Climate-related differences in the dominance of submerged macrophytes in shallow lakes

It has been suggested that shallow lakes in warm climates have a higher probability of being turbid, rather than macrophyte dominated, compared with lakes in cooler climates, but little field evidence exists to evaluate this hypothesis. We analyzed data from 782 lake years in different climate zones in North America, South America, and Europe. We tested if systematic differences exist in the relationship between the abundance of submerged macrophytes and environmental factors such as lake depth and nutrient levels. In the pooled dataset the proportion of lakes with substantial submerged macrophyte coverage (> 30% of the lake area) decreased in a sigmoidal way with increasing total phosphorus (TP) concentration, falling most steeply between 0.05 and 0.2 mg L⁻¹. Substantial submerged macrophyte coverage was also rare in lakes with total nitrogen (TN) concentrations above 1–2 mg L⁻¹, except for lakes with very low TP concentrations where macrophytes remain abundant until higher TN concentrations. The deviance reduction of logistic regression models predicting macrophyte coverage from nutrients and water depth was generally low, and notably lowest in tropical and subtropical regions (Brazil, Uruguay, and Florida), suggesting that macrophyte coverage was strongly influenced by other factors. The maximum TP concentration allowing substantial submerged macrophyte coverage was clearly higher in cold regions with more frost days. This is in agreement with other studies which found a large influence of ice cover duration on shallow lakes' ecology through partial fish kills that may improve light conditions for submerged macrophytes by cascading effects on periphyton and phytoplankton. Our findings suggest that, in regions where climatic warming is projected to lead to fewer frost days, macrophyte cover will decrease unless the nutrient levels are lowered.     

Invasive Crayfish Threaten the Development of Submerged Macrophytes in Lake Restoration

Submerged macrophytes enhance water transparency and aquatic biodiversity in shallow water ecosystems. Therefore, the return of submerged macrophytes is the target of many lake restoration projects. However, at present, north-western European aquatic ecosystems are increasingly invaded by omnivorous exotic crayfish. We hypothesize that invasive crayfish pose a novel constraint on the regeneration of submerged macrophytes in restored lakes and may jeopardize restoration efforts. We experimentally investigated whether the invasive crayfish (Procambarus clarkii Girard) affects submerged macrophyte development in a Dutch peat lake where these crayfish are expanding rapidly. Seemingly favourable abiotic conditions for macrophyte growth existed in two 0.5 ha lake enclosures, which provided shelter and reduced turbidity, and in one lake enclosure iron was added to reduce internal nutrient loading, but macrophytes did not emerge. We transplanted three submerged macrophyte species in a full factorial exclosure experiment, where we separated the effect of crayfish from large vertebrates using different mesh sizes combined with a caging treatment stocked with crayfish only. The three transplanted macrophytes grew rapidly when protected from grazing in both lake enclosures, demonstrating that abiotic conditions for growth were suitable. Crayfish strongly reduced biomass and survival of all three macrophyte species while waterfowl and fish had no additive effects. Gut contents showed that crayfish were mostly carnivorous, but also consumed macrophytes. We show that P. clarkii strongly inhibit macrophyte development once favourable abiotic conditions for macrophyte growth are restored. Therefore, expansion of invasive crayfish poses a novel threat to the restoration of shallow water bodies in north-western Europe. Prevention of introduction and spread of crayfish is urgent, as management of invasive crayfish populations is very difficult.     

Echosounding observations of coverage,height, PVI,and biomass of submerged macrophytes in the southern basin of Lake Biwa,Japan

We have developed a procedure to process echosounding data to map the distribution of submerged aquatic macrophytes in the southern basin of Lake Biwa, a water body that has a surface area of 52 km² and a mean depth of 4 m. Echosounding observations were made along 27 transect lines spaced at 500-m intervals on August 4 and September 2 and 30, 2003. Quantitative vegetation data including percent coverage, mean vegetation height, and percent vegetation infestation were directly determined using image data from the echosounder recorded digitally on videotape. Based on the image data from an echosounder, a regression model was developed for estimating biomass of submerged macrophytes. The regression model using the total echo strength as the explanatory variable could reliably estimate macrophyte biomass up to 300 g m⁻². Distribution maps of macrophyte height and biomass suggest that the recent summer decline of submerged macrophytes started earlier in shallow areas (<3 m of depth) than deep areas (>4 m) in the southern basin of Lake Biwa.     

