Water level drawdown induces a legacy effect on the seed bank and retains sediment chemistry in a eutrophic clay wetland.

The lack of extreme water level fluctuations in managed, non-peat forming wetland ecosystems can result in decreased productivity through the loss of heterogeneity of these ecosystems. Stochastic disruption, such as a water level drawdown, can effectively reverse this effect and return the wetland to a more productive state, associated with higher biodiversity through new vegetation development. Yet, aside from the effect on vegetation dynamics, little is known about longer-term effects (30 years) of a water level drawdown, hereafter referred to as legacy effects, and how this may impact future water level drawdowns. Here, we aim to unravel the legacy effects of a water level drawdown, stand alone and along a water level gradient, on seed bank properties and nutrient availability in a eutrophic clay wetland. To identify these, we studied the hydrologically managed nature reserve Oostvaardersplassen in the Netherlands. Here, one section was subjected to a multi-year water level drawdown and another section was kept inundated. We determined seed bank properties in both areas, spatially and along a soil elevation gradient (20 cm). Nutrient availability was measured by taking sediment samples along the water level gradient and through experimental manipulation of the water level in an indoor mesocosm experiment. Germination was higher in locations with a water level drawdown history, especially at relatively high elevations. Additionally, the proportion of pioneer species in the seed bank was higher in the water level drawdown area. Overall, nutrient concentrations were higher compared to other aquatic systems. Nutrient availability was higher in the inundated area and did not respond to the water level gradient. We conclude that 30 years after an induced water level drawdown there is no depletion of nutrients, while we still observe a legacy effect in the number of viable seeds in the seed bank.

Multispecies fish tracking across newly created shallow and deep habitats in a forward-restored lake.

BACKGROUND: Freshwater fish communities typically thrive in heterogenous ecosystems that offer various abiotic conditions. However, human impact increasingly leads to loss of this natural heterogeneity and its associated rich fish communities. To reverse this trend, we need guidelines on how to effectively restore or recreate habitats for multiple fish species. Lake Markermeer in the Netherlands is a human-created 70,000-ha lake with a uniform 4 m-water depth, steep shorelines, high wind-induced turbidity, and a declining fish community. In 2016, a forward-looking restoration project newly created a 1000-ha five-island archipelago in this degrading lake, which offered new sheltered shallow waters and deep sand excavations to the fish community. METHODS: In 2020, we assessed how omnivorous and piscivorous fish species used these new habitats by tracking 78 adult fish of five key species across local and lake-scales. We monitored spring arrival of adult fish and assessed local macro-invertebrate and young-of-the-year fish densities. RESULTS: Adult omnivorous Cyprinidae and piscivorous Percidae arrived at the archipelago in early spring, corresponding with expected spawning movements. During the productive summer season, 12 species of young-of-the-year fish appeared along the sheltered shorelines, with particularly high densities of common roach (Rutilus rutilus) and European perch (Perca fluviatilis). This suggests the sheltered, shallow, vegetated waters formed new suitable spawning and recruitment habitat for the fish community. Despite highest food densities for adult fish in the shallowest habitats (< 2-m), adult fish preferred minimally 2-m deep water. After spawning most Cyprinidae left the archipelago and moved long distances through the lake system, while most Percidae remained resident. This may be related to (1) high densities of young-of-the-year fish as food for piscivores, (2) medium food densities for omnivores compared to elsewhere in the lake-system, or (3) the attractiveness of 30-m deep sand excavations that were newly created and frequently used by one-third of all tracked fish. CONCLUSIONS: New littoral zones and a deep sand excavation constructed in a uniform shallow lake that lacked these habitat types attracted omnivorous and piscivorous fish species within four years. Both feeding guilds used the littoral zones for reproduction and nursery, and notably piscivorous fish became residents year-round.

Restoring gradual land-water transitions in a shallow lake improved phytoplankton quantity and quality with cascading effects on zooplankton production.

