, eutrophication is expected to increase the dominance of submerged macrophytes with low homeostatic regulation coefficients (H) relative to those with high H values, ultimately reducing macrophyte community stability. However, empirical evidence supporting this hypothesis is limited. In this study, we conducted a three-year tracking survey (seven sampling events) at 81 locations across three regions of Erhai Lake. We assessed the H values of submerged macrophyte species, revealing significant H values for phosphorus (P) and strong associations of HP values (range: 1.58-2.94) with species and community stability. Moreover, in plots simultaneously containing the dominant high-HP species, Potamogeton maackianus, and its low-HP counterpart, Ceratophyllum demersum, we explored the relationships among eutrophication, interspecific interaction shifts, and community dynamics. As the environmental P concentration increased, the dominance of P. maackianus decreased, while that of C. demersum increased. This shift coincided with reductions in community HP and stability. Our study underpins the effectiveness of H values for forecasting interspecific interactions among submerged macrophytes, thereby clarifying how eutrophication contributes to the decline in stability of the submerged macrophyte community.

Re-oligotrophication and warming stabilize phytoplankton networks.

Phytoplankton taxa are strongly interconnected as a network, which could show temporal dynamics and non-linear responses to changes in drivers at both seasonal and long-term scale. Using a high quality dataset of 20 Danish lakes (1989-2008), we applied extended Local Similarity Analysis to construct temporal network of phytoplankton communities for each lake, obtained sub-network for each sampling month, and then measured indices of network complexity and stability for each sub-network. We assessed how lake re-oligotrophication, climate warming and grazers influenced the temporal dynamics on network complexity and stability of phytoplankton community covering three aspects: seasonal trends, long-term trends and detrended variability. We found strong seasonality for the complexity and stability of phytoplankton network, an increasing trend for the average degree, modularity, nestedness, persistence and robustness, and a decreasing trend for connectance, negative:positive interactions and vulnerability. Our study revealed a cascading effect of lake re-oligotrophication, climate warming and zooplankton grazers on phytoplankton network stability through changes in network complexity characterizing diversity, interactions and topography. Network stability of phytoplankton increased with average degree, modularity, nestedness and decreased with connectance and negative:positive interactions. Oligotrophication and warming stabilized the phytoplankton network (enhanced robustness, persistence and decreased vulnerability) by enhancing its average degree, modularity, nestedness and by reducing its connectance, while zooplankton richness promoted stability of phytoplankton network through increases in average degree and decreases in negative interactions. Our results further indicate that the stabilization effects might lead to more closed, compartmentalized and nested interconnections especially in the deeper lakes, in the warmer seasons and during bloom periods. From a temporal dynamic network view, our findings highlight stabilization of the phytoplankton community as an adaptive response to lake re-oligotrophication, climate warming and grazers.

Consistent stoichiometric long-term relationships between nutrients and chlorophyll-a across shallow lakes.

Aquatic ecosystems are threatened by eutrophication from nutrient pollution. In lakes, eutrophication causes a plethora of deleterious effects, such as harmful algal blooms, fish kills and increased methane emissions. However, lake-specific responses to nutrient changes are highly variable, complicating eutrophication management. These lake-specific responses could result from short-term stochastic drivers overshadowing lake-independent, long-term relationships between phytoplankton and nutrients. Here, we show that strong stoichiometric long-term relationships exist between nutrients and chlorophyll a (Chla) for 5-year simple moving averages (SMA, median R² = 0.87) along a gradient of total nitrogen to total phosphorus (TN:TP) ratios. These stoichiometric relationships are consistent across 159 shallow lakes (defined as average depth < 6 m) from a cross-continental, open-access database. We calculate 5-year SMA residuals to assess short-term variability and find substantial short-term Chla variation which is weakly related to nutrient concentrations (median R² = 0.12). With shallow lakes representing 89% of the world's lakes, the identified stoichiometric long-term relationships can globally improve quantitative nutrient management in both lakes and their catchments through a nutrient-ratio-based strategy.