Effects of Spirogyra arcta on biomass and structure of submerged macrophyte communities

The presence of algae can greatly reduce the amount of light that reaches submerged macrophytes, but few experimental studies have been conducted to examine the effects of algae on biomass and structure of submerged macrophyte communities. We constructed communities with four submerged macrophytes (Hydrilla verticillata, Egeria densa, Ceratophyllum demersum, and Chara vulgaris) in three environments in which 0 (control), 50 and 100% of the water surface was covered by Spirogyra arcta. Compared to the control treatment, the 100% spirogyra treatment decreased biomass of the submerged macrophyte communities and of all the four macrophytes except C. demersum. Compared to the control and 50% treatments, the 100% treatment significantly increased relative abundance of C. demersum and decreased that of E. densa. Therefore, the presence of S. arcta can greatly affect the productivity and alter the structure of submerged macrophyte communities. To restore submerged macrophyte communities in conditions with abundant algae, assembling communities consisting of C. demersum or similar species may be a good practice.     

Loss of Submerged Macrophytes in Shallow Lakes in North‐Eastern Germany

During the 1950s, the submerged vegetation of shallow lakes in north‐eastern Germany was dominated by nutrient tolerant species, with Ceratophyllum demersum and Myriophyllum sp. being most common. Almost one third of 300 investigated lakes had already lost their submerged macrophytes at that time. Very shallow lakes showed either high or low macrophyte abundance. Increasing depth resulted in medium macrophyte abundances, which may contribute to the stabilisation of local or temporary clearwater states. Forty years later, the percentage of lakes without macrophytes had dramatically increased. Between 55 and 85% of the investigated lakes showed a low abundance. The decline was most pronounced in very shallow lakes. The majority of the investigated lakes showed summer TP concentrations below 100 μg L^–1, but no colonisation by submerged macrophytes, which indicates a resilience against re‐colonisation.     

Influence of submerged vegetation and fish abundance on water clarity in peri-urban eutrophic ponds

Despite the presence of high nutrient concentrations, most ponds located around Brussels (Belgium) show a considerable variation in turbidity. The importance of submerged macrophytes in maintaining the clear-water state requires identification of the main factors determining macrophyte abundance and diversity in ponds and small lakes. In this study, the inter-relationships between submerged macrophyte cover, fish abundance and turbidity were investigated in 13 eutrophic peri-urban ponds. Along a turbidity gradient, vegetation switched from dominance by Stoneworts (Chara and Nitella spp.) in the clearest ponds, to dominance by Potamogeton pectinatus in ponds with a slightly lower water transparency. Despite the presence of both P. pectinatus and Stoneworts in each of the vegetated ponds, only one became dominant. Only a very low abundance (around 20%) of submerged vegetation was found in ponds of intermediate turbidity, while macrophytes were absent in turbid ponds. Multi- and univariate analysis showed a marked difference in chemical, physical and biological properties between ponds deliberately used for fish stocking and ponds that were not. Macrophyte cover was significantly negatively correlated with turbidity and plankti-benthivorous fish abundance. No such correlation was observed with piscivorous fish abundance, except for pike that were associated with a charophyte vegetation in the study ponds. The strong relationship found between fish abundance and turbidity, its negative effect on submerged vegetation cover, and the importance of submerged vegetation in controlling phytoplankton abundance, should be taken into account when selecting ponds for fish stocking. It also suggests that the study ponds have a good potential for ecological quality restoration by biomanipulation.     