Land-water transition areas play a significant role in the functioning of aquatic ecosystems. However, anthropogenic pressures are posing severe threats on land-water transition areas, which leads to degradation of the ecological integrity of many lakes worldwide. Enhancing habitat complexity and heterogeneity by restoring land-water transition areas in lake systems is deemed a suitable method to restore lakes bottom-up by stimulating lower trophic levels. Stimulating productivity of lower trophic levels (phytoplankton, zooplankton) generates important food sources for declining higher trophic levels (fish, birds). Here, we study ecosystem restoration project Marker Wadden in Lake Markermeer, The Netherlands. This project involved the construction of a 700-ha archipelago of five islands in a degrading shallow lake, aiming to create additional sheltered land-water transition areas to stimulate food web development from its base by improving phytoplankton quantity and quality. We found that phytoplankton quantity (chlorophyll-a concentration) and quality (inversed carbon:nutrient ratio) in the shallow waters inside the Marker Wadden archipelago were significantly improved, likely due to higher nutrient availabilities, while light availability remained sufficient, compared to the surrounding lake. Higher phytoplankton quantity and quality was positively correlated with zooplankton biomass, which was higher inside the archipelago than in the surrounding lake due to improved trophic transfer efficiency between phytoplankton and zooplankton. We conclude that creating new land-water transition areas can be used to increase light and nutrient availabilities and thereby enhancing primary productivity, which in turn can stimulate higher trophic levels in degrading aquatic ecosystems.

Seagrass ecosystem multifunctionality under the rise of a flagship marine megaherbivore.

Large grazers (megaherbivores) have a profound impact on ecosystem functioning. However, how ecosystem multifunctionality is affected by changes in megaherbivore populations remains poorly understood. Understanding the total impact on ecosystem multifunctionality requires an integrative ecosystem approach, which is especially challenging to obtain in marine systems. We assessed the effects of experimentally simulated grazing intensity scenarios on ecosystem functions and multifunctionality in a tropical Caribbean seagrass ecosystem. As a model, we selected a key marine megaherbivore, the green turtle, whose ecological role is rapidly unfolding in numerous foraging areas where populations are recovering through conservation after centuries of decline, with an increase in recorded overgrazing episodes. To quantify the effects, we employed a novel integrated index of seagrass ecosystem multifunctionality based upon multiple, well-recognized measures of seagrass ecosystem functions that reflect ecosystem services. Experiments revealed that intermediate turtle grazing resulted in the highest rates of nutrient cycling and carbon storage, while sediment stabilization, decomposition rates, epifauna richness, and fish biomass are highest in the absence of turtle grazing. In contrast, intense grazing resulted in disproportionally large effects on ecosystem functions and a collapse of multifunctionality. These results imply that (i) the return of a megaherbivore can exert strong effects on coastal ecosystem functions and multifunctionality, (ii) conservation efforts that are skewed toward megaherbivores, but ignore their key drivers like predators or habitat, will likely result in overgrazing-induced loss of multifunctionality, and (iii) the multifunctionality index shows great potential as a quantitative tool to assess ecosystem performance. Considerable and rapid alterations in megaherbivore abundance (both through extinction and conservation) cause an imbalance in ecosystem functioning and substantially alter or even compromise ecosystem services that help to negate global change effects. An integrative ecosystem approach in environmental management is urgently required to protect and enhance ecosystem multifunctionality.

Evolutionary history of grazing and resources determine herbivore exclusion effects on plant diversity.

Ecological models predict that the effects of mammalian herbivore exclusion on plant diversity depend on resource availability and plant exposure to ungulate grazing over evolutionary time. Using an experiment replicated in 57 grasslands on six continents, with contrasting evolutionary history of grazing, we tested how resources (mean annual precipitation and soil nutrients) determine herbivore exclusion effects on plant diversity, richness and evenness. Here we show that at sites with a long history of ungulate grazing, herbivore exclusion reduced plant diversity by reducing both richness and evenness and the responses of richness and diversity to herbivore exclusion decreased with mean annual precipitation. At sites with a short history of grazing, the effects of herbivore exclusion were not related to precipitation but differed for native and exotic plant richness. Thus, plant species' evolutionary history of grazing continues to shape the response of the world's grasslands to changing mammalian herbivory.

Impacts of sediment resuspension on phytoplankton biomass production and trophic transfer: Implications for shallow lake restoration.

