Warming alters the network of physiological traits and their contribution to plant abundance.

The impact of global warming on plant abundance has been widely discussed, but it remains unclear how warming affects plant physiological traits, and how these traits contribute to the abundance of aquatic plants. We explored the adjustments in physiological traits of two common aquatic plant species (Potamogeton crispus L. and Elodea canadensis Michx.) and their links to plant abundance in three temperature treatments by determining twelve physiological traits and plant abundance over an 11-month period in outdoor mesocosms. This mesocosms facility has been running uninteruptedly for 16 years, rendering the plants a unique opportunity to adapt to the warming differences. We found that 1) warming reduced the starch storage in winter for P. crispus and in summer for E. canadensis while increased the nitrogenous substances (e.g., TN, FAA, and proline) in winter for P. crispus. 2) For E. canadensis, TC, starch, SC, and sucrose contents were higher in summer than in winter regardless of warming, while TC, SC, and sucrose contents were lower in summer for P. crispus. 3) Warming decreased the association strength between physiological traits and plant abundance for P. crispus but enhanced it for E. canadensis. 4) E. canadensis showed increased interaction strength among physiological traits under warming, indicating increased metabolic exertion in the response to warming, which contributed to the reduction in abundance. Trait interaction strength of P. crispus was reduced under warming, but with less impact on plant abundance compared with E. canadensis. Our study emphasizes that warming alters the network of plant physiological traits and their contribution to abundance and that different strengths of susceptibility to warming of the various plant species may alter the composition of plant communities in freshwater ecosystems.

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.

Climate-associated variation in the drivers of benthic macroinvertebrate species-area relationships across shallow freshwater lakes.

The island species-area relationship (ISAR) describes how species richness increases with increasing area of a given island or island-like habitat, such as freshwater lakes. While the ISAR is one of the most common phenomena observed in ecology, there is variation in both the form of the relationship and its underlying mechanisms. We compiled a global data set of benthic macroinvertebrates from 524 shallow freshwater lakes, ranging from 1 to 293,300 ha in area. We used individual-based rarefaction to determine the degree to which ISAR was influenced by mechanisms other than passive sampling (larger islands passively sample more individuals from the regional pool and, therefore, have more species than smaller islands), which would bias results away from expected relationships between rarefied species richness (and other measures that capture relative abundances) and lake area. We also examined how climate may alter the shape of the ISARs. We found that both rarefied species richness (the number of species standardized by area or number of individuals) and a measure of evenness emphasizing common species exhibit shallow slopes in relationships with lake area, suggesting that the expected ISARs in these lakes most likely result from passive sampling. While there was considerable variation among ISARs across the investigated lakes, we found an overall positive rarefied ISAR for lakes in warm (i.e. tropical/subtropical) regions (n = 195), and in contrast, an overall negative rarefied ISAR in cool (i.e. north temperate) lakes (n = 329). This suggested that mechanisms beyond passive sampling (e.g. colonization-extinction dynamics and/or heterogeneity) were more likely to operate in warm lakes. One possible reason for this difference is that the area-dependent intensity of fish predation, which can lead to flatter ISARs, is weaker in warmer relative to cooler lakes. Our study illustrates the importance of understanding both the pattern and potential processes underlying the ISARs of freshwater lakes in different climatic regions. Furthermore, it provides a baseline for understanding how further changes to the ecosystem (i.e. in lake area or climate) might influence biodiversity patterns.

Benthic primary production decreases internal phosphorus loading from lake sediments under light supplement.

In aquatic ecosystems, light penetrating the sediment surface in shallow lakes may regulate the internal phosphorus (P) release through benthic primary production, which subsequently affects oxidation, pH levels, and alkaline phosphatase activity in the upper sediment. To study the effects of light exposure on the P dynamics at the sediment-water interface under eutrophic conditions, a two-month mesocosm experiment was conducted in twelve cement tanks (1000 L each). The tanks were equipped with Light-Emitting Diode (LED) lights, and surface sediments collected from eutrophic Lake Nanhu (China) were exposed to four different light intensities (0, 50, 100, 200 µmol m-2 s-1). The results revealed that: 1) Both the total phosphorus concentration and the phosphorus release flux from the sediment were lower in the light treatments (mean value, 0.59-0.71 mg L-1 and 0.00-0.01 mg m-2 d-1, respectively) than in the control treatment (0.77 mg L-1 and 0.01 mg m-2 d-1, respectively), indicating that light supplement could decrease the internal P release. 2) Benthic primary production promoted by light directly absorbed soluble reactive phosphorus and decreased the internal P release. The resulting improved production could also increase dissolved oxygen concentrations at the sediment-water interface, thus indirectly inhibiting internal P release. 3) The relative contributions of direct absorption and indirect inhibition on the internal P release ranged between 23% to 69% and 31% to 77% depending on the light intensity.