Seasonality of macrophytes and interaction with flow in a New Zealand lowland stream

Introduced submerged macrophytes have come to dominate many shallow water bodies in New Zealand, and are a common component of many lowland streams. We investigated the seasonal variation of macrophyte abundance, its influence on flow and channel volume, and the implications of this on stream habitat and functioning in Whakapipi Stream, a typical lowland stream draining a predominantly agricultural catchment.Abundance of macrophytes over the summer was primarily controlled by the phenological cycles of the two dominant species. Mean minimum total macrophyte biomass (36 g m^–2) and cover (7%) occurred in winter (June and August, respectively), and mean maximum biomass (324 g m^–2), and cover (79%) occurred in late summer (March and February respectively). Egeria densa comprised the majority of both cover and biomass during the study period, except early summer (December) when Potamogeton crispus was prevalent in the shallow stream reaches.Macrophyte beds had a major impact on summer stream velocities, reducing average velocities by an estimated 41%. Stream cross-sectional area was maintained at relatively stable levels similar to that recorded over winter, when stream discharge was in the order of seven times greater. The mean velocity distribution coefficient (), and Manning's roughness coefficient (n) were dependent on and displayed a positive linear relationship with macrophyte abundance. The velocity distribution coefficient is recommended as a better indicator of macrophyte effects on velocity in natural streams, as it does not assume uniform velocity, channel depth and slope within the stream reach.Our study shows that submerged macrophytes play an important structuring role within the stream during the summer period, where macrophyte beds act as semi-permeable dams, retarding flow velocities and increasing stream depth and cross-sectional area. This promotes habitat heterogeneity by creating a greater range of flow velocity variation, and also provides large stable low-flow areas. Other likely ecosystem effects resulting from macrophyte/velocity interactions include increased sedimentation, potential for nutrient processing and increased primary production, both by macrophytes and attached epiphyton. The complex architecture of submerged macrophytes and their influence on stream flow may also provide an increased diversity of habitat for other aquatic biota. We propose that management of degraded lowland streams such as the Whakapipi Stream to maintain stretches with moderate quantities of submerged macrophytes interspersed with shaded areas would optimise stream health during low summer flows.     

Detritus Processing and Microbial Dynamics of an Aquatic Macrophyte and Terrestrial Leaf in a Thermally Constant,Spring-Fed Stream

Past studies of organic matter processing in stream ecosystems have focused on the fate of allochthonous terrestrial leaf detritus. In streams with a reduced canopy, submerged macrophytes may provide a significant source of organic matter to the microbial community and higher trophic levels. We compared mass loss patterns and microbial dynamics between a submerged macrophyte, Sagittaria platyphylla, and a deciduous leaf, Populus deltoides. Mass loss rates were higher for the submerged macrophyte, though exponential decay values indicated that both are "fast" decomposers. Bacterial abundance was not significantly different between plant types, but bacterial productivity was significantly higher in Sagittaria. Although fungal biomass was higher overall for Populus, it was not significantly different from that of Sagittaria until day 30. Relative to fungi, bacteria made up 4% and 7% of the peak microbial biomass on Populus and Sagittaria, respectively. Aquatic hyphomycete sporulation was detected only on Populus. These results suggest that in systems where submerged macrophytes are abundant, they can provide a carbon source quantitatively comparable to that of riparian leaf detritus, but that qualitative differences in leaf structural composition cause a shift toward a more significant role for bacterial decomposers.     

Factors affecting the structure of epiphytic gastropod communities in the St. Lawrence River (Quebec,Canada)

A study of epiphytic gastropods associated with two submerged macrophytes (Myriophyllum spicatum and Vallisneria americana) was conducted at a soft and at a hard water site in the St. Lawrence River during two non-consecutive years in order to compare effects of macrophyte species and biomass, site and year in gastropod community structure. There was no effect of macrophyte species on total gastropod abundance nor on gastropod diversity, and few gastropod species showed a marked preference for either macrophyte species. Inter-site and inter-annual variations in diversity, total gastropod abundance and gastropod community structure were greater than variations among macrophytes. However, analysis of variations of epiphytic gastropod communities in one of the two sites shows that abiotic factors are important in explaining epiphytic gastropod distribution. Our results and results of other studies on gastropod population dynamics in the St. Lawrence River indicate that intra- and interspecific competition between gastropods is important on macrophytes and that they must be carefully considered in order to understand epiphytic community structure and population dynamics.     

