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.     

Relationships between lake macrophyte cover and lake and landscape features

We examined the ability of lake and landscape features to predict a variety of macrophyte cover metrics using 54 north temperate lakes. We quantified submersed cover, emergent cover, floating leaf cover, Eurasian watermilfoil cover and total macrophyte cover. Measured lake features included lake physio-chemical and morphometric variables and landscape features included hydrologic, catchment and land use/cover variables. Univariate regression analyses demonstrated that these macrophyte cover metrics are predicted by a wide range of predictor variables, most commonly by: Secchi disk depth, maximum or mean depth, catchment morphometry, road density and the proportion of urban or agricultural land use/cover in the riparian zone or catchment (r² = 0.06–0.46). Using a combination of lake and landscape features in multiple regressions, we were able to explain 29–55% of the variation in macrophyte cover metrics. Total macrophyte cover and submersed cover were related to Secchi disk depth and mean depth, whereas the remaining metrics were best predicted by including at least one land use/cover variable (road density, proportion local catchment agriculture land use/cover, proportion cumulative catchment urban land use/cover, or proportion riparian agriculture land use/cover). The two main conclusions from our research are: (1) that different macrophyte growth forms and species are predicted by a different suite of variables and thus should be examined separately, and (2) that anthropogenic landscape features may override patterns in natural landscape or local features and are important in predicting present-day macrophytes in lakes.     

An assessment of the importance of emergent and floating-leaved macrophytes to trophic status in the Loosdrecht lakes (The Netherlands)

The potential importance of the six major emergent and floating-leaved macrophyte species in recycling of sediment phosphorus in the Loosdrecht lakes was studied. Representative plant samples were collected at the time of maximum biomass, and analysed for biomass and carbon, nitrogen and phosphorus contents. Species cover was determined by aerial photography.Total cover in the seven lakes studied ranged between 2 and 26 percent. For the four main species, biomass per unit area increased with lake trophic status. Consistent differences in C, N and P contents per unit biomass were not observed. Although cover values were small, significant amounts of C, N and P were contained in the macrophytes when compared with maximum sestonic content.Potential P loads from macrophyte decay were calculated. In Lake Loosdrecht, the P load represented 15 percent of current external P inputs. The potential importance of macrophyte decay to P recycling in the other lakes is greater.Decay of macrophyte species at the end of the growing season appears to affect autumnal nutrient and chlorophyll a levels in the water column of some lakes. The re-establishment of submerged species following lake restoration may increase the importance of this pathway in the lakes.     

Rotifer Nervous System Visualized by FMRFamide and 5-HT Immunocytochemistry and Confocal Laser Scanning Microscopy

The characteristic ecology of floodplain lakes is in part due to their relatively strong water-level fluctuations. We analyzed the factors determining water-level fluctuations in 100 floodplain lakes (during non-flooded conditions) in the active floodplains of the Lower Rhine in the Netherlands. Furthermore, we explored the relationship between water-level fluctuations and macrophyte species richness, and analyzed the suitability of artificially created lakes for macrophyte vegetation. During non-flooded conditions along the Rhine, lake water-level fluctuations are largely driven by groundwater connection to the river. Hence, water-level fluctuations are largest in lakes close to the main channel in strongly fluctuating sectors of the river and smallest in isolated lakes. Additionally, water-level fluctuations are usually small in old lakes, mainly due to reduced groundwater hydraulic conductivity resulting from accumulated clay and silt on the bottom. Species richness of floating-leaved and emergent macrophytes was reduced at both small and large water-level fluctuations, whereas species richness of submerged macrophytes was reduced at small water-level fluctuations only. In addition, species richness of submerged macrophytes was higher in lakes that experienced drawdown, whereas no similar pattern was detected for floating-leaved and emergent macrophytes. The decline in amplitude of lake water-level with lake age implies that the number of hydrologically dynamic lakes will decrease over time. Therefore, we suggest that excavation of new lakes is essential to conserve the successional sequence of floodplain water bodies including conditions of high biodiversity. Shallow, moderately isolated, lakes with occasional bottom exposure have the highest potential for creating macrophyte-rich floodplain lakes along large lowland rivers. The water-level regime of such lakes can in part be designed, through choice of the location along the river, the distance away from the river and the depth profile of the lake.     