Ecological implications of fish removal: Insights from gut-content analysis of roach (Rutilus rutilus) and European perch (Perca fluviatilis) in a eutrophic shallow lake.

Large reductions in fish biomass are common both as a method of managing lake ecosystems by fish removals (biomanipulation) and as naturally occurring fish kills. To further understand how fish reductions change feeding patterns of fish, we studied the diets of small- to medium-sized roach (Rutilus rutilus) and European perch (Perca fluviatilis) on a monthly basis using gut-content analysis during an 18-month period before and after a whole-lake fish removal in a eutrophic shallow lake. Further, we performed in-depth analyses of zoobenthos communities of the profundal and littoral zones, as well as analysed the zooplankton community in the littoral and pelagic parts of the lake to estimate abundance and biomass of potential diet items. We found that, in general, there was a trend toward increased zoobenthivory in both species and among all-sized fish after fish removal, regardless of prior diet preference. Reduced piscivory among larger perch (>150 mm) and reduced zooplanktivory among smaller perch and roach (<150 mm) were also observed. Moreover, during a short period of high zooplankton biomass after fish removal, both perch and roach (all sizes) shifted their diet toward daphnids, which likely caused a decrease in daphnid population. We suggest that such change toward periodical zooplanktivory across fish species and size groups may lead to unexpectedly high top-down control by fish after lake restoration by fish removal.

Bimodality and alternative equilibria do not help explain long-term patterns in shallow lake chlorophyll-a.

Since its inception, the theory of alternative equilibria in shallow lakes has evolved and been applied to an ever wider range of ecological and socioecological systems. The theory posits the existence of two alternative stable states or equilibria, which in shallow lakes are characterised by either clear water with abundant plants or turbid water where phytoplankton dominate. Here, we used data simulations and real-world data sets from Denmark and north-eastern USA (902 lakes in total) to examine the relationship between shallow lake phytoplankton biomass (chlorophyll-a) and nutrient concentrations across a range of timescales. The data simulations demonstrated that three diagnostic tests could reliably identify the presence or absence of alternative equilibria. The real-world data accorded with data simulations where alternative equilibria were absent. Crucially, it was only as the temporal scale of observation increased (>3 years) that a predictable linear relationship between nutrient concentration and chlorophyll-a was evident. Thus, when a longer term perspective is taken, the notion of alternative equilibria is not required to explain the response of chlorophyll-a to nutrient enrichment which questions the utility of the theory for explaining shallow lake response to, and recovery from, eutrophication.

The impact of climate change and eutrophication on phosphorus forms in sediment: Results from a long-term lake mesocosm experiment.

Characteristics of bottom sediments in lake mesocosms 11 years after starting the experiment were studied in order to determine the effects of nutrient loading, temperature increase and vegetation type on concentration and vertical distribution of phosphorus (P) forms. The experimental setup consisted of 24 outdoor flow-through mesocosms with two nutrient treatments - low (L) and high (H) and 3 temperature levels - ambient (T0), heated by 2-4 °C (T1) and 3-6 °C (T2) in four replicates. Thickness of the organic sediment was measured and the sediment analysed for dry weight, organic matter, and P fractions (according to a sequential extraction scheme) and organic P compounds (by 31P nuclear magnetic resonance spectroscopy). Higher nutrient loading led to increased sediment accumulation and higher concentration of total P and most P fractions, except P bound to aluminium and humic matter. The dominant vegetation type covaried with nutrient levels. Vertical gradients in Ca bound P and mobile P in low nutrient mesocosms was perhaps a result of P coprecipitation with calcite on macrophytes and P uptake by roots indicating that in macrophyte-rich lakes, plants can be important modifiers of early P diagenesis. Temperature alone did not significantly affect sediment accumulation rate but the interaction effect between nutrient and temperature treatments was significant. At high nutrient loading, sediment thickness decreased with increasing temperature, but at low nutrient loading, it increased with warming. The effect of warming on sediment composition became obvious only in nutrient enriched mesocosms showing that eutrophication makes shallow lake ecosystems more susceptible to climate change.