Submerged Macrophytes Mitigate Direct and Indirect Insecticide Effects in Freshwater Communities

Understanding how ecological interactions mitigate the impacts of perturbations such as pesticides in biological communities is an important basic and applied question for ecologists. In aquatic ecosystems, new evidence from microcosm experiments suggests that submerged macrophytes can buffer cladocerans from pulse exposures to the widely used insecticide malathion, and that mitigation increases with macrophyte density. However, whether these results scale up to more complex aquatic communities where ecological interactions such as competition can alter toxicity is unknown. Further, macrophyte abilities to mitigate different insecticide exposure scenarios (i.e. single versus repeated pulses) have never been tested. To address these gaps, we performed a factorial mesocosm experiment examining the influence of four macrophyte treatments (0, 10, 50, or 100 Elodea Canadensis shoots planted per mesocosm) crossed with three malathion exposure scenarios (no insecticide, single pulse, repeated pulses) on aquatic communities containing zooplankton, phytoplankton, periphyton, two snail species, and larval amphibians. In the absence of macrophytes, single malathion pulses caused short-term declines in cladoceran abundance followed by their rapid recovery, which precluded any indirect effects (i.e. trophic cascades). However, repeated malathion pulses caused cladoceran extinctions, resulting in persistent phytoplankton blooms and reduced abundance of one snail species. In contrast, with macrophytes present, even at low density, malathion had no effect on any taxa. We also discovered novel effects of macrophytes on the benthic food web. In the two highest macrophyte treatments, we observed trends of reduced periphyton biomass, decreased abundance of one snail species, and decreased amphibian time to and mass at metamorphosis. To our knowledge, this is the first evidence of negative submerged macrophyte effects on amphibians, a taxa of global conservation concern. Our findings suggest that facilitating macrophytes could be an important strategy for buffering freshwater communities from insecticides, though consideration of their impacts on animal species is necessary.     

Influence of nutrients, submerged macrophytes and zooplankton grazing on phytoplankton biomass and diversity along a latitudinal gradient in Europe

In order to evaluate latitudinal differences in the relationship of phytoplankton biomass and diversity with environmental conditions in shallow lakes, we sampled 98 shallow lakes from three European regions: Denmark (DK), Belgium/The Netherlands (BNL) and southern Spain (SP). Phytoplankton biomass increased with total phosphorus (TP) concentrations and decreased with submerged macrophyte cover across the three regions. Generic richness was significantly negatively related to submerged macrophyte cover and related environmental variables. Zooplankton:phytoplankton biomass ratios were positively related to submerged macrophyte cover and negatively to phytoplankton generic richness in DK and BNL, suggesting that the low generic richness in lakes with submerged macrophytes was due to a higher zooplankton grazing pressure in these regions. In SP, phytoplankton generic richness was not influenced by zooplankton grazing pressure but related to conductivity. We observed no relationship between phytoplankton generic richness and TP concentration in any of the three regions. The three regions differed significantly with respect to mean local and regional generic richness, with BNL being more diverse than the other two regions. Our observations suggest that phytoplankton diversity in European shallow lakes is influenced by submerged macrophyte cover indirectly by modulating zooplankton grazing. This influence of submerged macrophytes and zooplankton grazing on phytoplankton diversity decreases from north to south.     