Geomorphic legacy controls macrophyte distribution within and across disconnected floodplain lakes

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Chironomid stratigraphy in the shallow and eutrophic Lake Søbygaard, Denmark: chironomid–macrophyte co-occurrence

1. A sediment core from the shallow, hypertrophic Lake Søbygaard (mean depth ∼1 m; [TP] 310 μg P L⁻¹) was analysed for subfossil remains to reconstruct chironomid community changes in relation to the succession and disappearance of aquatic macrophytes. 2. Species composition in the 1.10 m core indicates a succession from a 'naturally' eutrophic state to a hypertrophic state during recent centuries. Radiometric dating (²¹⁰Pb) of the uppermost 20 cm of the sediment core (∼1932–93) indicates that sediment accumulation rate had doubled in recent decades. 3. Changes in chironomid assemblages were in close agreement with changes in both diatoms and macrophyte remains in the same core. Distinct changes in chironomid communities reflect the eutrophication process and macrophyte succession through Chara , Ceratophyllum and Potamogeton dominance to the present state, with complete loss of submerged vegetation and dominance by phytoplankton. 4. The co-occurrence and relationship between aquatic macrophyte diversity and recent subfossil chironomid assemblages were assessed from an additional 25 Danish lakes. There was good agreement between the macrophyte and chironomid-based lake groupings. Overall, a significant difference ( P <0.001) was found in chironomid assemblages among lakes in different macrophyte classes. In a pair-wise comparison, the poorly buffered mesotrophic lakes and the alkaline eutrophic lakes had significantly different chironomid assemblages. 5. Chironomid taxa commonly reported to be associated with macrophytes ( Cricotopus , Endochironomus and Glyptotendipes ) were shown also to be indicators of highly productive lakes lacking abundant submerged vegetation.     

Long Transients Near the Ghost of a Stable State in Eutrophic Shallow Lakes with Fluctuating Water Levels

Alternative stable states in shallow lakes have received much attention over the past decades, but less is known about transient dynamics of such lakes in the face of stochastic perturbations such as incidental extremes in water levels driven by climatic variability. Here, we report on the ecosystem dynamics of 70 lakes in the floodplains of the Lower Rhine in The Netherlands from 1999 to 2004. In any particular year, most lakes were either in a macrophyte-dominated clear state or in a contrasting state with turbid water and sparse submerged macrophyte cover. Macrophyte dominance was positively related to the occurrence of drawdown, and negatively to lake surface area and mean depth. We did not find a relation with nutrient levels. Remarkably, shifts between the two contrasting states were common, and episodes of low water levels appear to be an important external driver. A dry period before our study and the exceptionally dry summer of 2003 caused widespread drawdown of floodplain lakes, resulting in establishment of submerged macrophytes in the next year upon refill. In the 4 years without drawdown, many lakes returned to a macrophyte-poor turbid state. Although some lakes turned turbid again quickly, others took several years to shift into the turbid state. A model analysis suggests that such prolonged transient vegetated states may be explained by the fact that the system dynamics slow down in the vicinity of the “almost stable” macrophyte-dominated state. Such a “ghost” of an equilibrium causes the system to stick around that state relatively long before slipping into the only true stable state. Our results support the idea that transient dynamics rather than equilibrium may be the key to understanding the overall state of some ecosystems. A practical implication of our findings is that artificial stabilization of the water level in shallow lakes may have been an important factor aggravating the permanent loss of submerged macrophytes due to cultural eutrophication.     