Diet and food selection by fish larvae in turbid and clear water shallow temperate lakes.

Fish larvae play an important structuring role for their prey and show ontogenetic shifts in diet. Changes in diet differ between species and habitats and may also be affected by turbidity (eutrophication). We investigated the diet (stomach content) and the food selection (ratio of ingested prey and prey availability) of roach and perch larvae in a clear lake and of roach, perch and pikeperch larvae in a turbid lake multiple times during spring to autumn. The diet of the fish larvae changed with size, and for roach and perch larvae between the lakes. Coexisting species of fish larvae had different diets in the two lakes, pointing to resource partitioning; yet, in the clear lake, medium-sized larvae had a high diet overlap, suggesting a competitive relationship at this developmental stage. In the clear lake, roach larvae showed diel differentiation in diet, while perch demonstrated diet shifts between habitats, which probably aided in reducing competition and also evidenced an effect of light on the larval prey capture and/or predator-fish larvae interactions. In the turbid lake, roach and perch larvae did not reveal differences in diet between habitats or time of the day, owing to homogeneity of food items and poor light conditions. However, the diet of pikeperch larvae differed between day and night following daily variations in the abundance of its preferred prey. The roach larvae were highly selective for Bosmina, Daphnia and benthic cladocerans, perch larvae generally consumed what was available, while pikeperch primarily preyed on cyclopoid copepodites. We conclude that turbidity acted as a cover for fish larvae in the turbid lake. Under eutrophication-induced turbidity scenarios the effects of fish larvae on their prey are stronger (i.e., high selectivity for several resources) than that of larvae in clear waters, creating a negative feedback on the path to restore water clarity.

Abiotic and biotic drivers of temporal dynamics in the spatial heterogeneity of zooplankton communities across lakes in recovery from eutrophication.

Seasonal and annual dynamics of the zooplankton community in lakes are affected by changes in abiotic drivers, trophic interactions (e.g., changes in phytoplankton and fish communities and abundances) and habitat characteristics (e.g. macrophyte abundance and composition). However, little is known about the temporal responses of the zooplankton community to abiotic and biotic drivers across lakes at the regional scale. Using a comprehensive 20-year dataset from 20 Danish lakes in recovery from eutrophication, we assessed the seasonal and annual trends in the spatial heterogeneity of zooplankton community across lakes and related it to abiotic and biotic drivers. We found significant seasonality and inter-annual decreases in spatial zooplankton heterogeneity in both shallow and deep lakes, with the decrease in the spatial turnover dominating the temporal dynamics of the beta diversity. For the inter-annual changes, decreased spatial heterogeneity of phytoplankton, macrophytes and fish were important biotic drivers at the regional scale. Using a series of ordinary least squares regressions and model selection with model averaging approaches, we revealed that both local (e.g., total phosphorus, total nitrogen, pH, Secchi depth, alkalinity, Schmidt stability, water temperature) and regional drivers (e.g., air temperature, solar irradiance) were important variables influencing the spatial zooplankton heterogeneity, although the directions depended on the beta diversity measures and water depth. Our results highlight an important role of bottom-up forces through phytoplankton community as well as macrophytes and top-down forces via fishes in driving the temporal changes in zooplankton community composition patterns at the regional scale.

Seasonal and long-term trends in the spatial heterogeneity of lake phytoplankton communities over two decades of restoration and climate change.