Mechanisms of high ammonium loading promoted phosphorus release from shallow lake sediments: A five-year large-scale experiment.

The unprecedented global increase in the anthropogenic-derived nitrogen (N) input may have profound effects on phosphorus (P) dynamics and may potentially lead to enhanced eutrophication as demonstrated in short-term mesocosm experiments. However, the role of N-influenced P release is less well studied in large-scale ecosystems. To gain more insight into ecosystem effects, we conducted a five-year large-scale experiment in ten ponds (700-1000 m2 each) with two types of sediments and five targeted total N concentrations (TN) by adding NH4Cl fertilizer (0.5, 1, 5, 10, and 25 mg N L-1). The results showed that: (ⅰ) The sediment P release increased significantly when TN exceeded 10-25 mg N L-1. (ⅱ) The most pronounced sediment P release increase occurred in summer and from sediments rich in organic matter (OMSed). (ⅲ) TN, algal biomass, fish biomass, non-algal turbidity, sediment pH, and OMSed were the dominant factors explaining the sediment P release, as suggested by piecewise structural equation modeling. We propose several mechanisms that may have stimulated P release, i.e. high ammonium input causes a stoichiometric N:P imbalance and induce alkaline phosphatase production and dissolved P uptake by phytoplankton, leading to enhanced inorganic P diffusion gradient between sediment and water; higher pelagic fish production induced by the higher phytoplankton production may have led increased sediment P resuspension through disturbance; low oxygen level in the upper sediment caused by nitrification and organic decomposition of the settled phytoplankton and, finally, long-term N application-induced sediment acidification as a net effect of ammonium hydrolysis, nitrification, denitrification; The mechanisms revealed by this study shed new light on the complex processes underlying the N-stimulated sediment P release, with implications also for the strategies used for restoring eutrophicated lakes.

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.

Effects of DOC addition from different sources on phytoplankton community in a temperate eutrophic lake: An experimental study exploring lake compartments.

A mesocosm experiment was conducted in a temperate eutrophic lake with the hypotheses: 1) the addition of a labile form of DOC would trigger a more pronounced response in phytoplankton biomass and composition compared with a non-labile form; 2) DOC addition would increase phytoplankton biomass by co-inserting organic nutrients for phytoplankton growth; 3) DOC addition would change phytoplankton composition, in particular towards mixotrophic taxa due to higher DOC availability; and that 4) there would be differences in phytoplankton responses to DOC addition, depending on whether sediment was included or not. We used two types of mesocosms: pelagic mesocosms with closed bottom, and benthic mesocosms open to the sediment. The experiment ran for 29 days in total. The DOC addition occurred once, at Day 1. Besides the control, there were two treatments: HuminFeed® (non-labile DOC) at a concentration of 2 mg L-1, and a combination of 2 mg L-1 HuminFeed® and 2 mg L-1 DOC from alder leaf leachate (labile). Responses were detected only in the treatment with alder leaf extract. Ecosystem processes responded immediately to DOC addition, with the fall in dissolved oxygen and pH indicating an increase in respiration, relative to primary production (Day 2). In contrast, there was a delay of a few days in structural responses in the phytoplankton community (Day 6). Phytoplankton biomass increased after DOC addition, probably boosted by the phosphorus released from alder leaf extract. Changes in phytoplankton composition towards mixotrophic taxa were not as strong as changes in biomass, and happened only in the pelagic mesocosms. With the DOC addition, diatoms prevailed in benthic mesocosms, while the contribution of colonial buoyant cyanobacteria increased in the pelagic ones. This study points towards the necessity to look in greater detail at specific responses of phytoplankton to DOC concentration increases considering lake-habitat and sediment influence.

Responses of coastal sediment phosphorus release to elevated urea loading.