Effects of land use on aquatic macrophyte diversity and water quality of ponds

1. Aquatic macrophyte diversity and water quality of 55 ponds in western Japan were related to land use and morphometric variables to identify the environmental factors that sustain biodiversity and the spatial extent at which these factors operate. 2. The relevant spatial extent for floating‐leaved macrophyte richness (500 m from pond edge) was larger than that for submerged macrophyte occurrence (10, 75 and 100 m), whereas emergent macrophyte richness was best explained at much larger extents (1000 m). Total macrophyte richness was explained at the extent of 500, 750 and 1000 m. The extents relevant for explaining the physicochemical condition of pond water (100 and 250 m) were similar to those for submerged and floating‐leaved macrophytes, suggesting that these two growth forms are more sensitive to water quality compared to emergent macrophytes. 3. Diversity of all three growth forms and that of total macrophytes collectively were inversely related to turbidity and nutrient concentration; among the three growth forms, submerged macrophytes were most affected by water quality. 4. Negative relationships were found between the proportion of urban area and the diversity of the three growth forms and that of total macrophytes and water quality. Species richness of emergent, floating‐leaved and total macrophytes decreased with depth and increased with surface area up to about 5000 m², above which it declined. 5. Urbanisation and enlargement of ponds were the two main factors that decreased aquatic macrophyte diversity in irrigation ponds. To alleviate the adverse effects of urban areas on aquatic macrophyte diversity, our results suggest that management efforts should focus on the creation of buffer zones within the relevant spatial extent from the pond edge.     

Effects of common carp (Cyprinus carpio) on macrophytes and invertebrate communities in a shallow lake

1. We performed two field experiments using different cage sizes to determine the direct and indirect effects of carp on macrophytes and invertebrate community composition in a shallow lake. 2. The presence of carp produced a significant decrease in macrophyte abundance, changes in species composition and decreased abundance and diversity of invertebrates in small (2.5 × 2.5 × 2.0 m) cages. In large exclosures (15 × 5 m), only macrophyte stem length was significantly affected by carp. 3. There was considerable variation in the macrophyte and invertebrate responses to carp on different sides of the lake in the large exclosures, suggesting that wind and wave action are also important variables affecting macrophyte growth and persistence. 4. The data suggest that carp can significantly affect species abundance and diversity of macrophytes and some macroinvertebrates.     

Changes in benthic macroinvertebrate abundance and lake isotope (C,N) signals following biomanipulation: an 18-year study in shallow Lake Vaeng,Denmark

Change in the abundance of benthic macroinvertebrates and the stable isotope composition (C, N) of benthic invertebrates and zooplankton in Lake Vaeng, Denmark, was investigated over an 18-year period following biomanipulation (removal of cyprinids). During the first nine years after biomanipulation, the lake was clear and submerged macrophytes were abundant; after this period, a shift occurred to low plant abundance and high turbidity. Two years after the biomanipulation, total density of benthic macroinvertebrates reached a maximum of 17042 (±2335 SE) individuals m⁻² and the density was overall higher when the lake was in a clear state. Redundancy analysis (RDA) suggested macrophyte abundance and total nitrogen (TN) concentration were the dominant structuring forces on the benthic macroinvertebrate assemblage. Stable isotope analysis revealed that δ¹³C of macroinvertebrates and zooplankton was markedly higher in years with high submerged macrophyte abundance than in years without macrophytes, most likely reflecting elevated δ¹³C of phytoplankton and periphyton mediated by a macrophyte-induced lowering of lake water CO₂ concentrations. We conclude that the strong relationship between macrophyte coverage and δ¹³C of macroinvertebrates and cladocerans may be useful in paleoecological studies of past changes in the dynamics of shallow lakes, as change in macrophyte abundance may be tracked by the δ¹³C of invertebrate remains in the sediment.     