World-wide, reducing the external nutrient loading to lakes has been the primary priority of lake management in the restoration of eutrophic lakes over the past decades, and as expected this has resulted in an increase in the local environmental heterogeneity, and thus biotic heterogeneity, within lakes. However, little is known about how the regional spatial heterogeneity of lake biotic communities changes with restoration across a landscape. Using a long-term monitoring dataset from 20 Danish lakes, we elucidated the seasonal and long-term trends in the spatial heterogeneity of climate, local abiotic variables and phytoplankton communities over two decades of restoration and climate change at landscape level. We found significant seasonality in the spatial heterogeneity of most climatic and local drivers as well as in the total beta diversity (Sørensen coefficient) and its turnover components (Simpson coefficient) of phytoplankton communities among the lakes. The seasonality tended to be less marked in deep than in shallow lakes. We found significant spatial homogenisation of most local drivers (except for alkalinity) and phytoplankton communities after two decades of restoration and that turnover dominated the temporal responses of the total beta diversity of phytoplankton communities. Path analyses showed that the homogenisation of phytoplankton communities was mainly due to a decrease in spatial heterogeneity of total phosphorus and Schmidt stability in shallow lakes and to a decrease in spatial total phosphorus and total nitrogen heterogeneity in deep lakes. However, albeit weakly, the spatial heterogeneity of the phytoplankton communities was affected indirectly by climatic warming in both shallow and deep lakes and directly by wind speed in shallow lakes. We conclude that restoration of eutrophic lakes may lead to an increase in the local heterogeneity of phytoplankton communities at lake scale and an increase in homogeneity at landscape scale.

Effects of warming and nutrients on the microbial food web in shallow lake mesocosms.

We analysed changes in the abundance, biomass and cell size of the microbial food web community (bacteria, heterotrophic nanoflagellates, ciliates) at contrasting nutrient concentrations and temperatures during a simulated heat wave. We used 24 mesocosms mimicking shallow lakes in which two nutrient levels (unenriched and enriched by adding nitrogen and phosphorus) and three different temperature scenarios (ambient, IPCC A2 scenario and A2+%50) are simulated (4 replicates of each). Experiments using the mesocosms have been running un-interrupted since 2003. A 1-month heat wave was imitated by an extra 5 °C increase in the previously heated mesocosms (from 1st July to 1st August 2014). Changes in water temperature induced within a few days a strong effect on the microbial food web functioning, demonstrating a quick response of microbial communities to the changes in environment, due to their short generation times. Warming and nutrients showed synergistic effects. Microbial assemblages of heterotrophic nanoflagellates and ciliates responded positively to the heating, the increase being largest in the enriched mesocosms. The results indicate that warming and nutrients in combination can set off complex interactions in the microbial food web functioning.

Response of Submerged Macrophyte Communities to External and Internal Restoration Measures in North Temperate Shallow Lakes.

Submerged macrophytes play a key role in north temperate shallow lakes by stabilizing clear-water conditions. Eutrophication has resulted in macrophyte loss and shifts to turbid conditions in many lakes. Considerable efforts have been devoted to shallow lake restoration in many countries, but long-term success depends on a stable recovery of submerged macrophytes. However, recovery patterns vary widely and remain to be fully understood. We hypothesize that reduced external nutrient loading leads to an intermediate recovery state with clear spring and turbid summer conditions similar to the pattern described for eutrophication. In contrast, lake internal restoration measures can result in transient clear-water conditions both in spring and summer and reversals to turbid conditions. Furthermore, we hypothesize that these contrasting restoration measures result in different macrophyte species composition, with added implications for seasonal dynamics due to differences in plant traits. To test these hypotheses, we analyzed data on water quality and submerged macrophytes from 49 north temperate shallow lakes that were in a turbid state and subjected to restoration measures. To study the dynamics of macrophytes during nutrient load reduction, we adapted the ecosystem model PCLake. Our survey and model simulations revealed the existence of an intermediate recovery state upon reduced external nutrient loading, characterized by spring clear-water phases and turbid summers, whereas internal lake restoration measures often resulted in clear-water conditions in spring and summer with returns to turbid conditions after some years. External and internal lake restoration measures resulted in different macrophyte communities. The intermediate recovery state following reduced nutrient loading is characterized by a few macrophyte species (mainly pondweeds) that can resist wave action allowing survival in shallow areas, germinate early in spring, have energy-rich vegetative propagules facilitating rapid initial growth and that can complete their life cycle by early summer. Later in the growing season these plants are, according to our simulations, outcompeted by periphyton, leading to late-summer phytoplankton blooms. Internal lake restoration measures often coincide with a rapid but transient colonization by hornworts, waterweeds or charophytes. Stable clear-water conditions and a diverse macrophyte flora only occurred decades after external nutrient load reduction or when measures were combined.