Wind-induced sediment resuspension in shallow lakes may enhance nutrient availability while reducing light availability for phytoplankton growth, thereby affecting the entire food-web. Lake restoration projects that reduce wind-induced resuspension are expected to enhance trophic transfer efficiencies, thereby improving food-web structure and functioning. Yet, reduced resuspension may also lead to lower nutrient concentrations in the water column, promote benthic algae development, reduce phytoplankton biomass production and thereby reduce secondary production by zooplankton. Lake Markermeer is a shallow delta lake in The Netherlands subject to wind-induced sediment resuspension. Restoration project Marker Wadden consists of newly built islands aiming to reduce sediment resuspension and promote higher trophic levels. Here, we tested the effects of reduced sediment resuspension on phytoplankton biomass build-up, benthic algae development, and zooplankton abundances at different temperatures in a 14-day indoor microcosm experiment. We used Marker Wadden sediment with three resuspension intensities combined with three temperatures, to also test effects of higher temperatures in shallow sheltered waters. Reduced sediment resuspension decreased nutrient concentrations and phytoplankton biomass build-up, while increasing light availability and enhancing benthic algae biomass development. Reduced sediment resuspension furthermore increased zooplankton biomass. Enhanced sediment resuspension and higher temperatures synergistically interacted, maintaining a high level of inorganic suspended solids. Our experimental results are in line with long-term seasonal observations from Lake Markermeer. Our findings demonstrate that for shallow lakes suffering from wind effects, measures such as Marker Wadden aimed at reducing sediment resuspension can be effective in restoring secondary production and supporting higher trophic levels.

Species identity and diversity effects on invasion resistance of tropical freshwater plant communities.

Biotic resistance mediated by native plant diversity has long been hypothesized to reduce the success of invading plant species in terrestrial systems in temperate regions. However, still little is known about the mechanisms driving invasion patterns in other biomes or latitudes. We help to fill this gap by investigating how native plant community presence and diversity, and the presence of native phylogenetically closely related species to an invader, would affect invader Hydrilla verticillata establishment success in tropical freshwater submerged plant communities. The presence of a native community suppressed the growth of H. verticillata, but did not prevent its colonisation. Invader growth was negatively affected by native plant productivity, but independent of native species richness and phylogenetic relatedness to the invader. Native plant production was not related to native species richness in our study. We show that resistance in these tropical aquatic submerged plant communities is mainly driven by the presence and biomass of a native community independent of native species diversity. Our study illustrates that resistance provided by these tropical freshwater submerged plant communities to invasive species contrasts to resistance described for other ecosystems. This emphasizes the need to include understudied systems when predicting patterns of species invasiveness and ecosystem invasibility across biomes.

Interactive Effects of Rising Temperature and Nutrient Enrichment on Aquatic Plant Growth, Stoichiometry, and Palatability.

The abundance and stoichiometry of aquatic plants are crucial for nutrient cycling and energy transfer in aquatic ecosystems. However, the interactive effects of multiple global environmental changes, including temperature rise and eutrophication, on aquatic plant stoichiometry and palatability remain largely unknown. Here, we hypothesized that (1) plant growth rates increase faster with rising temperature in nutrient-rich than nutrient-poor sediments; (2) plant carbon (C): nutrient ratios [nitrogen (N) and phosphorus (P)] respond differently to rising temperatures at contrasting nutrient conditions of the sediment; (3) external nutrient loading to the water column limits the growth of plants and decreases plant C:nutrient ratios; and that (4) changes in plant stoichiometry affect plant palatability. We used the common rooted submerged plant Vallisneria spiralis as a model species to test the effects of temperature and nutrient availability in both the sediment and the water column on plant growth and stoichiometry in a full-factorial experiment. The results confirmed that plants grew faster in nutrient-rich than nutrient-poor sediments with rising temperature, whereas external nutrient loading decreased the growth of plants due to competition by algae. The plant C: N and C: P ratios responded differently at different nutrient conditions to rising temperature. Rising temperature increased the metabolic rates of organisms, increased the nutrient availability in the sediment and enhanced plant growth. Plant growth was limited by a shortage of N in the nutrient-poor sediment and in the treatment with external nutrient loading to the water column, as a consequence, the limited plant growth caused an accumulation of P in the plants. Therefore, the effects of temperature on aquatic plant C:nutrient ratios did not only depend on the availability of the specific nutrients in the environment, but also on plant growth, which could result in either increased, unaltered or decreased plant C:nutrient ratios in response to temperature rise. Plant feeding trial assays with the generalist consumer Lymnaea stagnalis (Gastropoda) did not show effects of temperature or nutrient treatments on plant consumption rates. Overall, our results implicate that warming and eutrophication might interactively affect plant abundance and plant stoichiometry, and therefore influence nutrient cycling in aquatic ecosystems.