Increased urea is one of the common nitrogen forms polluting coastal waters and affecting nutrient dynamics. To investigate the effects of urea on sediment phosphorus (P) release, we carried out a 2-month mesocosm experiment with six targeted loadings of urea (0-0.6 mg N L-1 d-1). Results showed that: i) urea was rapidly transformed into ammonium and then nitrate (NO3-). ii) When nitrogen occurred as urea or ammonium, minor P release was observed. iii) After urea were mostly converted to NO3-, P release became clearer. iv) NO3- had a dual effect by promoting P release through decreasing sediment pH and increasing alkaline phosphatase activity or by inhibiting P release through improving sediment oxidation. v) The overall effects of urea on P release depended on the ultimate NO3- concentrations, being prominent when NO3- ≥ 11 mg N L-1. Our findings are of relevance when determining nitrogen reduction targets needed for combating eutrophication.

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.

Effects of nitrate on phosphorus release from lake sediments.

Phosphorus (P) release from sediment is a key process affecting the effectiveness of eutrophication mitigation. We hypothesized that high nitrate (NO3-) input may have dual effect on sediment P release: reduce the sediment P release by improving the oxidation of sediment or promote P release by stimulating the growth of phytoplankton and increase the decomposition rates and oxygen consumption at the sediment water interface. To test the effect of different NO3- concentrations, we conducted a three-month experiment in 15 cement tanks (1 m3), with five targeted concentrations of NO3-: control, 2 mg L-1, 5 mg L-1, 10 mg L-1, and 15 mg L-1. The results showed that: i) when NO3- was maintained at high levels: NO3-≥5-7 mg L-1 (range of median values), there was no effect of NO3- on net P release from the sediment, likely because the positive effects of NO3- (increasing oxidation) was counteracted by a promotion of phytoplankton growth. ii) after NO3- addition was terminated NO3- dropped sharply to a low level (NO3-≤0.4 mg L-1), followed by a minor P release in the low N treatments but a significant P release in the high N treatments, which likely reflect that the inhibition effect of NO3- on P release decreased, while the promotion effects at high NO3- concentrations continued. The results thus supported our hypotheses of a dual effect on sediment P release and suggest dose-dependent effect of NO3- loading on stimulating P release from the sediment, being clear at high NO3- exceeding 5-7 mg L-1.

Impact of nutrients and water level changes on submerged macrophytes along a temperature gradient: A pan-European mesocosm experiment.

Submerged macrophytes are of key importance for the structure and functioning of shallow lakes and can be decisive for maintaining them in a clear water state. The ongoing climate change affects the macrophytes through changes in temperature and precipitation, causing variations in nutrient load, water level and light availability. To investigate how these factors jointly determine macrophyte dominance and growth, we conducted a highly standardized pan-European experiment involving the installation of mesocosms in lakes. The experimental design consisted of mesotrophic and eutrophic nutrient conditions at 1 m (shallow) and 2 m (deep) depth along a latitudinal temperature gradient with average water temperatures ranging from 14.9 to 23.9°C (Sweden to Greece) and a natural drop in water levels in the warmest countries (Greece and Turkey). We determined percent plant volume inhabited (PVI) of submerged macrophytes on a monthly basis for 5 months and dry weight at the end of the experiment. Over the temperature gradient, PVI was highest in the shallow mesotrophic mesocosms followed by intermediate levels in the shallow eutrophic and deep mesotrophic mesocosms, and lowest levels in the deep eutrophic mesocosms. We identified three pathways along which water temperature likely affected PVI, exhibiting (a) a direct positive effect if light was not limiting; (b) an indirect positive effect due to an evaporation-driven water level reduction, causing a nonlinear increase in mean available light; and (c) an indirect negative effect through algal growth and, thus, high light attenuation under eutrophic conditions. We conclude that high temperatures combined with a temperature-mediated water level decrease can counterbalance the negative effects of eutrophic conditions on macrophytes by enhancing the light availability. While a water level reduction can promote macrophyte dominance, an extreme reduction will likely decrease macrophyte biomass and, consequently, their capacity to function as a carbon store and food source.

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.

Predicting ecosystem state changes in shallow lakes using an aquatic ecosystem model: Lake Hinge, Denmark, an example.