Gravel pit lakes in Denmark: Chemical and biological state.

Mining of gravel and sand for construction purposes is big business and gravel pit lakes have become increasingly common all over the world. In Denmark, hundreds of gravel pit lakes have been created during the past decades. We investigated the chemical and biological status of 33-52 gravel pit lakes and compared the results with data from similar-sized natural Danish lakes. The area of the lakes ranged from 0.2 to 13ha and their age from 0.5 to 26years. Generally, the gravel pit lakes were clear with low nutrient concentrations, the median concentrations of total phosphorus and total nitrogen being 0.023mg/l and 0.30mg/l compared with 0.115mg/l and 1.29mg/l, respectively, in natural lakes. Correspondingly, median chlorophyll a was 5µg/l in the gravel pit lakes and 36µg/l in the natural lakes. Submerged macrophytes were found in all gravel pit lakes, with particularly high cover in the shallow ones. Most gravel pit lakes were deeper than the natural lakes, which may restrict the area potentially to be covered by submerged macrophytes, with implications also for the biological quality of the lakes. Fish were found in most of the gravel pit lakes, roach (Rutilus rutilus), perch (Perca fluviatilis) and rudd (Scardinius erythrophalmus) being the most frequently observed species. Fish stocking was common and included also non-native species such as carp (Cyprinus carpio) and rainbow trout (Oncorchynchus mykiss). Compared with the natural lakes, fish species richness and catch per gillnet were overall lower in the gravel pit lakes. Groundwater-fed gravel pit lakes add importantly to the number of high-quality lakes in Denmark and with an optimised design and by avoiding negative side effects, they can be positive for both nature and society.

Spatial and temporal variations reveal the response of zooplankton to cyanobacteria.

The effects of cyanobacteria on zooplankton abundance, structure and diversity were investigated, based on a systematic study on spatial and temporal variations of cyanobacteria and zooplankton in Lake Taihu from 1998 to 2007. It was found that similar increasing trends of cyanobacteria/phytoplankton ratios were accompanied by different trends in biomass, composition and biodiversity of zooplankton in different regions of the lake; the cladocerans benefitted from the increase in cyanobacteria; however, rotifers and protozoans were negatively affected by cyanobacteria. The biomass-based biodiversity of phytoplankton and zooplankton was negatively affected by cyanobacteria as well, and the adverse effects were in proportion to the cyanobacteria/phytoplankton ratio. These results indicated interestingly that higher amounts of cyanobacteria do not necessarily reduce zooplankton biomass, as the biomass of larger zooplankton such as cladocerans was positively related to cyanobacteria. The findings are essential to understand the complex ecological effects of cyanobacteria blooms in lakes.

Response of fish communities to multiple pressures: Development of a total anthropogenic pressure intensity index.

Lakes in Europe are subject to multiple anthropogenic pressures, such as eutrophication, habitat degradation and introduction of alien species, which are frequently inter-related. Therefore, effective assessment methods addressing multiple pressures are needed. In addition, these systems have to be harmonised (i.e. intercalibrated) to achieve common management objectives across Europe. Assessments of fish communities inform environmental policies on ecological conditions integrating the impacts of multiple pressures. However, the challenge is to ensure consistency in ecological assessments through time, across ecosystem types and across jurisdictional boundaries. To overcome the serious comparability issues between national assessment systems in Europe, a total anthropogenic pressure intensity (TAPI) index was developed as a weighted combination of the most common pressures in European lakes that is validated against 10 national fish-based water quality assessment systems using data from 556 lakes. Multi-pressure indices showed significantly higher correlations with fish indices than single-pressure indices. The best-performing index combines eutrophication, hydromorphological alterations and human use intensity of lakes. For specific lake types also biological pressures may constitute an important additional pressure. The best-performing index showed a strong correlation with eight national fish-based assessment systems. This index can be used in lake management for assessing total anthropogenic pressure on lake ecosystems and creates a benchmark for comparison of fish assessments independent of fish community composition, size structure and fishing-gear. We argue that fish-based multiple-pressure assessment tools should be seen as complementary to single-pressure tools offering the major advantage of integrating direct and indirect effects of multiple pressures over large scales of space and time.