Nutrient availability controls the impact of mammalian herbivores on soil carbon and nitrogen pools in grasslands.

Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature - herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local-scale herbivory, and its interaction with nutrient enrichment and climate, within global-scale models to better predict land-atmosphere interactions under future climate change.

Success of lake restoration depends on spatial aspects of nutrient loading and hydrology.

Many aquatic ecosystems have deteriorated due to human activities and their restoration is often troublesome. It is proposed here that the restoration success of deteriorated lakes critically depends on hitherto largely neglected spatial heterogeneity in nutrient loading and hydrology. A modelling approach is used to study this hypothesis by considering four lake types with contrasting nutrient loading (point versus diffuse) and hydrology (seepage versus drainage). By comparing the longterm effect of common restoration measures (nutrient load reduction, lake flushing or biomanipulation) in these four lake types, we found that restoration through reduction of nutrient loading is effective in all cases. In contrast, biomanipulation only works in seepage lakes with diffuse nutrient inputs, while lake flushing will even be counterproductive in lakes with nutrient point sources. The main conclusion of the presented analysis is that a priori assessment of spatial heterogeneity caused by nutrient loading and hydrology is essential for successful restoration of lake ecosystems.

High Grazing Pressure of Geese Threatens Conservation and Restoration of Reed Belts.

Reed (Phragmites australis (Cav.) Trin. ex Steud.) beds are important habitat for marsh birds, but are declining throughout Europe. Increasing numbers of the native marsh bird, the Greylag goose (Anser anser L.), are hypothesized to cause reed bed decline and inhibit restoration of reed beds, but data are largely lacking. In this study, we experimentally tested the effect of grazing by Greylag geese on the growth and expansion of reed growing in belts along lake shorelines. After 5 years of protecting reed from grazing with exclosures, reed stems were over 4-fold denser and taller than in the grazed plots. Grazing pressure was intense with 50-100% of the stems being grazed among years in the control plots open to grazing. After 5 years of protection we opened half of the exclosures and the geese immediately grazed almost 100% of the reed stems. Whereas this did not affect the reed stem density, the stem height was strongly reduced and similar to permanently grazed reed. The next year geese were actively chased away by management from mid-March to mid-June, which changed the maximum amount of geese from over 2300 to less than 50. As a result, reed stem density and height increased and the reed belt had recovered over the full 6 m length of the experimental plots. Lastly, we introduced reed plants in an adjacent lake where no reed was growing and geese did visit this area. After two years, the density of the planted reed was six to nine-fold higher and significantly taller in exclosures compared to control plots where geese had access to the reed plants. We conclude that there is a conservation dilemma regarding how to preserve and restore reed belts in the presence of high densities of Greylag geese as conservation of both reed belts and high goose numbers seems infeasible. We suggest that there are three possible solutions for this dilemma: (1) effects of the geese can be mediated by goose population management, (2) the robustness of the reed marshes can be increased, and (3) at the landscape level, spatial planning can be used to configure landscapes with large reed bed reserves surrounded by unmown, unfertilized meadows.

Change in dominance determines herbivore effects on plant biodiversity.