Macrophyte diversity and composition in relation to substratum characteristics in regulated and unregulated Danish streams

1. The objective of the present study was to examine how the physical stream environment in regulated and unregulated lowland streams affects the diversity and distribution of macrophyte communities. We analysed the abundance, distribution and composition of macrophytes, together with physical parameters, in seven regulated and seven unregulated unshaded Danish stream reaches. 2. Total macrophyte coverage was similar in the regulated and unregulated streams, but species richness and Shannon diversity were higher in the unregulated streams. Overall, we found fifty-two different species in the regulated stream reaches and sixty-two in the unregulated stream reaches. The spatial distribution of macrophytes on the stream bottom was more heterogeneous in the unregulated streams. 3. We found positive correlations between the coverage and diversity of macrophytes and the coverage of coarse-textured substratum types on the stream bottom, as well as between macrophyte coverage and diversity and substratum heterogeneity. We also found that the macrophytes were more heterogeneously distributed where substratum heterogeneity was greater. 4. The species growing both submerged and emergent were more abundant in the regulated streams, whereas species growing only submerged were more abundant in the unregulated streams. Species growing submerged, species growing both submerged and emergent, and species only growing emergent segregated differently in a canonical correspondence analysis ordination. The submerged species were primarily associated with coarser-textured substrata, whereas species growing both submerged and emergent, and species growing only emergent were associated with finer-textured substrata. 5. The most abundant species in the regulated streams, Sparganium emersum, accounting for almost one-third of the total macrophyte coverage, was primarily associated with clay and sandy bottom substrata, whereas the most abundant species in the unregulated streams, Batrachium peltatum, was primarily associated with gravel and stony substrata.     

沉水植物重建对富营养水体氮磷营养水平的影响

利用富营养浅水湖泊(武汉东湖)中所建立的大型实验围隔系统,研究了沉水植物对水体N、P营养水平的影响．结果表明,沉水植物重建后N、P营养水平显著降低．在研究期间,水生植物围隔总N和总P水平均显著低于对照围隔和大湖水体,而且水生植物围隔的总P含量一般维持在0．1mg·L^-1左右。季节性波动远低于对照围隔和大湖水体．水生植物围隔水体中氨态氮和亚硝态氮含量较低．而硝态氮含量与对照围隔和和大湖水体差别不大．由此可见。恢复以沉水植物为主的水生植被,可以有效地降低N、P营养循环速度,控制浮游植物过度增长,是重建富营养湖泊生态系统的重要措施．     

Wave exposure related growth of epiphyton: implications for the distribution of submerged macrophytes in eutrophic lakes

The distribution of submerged macrophytes in eutrophic lakes has been found to be skewed towards sites with intermediate exposure to waves. Low submerged macrophyte biomass at exposed sites has been explained by, for instance, physical damage from waves. The aim of this study was to investigate if lower biomass at sheltered sites compared to sites with intermediate exposure to waves can be caused by competition from epiphyton.Investigations were performed in eutrophic lakes in southern Sweden. Samples of submerged macrophytes and epiphytic algae on the macrophytes were taken along a wave exposure gradient. The amount of epiphyton (AFDW) per macrophyte biomass decreased with increased exposure. Biomass of submerged macrophytes, on the other hand, increased with increased exposure until a relatively abrupt disappearance of submerged vegetation occurred at high exposures. Production of epiphytic algae was monitored on artificial substrates from June to September at a sheltered and an exposed site in three lakes. It was higher at sheltered sites compared with exposed sites.We suggest that epiphytic algae may be an important factor in limiting the distribution of submerged macrophytes at sheltered sites in eutrophic lakes.     

Interactive effects of light and snail herbivory rather than nutrient loading determine early establishment of submerged macrophytes

Submerged macrophytes play a key role in maintaining a clear‐water phase and promoting biodiversity in shallow aquatic ecosystems. Since their abundance has declined globally due to anthropogenic activities, it is important to include them in aquatic ecosystem restoration programs. Macrophytes establishment in early spring is crucial for the subsequent growth of other warm‐adapted macrophytes. However, factors affecting this early establishment of submerged macrophytes have not been fully explored yet. Here, we conducted an outdoor experiment from winter to early spring using the submerged macrophytes Potamogeton crispus and Vallisneria spinulosa to study the effects of shading, nutrient loading, snail herbivory (Radix swinhoei), and their interactions on the early growth and stoichiometric characteristics of macrophytes. The results show that the effects strongly depend on macrophyte species. Biomass and number of shoots of P. crispus decreased, and internode length increased during low light conditions, but were not affected by nutrient loading. P. crispus shoot biomass and number showed hump‐shaped responses to increased snail biomass under full light. In contrast, the biomass of the plant linearly decreased with snail biomass under low light. This indicates an interaction of light with snail herbivory. Since snails prefer grazing on periphyton over macrophytes, a low density of snails promoted growth of P. crispus by removing periphyton competition, while herbivory on the macrophyte increased during a high density of snails. The growth of V. spinulosa was not affected by any of the factors, probably because of growth limitation by low temperature. Our study demonstrates that the interaction of light with snail herbivory may affect establishment and growth of submerged macrophytes in early spring. Macrophyte restoration projects may thus benefit from lowering water levels to increase light availability and making smart use of cold‐adapted herbivores to reduce light competition with periphyton.     