Warming and nutrient enrichment in combination increase stochasticity and beta diversity of bacterioplankton assemblages across freshwater mesocosms.

The current climate warming and eutrophication are known to interactively threaten freshwater biodiversity; however, the interactive effects on lacustrine bacterioplankton diversity remain to be determined. Here, we analyzed the spring bacterioplankton community composition (BCC) in 24 outdoor, flow-through mesocosms (mimicking shallow lake environments) under 3 temperature scenarios and 2 nutrient regimes. Our results revealed that neither long-term warming (8.5 years) nor nutrient enrichment had significant effects on bacterioplankton alpha diversity, whereas long-term enhanced warming (elevated 50% above the IPCC A2 climate scenario) and nutrient enrichment in combination increased bacterioplankton beta diversity. We also found that BCC shifted significantly under enhanced warming and nutrient-enriched conditions towards decreased relative abundances of Actinobacteria, Bacteroidetes and Betaproteobacteria, whereas the percentages of Cyanobacteria, total rare phyla and unclassified phyla significantly increased. Null-model tests indicated that deterministic processes played a more important role than stochastic processes in determining BCC. However, the relative importance of stochasticity, primarily ecological drift, was enhanced and contributed to the increased beta diversity of BCC under enhanced warming and nutrient-enriched conditions. Overall, our study suggests that the synergetic effects of warming and nutrient enrichment may result in high variability in the composition of bacterioplankton communities in lacustrine water bodies.

A Systematic Investigation into the Environmental Fate of Microcystins and The Potential Risk: Study in Lake Taihu.

A systematic investigation was conducted in Lake Taihu in autumn of 2013 and 2014, in order to understand the environmental fate of microcystins (MCs) and evaluate the health risk from MCs. Samples of water, algal cells, macrophytes, shrimps and fish were taken to detect MCs by HPLC-MS/MS after solid phase extraction. Widespread MC contamination in water, algal cells, macrophytes, shrimps and fish was found in Lake Taihu. The ubiquitous presence of MCs in water, algal cells and biota was found in 100% of samples. MC accumulation was in the order of primary producer > tertiary consumer > secondary consumer > primary consumer. The highest levels of MCs in macrophytes, shrimps and fish tissue were found in Potamogeton maackianus, Exopalaemon modestus, and Hyporhamphus intermedius, respectively. The MCs level in shrimps and the tissues of three fish species, Neosalanx tangkahkeii taihuensis, Coilia ectenes and silver carp, was closely linked to their dietary exposure. Ceratophyllum demersum L. was an ideal plant for introduction into lakes to protect against Microcystis blooms and MCs, due to its ability to absorb nutrients, accumulate large amounts of MCs and tolerate these toxins compared to other macrophytes. The average daily intakes (ADIs) of MCs for Exopalaemon modestus and three fish species, Coilia ectenes, Hyporhamphus intermedius and Carassius carassius, were all above the tolerable daily intakes (TDI) set by the World Health Organization (WHO), implying there existed potential threats to human health.

Eutrophication effects on greenhouse gas fluxes from shallow-lake mesocosms override those of climate warming.