Herbivores alter plant biodiversity (species richness) in many of the world's ecosystems, but the magnitude and the direction of herbivore effects on biodiversity vary widely within and among ecosystems. One current theory predicts that herbivores enhance plant biodiversity at high productivity but have the opposite effect at low productivity. Yet, empirical support for the importance of site productivity as a mediator of these herbivore impacts is equivocal. Here, we synthesize data from 252 large-herbivore exclusion studies, spanning a 20-fold range in site productivity, to test an alternative hypothesis-that herbivore-induced changes in the competitive environment determine the response of plant biodiversity to herbivory irrespective of productivity. Under this hypothesis, when herbivores reduce the abundance (biomass, cover) of dominant species (for example, because the dominant plant is palatable), additional resources become available to support new species, thereby increasing biodiversity. By contrast, if herbivores promote high dominance by increasing the abundance of herbivory-resistant, unpalatable species, then resource availability for other species decreases reducing biodiversity. We show that herbivore-induced change in dominance, independent of site productivity or precipitation (a proxy for productivity), is the best predictor of herbivore effects on biodiversity in grassland and savannah sites. Given that most herbaceous ecosystems are dominated by one or a few species, altering the competitive environment via herbivores or by other means may be an effective strategy for conserving biodiversity in grasslands and savannahs globally.

Aquatic omnivores shift their trophic position towards increased plant consumption as plant stoichiometry becomes more similar to their body stoichiometry.

Human induced eutrophication has strongly altered aquatic ecosystems. With increasing eutrophication, plant nutrient concentrations increase, making them more attractive as food for herbivores. However, most aquatic consumers are omnivorous. Ecological stoichiometry theory predicts that animals prefer to consume food which has a similar nutrient (N and P) composition or C:nutrient ratio compared to their own bodies, hence omnivorous animals may prefer to eat animal prey instead of plants. We asked whether aquatic omnivores would shift their diet towards more plant consumption when plants are more nutritious and their stoichiometry becomes more similar to the stoichiometry of the omnivore. We hypothesized that: (1) the omnivore increases plant consumption as the plant C:nutrient ratio decreases when there is only plant material available; (2) the omnivore generally prefers animal food over plant material; (3) the omnivore will increase its relative plant consumption as the plant C:nutrient ratio decreases, in the presence of animal food. As a model system, we used the pond snail Lymnaea stagnalis (omnivorous consumer), the aquatic plant Potamogeton lucens (plant food to the consumer, cultured at different nutrient regimes to obtain different plant C:nutrient ratios), and the crustacean Gammarus pulex (animal food to the consumer, using freshly dead individuals). When there was only plant material available, the consumers increased their relative consumption rate with decreasing plant C:nutrient ratio from no measurable amount to about 102 mg g-1 day-1. When plant material was offered simultaneously with animal food, even though the omnivores always preferred animal food over plant material, the omnivores still increased their relative intake of plant material as plant C:nutrient ratio decreased, from virtually nothing at the highest to on average 16% of their diet at the lowest plant C:nutrient ratio, with a maximum of 28%. Therefore, we conclude that as nutrient loading increases in aquatic ecosystems, plant-eating omnivorous animals may shift their trophic position towards increased plant consumption and alter the food web structure. As a result, we may observe increased top-down control on aquatic plants.

Warming enhances sedimentation and decomposition of organic carbon in shallow macrophyte-dominated systems with zero net effect on carbon burial.

Temperatures have been rising throughout recent decades and are predicted to rise further in the coming century. Global warming affects carbon cycling in freshwater ecosystems, which both emit and bury substantial amounts of carbon on a global scale. Currently, most studies focus on the effect of warming on overall carbon emissions from freshwater ecosystems, while net effects on carbon budgets may strongly depend on burial in sediments. Here, we tested whether year-round warming increases the production, sedimentation, or decomposition of particulate organic carbon and eventually alters the carbon burial in a typical shallow freshwater system. We performed an indoor experiment in eight mesocosms dominated by the common submerged aquatic plant Myriophyllum spicatum testing two temperature treatments: a temperate seasonal temperature control and a warmed (+4°C) treatment (n = 4). During a full experimental year, the carbon stock in plant biomass, dissolved organic carbon in the water column, sedimented organic matter, and decomposition of plant detritus were measured. Our results showed that year-round warming nearly doubled the final carbon stock in plant biomass from 6.9 ± 1.1 g C in the control treatment to 12.8 ± 0.6 g C (mean ± SE), mainly due to a prolonged growing season in autumn. DOC concentrations did not differ between the treatments, but organic carbon sedimentation increased by 60% from 96 ± 9.6 to 152 ± 16 g C m-2  yaer-1 (mean ± SE) from control to warm treatments. Enhanced decomposition of plant detritus in the warm treatment, however, compensated for the increased sedimentation. As a result, net carbon burial was 40 ± 5.7 g C m-2  year-1 in both temperature treatments when fluxes were combined into a carbon budget model. These results indicate that warming can increase the turnover of organic carbon in shallow macrophyte-dominated systems, while not necessarily affecting net carbon burial on a system scale.