Effects of an artificial protection structure on the sandy shore macrofaunal community: the special case of Lido di Dante (Northern Adriatic Sea)

Biomanipulation of eutropicated peaty lakes has rarely been successful; clear water with dense macrophyte stands fails to develop in most cases. It was unclear whether (1) high turbidity due to resuspension by benthivorous fish or wind is the major cause of low macrophyte density or whether (2) the establishment of submerged macrophyte stands is prevented by a lack of propagules, low cohesive strength of the lake sediment, high concentrations of phytotoxics, grazing by waterfowl and/or shading by periphyton growth. These hypotheses were tested in an experiment in a shallow peat lake in the Netherlands (Terra Nova). Removal of fish from a 0.5 ha experimental site resulted in clear water and the development of a dense (90% coverage) and species-rich (10 species) submerged vegetation. At a fish-stocked site and a control site the water remained turbid and dense macrophyte stands did not develop. The establishment of submerged macrophytes appeared not to be limited by a lack of propagules. Introduced plants grew poorly in turbid water, but very well in clear water. Exclosures showed that bird grazing reduced the plant biomass. In clear water grazing seemed to enhance the vegetation diversity. Periphyton development did not prevent plant growth in clear water. After the experiment, the fish stock was greatly reduced in the whole lake (85 ha), to test if (3) in a large lake, submerged macrophyte stands will not develop after biomanipulation. In the first season after fish reduction, transparency increased and species-rich submerged macrophyte stands developed, covering 60% of the shallow parts of the lake. Most of the species known to have occurred in the past re-established. The results indicate that high turbidity caused by benthivorous fish in combination with bird grazing were the major causes of the absence of submerged macrophyte stands in this lake. Abiotic conditions after the clearing of the lake were suitable for the growth of macrophytes. We infer that the restoration potential of submerged macrophyte stands in eutrophicated peaty lakes can be high, and results can be obtained quickly.     

Differential response to climatic variation of free‐floating and submerged macrophytes in ditches

1. Experimental studies have indicated in freshwater ecosystems that a shift in dominance from submerged to free‐floating macrophytes may occur with climate change because of increased water surface temperatures and eutrophication. Field evidence is, however, rare. 2. Here, we analysed long‐term (26 years) dynamics of macrophyte cover in Dutch ditches in relation to Dutch weather variables and the North Atlantic Oscillation (NAO) winter index. The latter appears to be a good proxy for Dutch weather conditions. 3. Cover of both free‐floating macrophytes and evergreen overwintering submerged macrophytes was positively related to mild winters (positive NAO winter index). On the other hand, high cover of submerged macrophytes that die back in winter coincided with cold winters (negative NAO winter index). Our results therefore suggest that the effect of weather on macrophyte species depends strongly on their overwintering strategy. 4. The positive relation of free‐floating macrophytes with the NAO winter index was significantly stronger in ditches in organic soil than in those in inorganic soil. This may be because of increased nutrient loading associated with increased decomposition of organic matter and increased run‐off to these ditches during mild wet winters. 5. Our results suggest that mild winters in a changing climate may cause submerged macrophytes with an evergreen overwintering strategy and free‐floating macrophytes to outcompete submerged macrophytes that die back in winter.