Fresh waters make a disproportionately large contribution to greenhouse gas (GHG) emissions, with shallow lakes being particular hot spots. Given their global prevalence, how GHG fluxes from shallow lakes are altered by climate change may have profound implications for the global carbon cycle. Empirical evidence for the temperature dependence of the processes controlling GHG production in natural systems is largely based on the correlation between seasonal temperature variation and seasonal change in GHG fluxes. However, ecosystem-level GHG fluxes could be influenced by factors, which while varying seasonally with temperature are actually either indirectly related (e.g. primary producer biomass) or largely unrelated to temperature, for instance nutrient loading. Here, we present results from the longest running shallow-lake mesocosm experiment which demonstrate that nutrient concentrations override temperature as a control of both the total and individual GHG flux. Furthermore, testing for temperature treatment effects at low and high nutrient levels separately showed only one, rather weak, positive effect of temperature (CH4 flux at high nutrients). In contrast, at low nutrients, the CO2 efflux was lower in the elevated temperature treatments, with no significant effect on CH4 or N2 O fluxes. Further analysis identified possible indirect effects of temperature treatment. For example, at low nutrient levels, increased macrophyte abundance was associated with significantly reduced fluxes of both CH4 and CO2 for both total annual flux and monthly observation data. As macrophyte abundance was positively related to temperature treatment, this suggests the possibility of indirect temperature effects, via macrophyte abundance, on CH4 and CO2 flux. These findings indicate that fluxes of GHGs from shallow lakes may be controlled more by factors indirectly related to temperature, in this case nutrient concentration and the abundance of primary producers. Thus, at ecosystem scale, response to climate change may not follow predictions based on the temperature dependence of metabolic processes.

Planktonic ciliate community structure in shallow lakes of lowland Western Europe.

Temperate shallow meso- to eutrophic lakes can exist in one of two alternative states with contrasting foodwebs, referred to as the clear-water and the turbid state. We describe the planktonic ciliate communities of such lakes based on a survey of 66 northwestern European lakes. Ciliates were enumerated and identified to species level according to the quantitative protargol staining technique. Ciliate biomass was on average twice as high in the turbid than in the clear-water lakes. The ciliate communities were dominated by oligotrichs and protostomatids, and no differences in functional composition or α-diversity could be detected between turbid and clear-water lakes, although ß-diversity tended to be higher in the latter. At the species level, however, community structure strongly differed between turbid and clear-water lakes, and several indicator species could be identified for the different lake categories. Variation partitioning showed that nutrient status did not explain ciliate community structure independent of the alternative states, while lake area was identified as an additional structuring factor for the ciliate communities. These results stress the importance of the ecosystem structure in shaping ciliate communities in temperate shallow lakes and suggest that nutrient status has little direct effect on ciliate community structure in such lakes.

Watershed land use effects on lake water quality in Denmark.

Mitigating nutrient losses from anthropogenic nonpoint sources is today of particular importance for improving the water quality of numerous freshwater lakes worldwide. Several empirical relationships between land use and in-lake water quality variables have been developed, but they are often weak, which can in part be attributed to lack of detailed information about land use activities or point sources. We examined a comprehensive data set comprising land use data, point-source information, and in-lake water quality for 414 Danish lakes. By excluding point-source-influenced lakes (n = 210), the strength in relationship (R2) between in-lake total nitrogen (TN) and total phosphorus (TP) concentrations and the proportion of agricultural land use in the watershed increased markedly, from 10-12% to 39-42% for deep lakes and from 10-12% to 21-23% for shallow lakes, with the highest increase for TN. Relationships between TP and agricultural land use were even stronger for lakes with rivers in their watershed (55%) compared to lakes without (28%), indicating that rivers mediate a stronger linkage between landscape activity and lake water quality by providing a "delivery" mechanism for excess nutrients in the watershed. When examining the effect of different near-freshwater land zones in contrast to the entire watershed, relationships generally improved with size of zone (25, 50, 100, 200, and 400 m from the edge of lake and streams) but were by far strongest using the entire watershed. The proportion of agricultural land use in the entire watershed was best in explaining lake water quality, both relative to estimated nutrient surplus at agricultural field level and near-lake land use, which somewhat contrasts typical strategies of management policies that mainly target agricultural nutrient applications and implementation of near-water buffer zones. This study suggests that transport mechanisms within the whole catchment are important for the nutrient export to lakes. Hence, the whole watershed should be considered when managing nutrient loadings to lakes, and future policies should ideally target measures that reduce the proportion of cultivated land in the watershed to successfully improve lake water quality.