Managing Successional Stage Heterogeneity to Maximize Landscape-Wide Biodiversity of Aquatic Vegetation in Ditch Networks.

The presence of a high diversity of different successional stages in a landscape may help to conserve and promote landscape-wide biodiversity. A strategy to achieve this is using Cyclic Rejuvenation through Management (CRM), an approach employed in a variety of different ecosystems. CRM periodically resets the successional stages in a landscape. For aquatic systems this constitutes vegetation removal and dredging. For this approach to be useful (a) successional stages are required to be different in community composition and (b) these differences need to be caused by true replacement of species between stages. While potentially valid, these assumptions are not generally tested prior to application of CMR. In this study we test these assumptions to explore the usefulness of managing on successional stage heterogeneity for maximizing landscape-wide aquatic plant diversity. We carried out vegetation surveys in the ditch networks of 21 polder landscapes in Netherlands, each containing 24 ditch reaches. Using a clustering approach combined with insight from literature on vegetation succession in these systems we assigned our sampled communities to defined successional stages. After partitioning landscape diversity into its alpha and beta components, we quantified the relative importance of replacement among successional stages. Next, through scenario analyses based on simulations we studied the effects of reducing successional stage heterogeneity on landscape-wide biodiversity. Results showed that differences in community composition among successional stages were a potentially important factor contributing to landscape diversity. Early successional stages were characterized by higher replacement of species compared to late successional stages. In a scenario of gradual decrease of heterogeneity through the systematic loss of the earliest successional stages we found 20% of the species richness in a polder was lost, pointing toward the importance of maintaining early successional stages in a polder. This makes a compelling case for application of CRM within agricultural drainage ditch landscapes to maximize regional aquatic plant diversity. While applied to drainage ditch systems, our data-driven approach is broadly applicable to other systems and may help in providing first indications of the potential of the CRM approach. We argue that CRM may maintain and promote regional biodiversity without compromising the hydrological function of the systems.

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.

Mechanisms of Invasion Resistance of Aquatic Plant Communities.

Invasive plant species are among the major threats to freshwater biodiversity. Few experimental studies have investigated whether native plant diversity can provide biotic resistance to invaders in freshwater ecosystems. At small spatial scales, invasion resistance may increase with plant species richness due to a better use of available resources, leaving less available for a potential invader (Complementarity effect) and/or the greater probability to have a highly competitive (or productive) native species in the community (Selection effect). In submerged aquatic plant communities, we tested the following hypotheses: (1) invader establishment success is greatest in the absence of a native plant community; (2) lower in plant communities with greater native species richness, due to complementary and/or selection effects; and (3) invader establishment success would be lowest in rooted plant communities, based on the limiting similarity theory as the invader is a rooted submerged species. In a greenhouse experiment, we established mesocosms planted with 0 (bare sediment), 1, 2, and 4 submerged plant species native to NW Europe and subjected these to the South African invader Lagarosiphon major (Ridl.) Moss. We used two rooted (Myriophyllum spicatum L., Potamogeton perfoliatus L.) and two non-rooted native species (Ceratophyllum demersum L., Utricularia vulgaris L.) representing two distinct functional groups considering their nutrient acquisition strategy which follows from their growth form, with, respectively, the sediment and water column as their main nutrient source. We found that the presence of native vegetation overall decreased the establishment success of an alien aquatic plant species. The strength of this observed biotic resistance increased with increasing species richness of the native community. Mainly due to a selection effect, the native biomass of mixed communities overyielded, and this further lowered the establishment success of the invader in our experiment. The strongest biotic resistance was caused by the two native plant species that were of the same functional group, i.e., functionally most similar to the invader. These results support the prediction of Elton's biotic resistance hypothesis in aquatic ecosystems and indicate that both species richness and functional group identity can play an important role in decreasing establishment success of alien plant species.