In recent years, considerable efforts have been made to restore turbid, phytoplankton-dominated shallow lakes to a clear-water state with high coverage of submerged macrophytes. Various dynamic lake models with simplified physical representations of vertical gradients, such as PCLake, have been used to predict external nutrient load thresholds for such nonlinear regime shifts. However, recent observational studies have questioned the concept of regime shifts by emphasizing that gradual changes are more common than sudden shifts. We investigated if regime shifts would be more gradual if the models account for depth-dependent heterogeneity of the system by including the possibility of vertical gradients in the water column and sediment layers for the entire depth. Hence, bifurcation analysis was undertaken using the 1D hydrodynamic model GOTM, accounting for vertical gradients, coupled to the aquatic ecosystem model PCLake, which is implemented in the framework for aquatic biogeochemical modeling (FABM). First, the model was calibrated and validated against a comprehensive data set covering two consecutive 7-yr periods from Lake Hinge, a shallow, eutrophic Danish lake. The autocalibration program Auto-Calibration Python (ACPy) was applied to achieve a more comprehensive adjustment of model parameters. The model simulations showed excellent agreement with observed data for water temperature, total nitrogen, and nitrate and good agreement for ammonium, total phosphorus, phosphate, and chlorophyll a concentrations. Zooplankton and macrophyte coverage were adequately simulated for the purpose of this study, and in general the GOTM-FABM-PCLake model simulations performed well compared with other model studies. In contrast to previous model studies ignoring depth heterogeneity, our bifurcation analysis revealed that the spatial extent and depth limitation of macrophytes as well as phytoplankton chlorophyll-a responded more gradually over time to a reduction in the external phosphorus load, albeit some hysteresis effects still appeared. In a management perspective, our study emphasizes the need to include depth heterogeneity in the model structure to more correctly determine at which external nutrient load a given lake changes ecosystem state to a clear-water condition.

Macrophyte assessment in European lakes: Diverse approaches but convergent views of 'good' ecological status.

The European Water Framework Directive has been adopted by Member States to assess and manage the ecological integrity of surface waters. Specific challenges include harmonizing diverse assessment systems across Europe, linking ecological assessment to restoration measures and reaching a common view on 'good' ecological status. In this study, nine national macrophyte-based approaches for assessing ecological status were compared and harmonized, using a large dataset of 539 European lakes. A macrophyte common metric, representing the average standardized view of each lake by all countries, was used to compare national methods. This was also shown to reflect the total phosphorus (r2 = 0.32), total nitrogen (r2 = 0.22) as well as chlorophyll-a (r2 = 0.35-0.38) gradients, providing a link between ecological data, stressors and management decisions. Despite differing assessment approaches and initial differences in classification, a consensus was reached on how type-specific macrophyte assemblages change across the ecological status gradient and where ecological status boundaries should lie. A marked decline in submerged vegetation, especially Charophyta (characterizing 'good' status), and an increase in abundance of free-floating plants (characterizing 'less than good' status) were the most significant changes along the ecological status gradient. Macrophyte communities of 'good' status lakes were diverse with many charophytes and several Potamogeton species. A large number of taxa occurred across the entire gradient, but only a minority dominated at 'less than good' status, including filamentous algae, lemnids, nymphaeids, and several elodeids (e.g., Zannichellia palustris and Elodea nuttallii). Our findings establish a 'guiding image' of the macrophyte community at 'good' ecological status in hard-water lakes of the Central-Baltic region of Europe.

Global patterns in the metacommunity structuring of lake macrophytes: regional variations and driving factors.

We studied community-environment relationships of lake macrophytes at two metacommunity scales using data from 16 regions across the world. More specifically, we examined (a) whether the lake macrophyte communities respond similar to key local environmental factors, major climate variables and lake spatial locations in each of the regions (i.e., within-region approach) and (b) how well can explained variability in the community-environment relationships across multiple lake macrophyte metacommunities be accounted for by elevation range, spatial extent, latitude, longitude, and age of the oldest lake within each metacommunity (i.e., across-region approach). In the within-region approach, we employed partial redundancy analyses together with variation partitioning to investigate the relative importance of local variables, climate variables, and spatial location on lake macrophytes among the study regions. In the across-region approach, we used adjusted R2 values of the variation partitioning to model the community-environment relationships across multiple metacommunities using linear regression and commonality analysis. We found that niche filtering related to local lake-level environmental conditions was the dominant force structuring macrophytes within metacommunities. However, our results also revealed that elevation range associated with climate (increasing temperature amplitude affecting macrophytes) and spatial location (likely due to dispersal limitation) was important for macrophytes based on the findings of the across-metacommunities analysis. These findings suggest that different determinants influence macrophyte metacommunities within different regions, thus showing context dependency. Moreover, our study emphasized that the use of a single metacommunity scale gives incomplete information on the environmental features explaining variation in macrophyte communities.