Effects of Rising Temperature on the Growth, Stoichiometry, and Palatability of Aquatic Plants.

Global warming is expected to strengthen herbivore-plant interactions leading to enhanced top-down control of plants. However, latitudinal gradients in plant quality as food for herbivores suggest lower palatability at higher temperatures, but the underlying mechanisms are still unclear. If plant palatability would decline with temperature rise, then this may question the expectation that warming leads to enhanced top-down control. Therefore, experiments that directly test plant palatability and the traits underlying palatability along a temperature gradient are needed. Here we experimentally tested the impact of temperature on aquatic plant growth, plant chemical traits (including stoichiometry) and plant palatability. We cultured three aquatic plant species at three temperatures (15, 20, and 25°C), measured growth parameters, determined chemical traits and performed feeding trial assays using the generalist consumer Lymnaea stagnalis (pond snail). We found that rising temperature significantly increased the growth of all three aquatic plants. Plant nitrogen (N) and phosphorus (P) content significantly decreased, and carbon (C):N and C:P stoichiometry increased as temperature increased, for both Potamogeton lucens and Vallisneria spiralis, but not for Elodea nuttallii. By performing the palatability test, we found that rising temperatures significantly decreased plant palatability in P. lucens, which could be explained by changes in the underlying chemical plant traits. In contrast, the palatability of E. nuttallii and V. spiralis was not affected by temperature. Overall, P. lucens and V. spiralis were always more palatable than E. nuttallii. We conclude that warming generally stimulates aquatic plant growth, whereas the effects on chemical plant traits and plant palatability are species-specific. These results suggest that the outcome of the impact of temperature rise on macrophyte stoichiometry and palatability from single-species studies may not be broadly applicable. In contrast, the plant species tested consistently differed in palatability, regardless of temperature, suggesting that palatability may be more strongly linked to species identity than to intraspecific variation in plant stoichiometry.

Impact of Temperature and Nutrients on Carbon: Nutrient Tissue Stoichiometry of Submerged Aquatic Plants: An Experiment and Meta-Analysis.

Human activity is currently changing our environment rapidly, with predicted temperature increases of 1-5°C over the coming century and increased nitrogen and phosphorus inputs in aquatic ecosystems. In the shallow parts of these ecosystems, submerged aquatic plants enhance water clarity by resource competition with phytoplankton, provide habitat, and serve as a food source for other organisms. The carbon:nutrient stoichiometry of submerged aquatic plants can be affected by changes in both temperature and nutrient availability. We hypothesized that elevated temperature leads to higher carbon:nutrient ratios through enhanced nutrient-use efficiency, while nutrient addition leads to lower carbon:nutrient ratios by the luxurious uptake of nutrients. We addressed these hypotheses with an experimental and a meta-analytical approach. We performed a full-factorial microcosm experiment with the freshwater plant Elodea nuttallii grown at 10, 15, 20, and 25°C on sediment consisting of pond soil/sand mixtures with 100, 50, 25, and 12.5% pond soil. To address the effect of climatic warming and nutrient addition on the carbon:nutrient stoichiometry of submerged freshwater and marine plants we performed a meta-analysis on experimental studies that elevated temperature and/or added nutrients (nitrogen and phosphorus). In the microcosm experiment, C:N ratios of Elodea nuttallii decreased with increasing temperature, and this effect was most pronounced at intermediate nutrient availability. Furthermore, higher nutrient availability led to decreased aboveground C:P ratios. In the meta-analysis, nutrient addition led to a 25, 22, and 16% reduction in aboveground C:N and C:P ratios and belowground C:N ratios, accompanied with increased N content. No consistent effect of elevated temperature on plant stoichiometry could be observed, as very few studies were found on this topic and contrasting results were reported. We conclude that while nutrient addition consistently leads to decreased carbon:nutrient ratios, elevated temperature does not change submerged aquatic plant carbon:nutrient stoichiometry in a consistent manner. This effect is rather dependent on nutrient availability and may be species-specific. As changes in the carbon:nutrient stoichiometry of submerged aquatic plants can impact the transfer of energy to higher trophic levels, these results suggest that eutrophication may enhance plant consumption and decomposition, which could in turn have consequences for carbon sequestration.