Seasonality of chlorophyll and nutrients in Lake Erken – effects of weather conditions

The effect of submerged macrophytes on interactions among epilimnetic phosphorus, phytoplankton, and heterotrophic bacterioplankton has been acknowledged, but remains poorly understood. Here, we test the hypotheses that the mean summer phytoplankton biomass (chlorophyll a): phosphorus ratios decrease with increased macrophyte cover in a series of nine lakes. Further, we test that both planktonic respiration and bacterioplankton production increase with respect to phytoplankton biomass along the same gradient of increasing macrophyte cover. Increased macrophyte cover was associated with a lower fraction of particulate phosphorus in epilimnia, with total particulate phosphorus declining from over 80% of total phosphorus in a macrophyte free lake to less than 50% in a macrophyte rich lake. Phytoplankton biomass (chlorophyll a) too was lower in macrophyte dominated lakes, despite relatively high levels of total dissolved phosphorus. Planktonic respiration and bacterioplankton production were higher in macrophyte rich lakes than would be expected from phytoplankton biomass alone, pointing to a subsidy of bacterioplankton metabolism by macrophyte beds at the whole lake scale. The results suggest that the classical view of pelagic interactions, which proposes phosphorus determines phytoplankton abundance, which in turn determines bacterial abundance through the production of organic carbon, becomes less relevant as macrophyte cover increases.     

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 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.     

极端暴雨下山地丘陵区小流域洪水淹没强度对景观特征的响应——以河南“7·20”暴雨为例

通过分析郑州7·20暴雨事件中贾峪河山地丘陵区小流域的洪水过程,探究景观特征对洪水淹没强度影响的时空分布规律,并提出增强流域洪水韧性的规划建议,以缓解河南省山区所面临的社会经济发展、生态环境改善等问题。基于高分6号遥感数据、先进陆地观测卫星(ALOS,Advance Land Observing Satellite)相控阵L频段合成孔径雷达(PALSAR)的地表高程数据和小时降雨量数据,利用MIKE 21水动力模型构建贾峪河流域二维水文模型,分析其2021年7月20日0-24时期上、中、下游的洪水淹没深度和面积,并结合双变量空间自相关模型方法,探究洪水淹没强度与各景观组成和地形因素在时间和空间上的相关性的差异以及其空间聚类类型。研究表明:(1)贾峪河流域淹没面积在0-6时快速增长,于18时达到最大9.59km²,此时各区域淹没面积占比从大到小依次为下游18.88%、上游8.25%、中游12.03%,淹没深度在3m以上的面积占36.11%。(2)地形因素(平均Moran''s I=0.159)对洪水强度的影响大于土地类型(平均Moran''s I=0.096),主要影响因子相对高程、地形湿度指数、矿坑面积百分比、水体面积百分比、建设用地面积百分比、耕地面积百分比以及林地和草地面积百分比与洪水淹没强度之间的相关性随时间变化呈增大趋势,均在暴雨中后期18-24时达到最强。(3)交互探测结果表明,多因子叠加会增强各景观特征对洪水淹没强度的影响。上游影响洪水淹没强度的主要驱动力为矿坑和相对高程,中游和下游的主要影响为水体和地形湿度指数。(4)洪水淹没强度24时的平均值与景观特征指数之间的"高-高"和"高-低"地区的面积占比约0.47%-9.85%,主要分布在上游的中部山区和北部河道周围、中游的河道两侧和下游的河道以及常庄水库周边地区。研究结论建议在上游露天矿坑就地改造为蓄水池并恢复植被,中游和下游应提升河岸带绿地质量,增加下游城区绿色基础设施,减轻城市洪水风险。     

Movement of plankton through lake-stream systems

1. River plankton are often assumed to come from upstream lakes, but the factors controlling the movement of plankton between lakes and rivers into outflow streams are unclear. We tested the possibility that the physical structure of the littoral zone near the lake outlet (depth, presence of macrophytes) and diurnal differences in plankton composition at the lake surface influence the movement of plankton from the lake into the stream and determine their persistence downstream. 2. Zooplankton and phytoplankton biomass, community composition and mean body size were compared between two deep lakes without macrophytes at the lake edge and two shallow lakes with macrophytes at the lake edge. Samples were collected day and night on three dates, in the lake centre, in the littoral zone adjacent to the lake outlet, at the outlet and at two sites downstream in Algonquin Park, Ontario, Canada. 3. The morphology of lake edges clearly affects the movement of lake zooplankton into outlet streams. Outlets draining deeper littoral zones had higher zooplankton biomass than shallow littoral outlets (P < 0.0001), but these differences disappeared within 50 m downstream of the lake. There was no difference in mean zooplankton body size among lake outlets or between littoral and outlet samples. However, shallow littoral zones were dominated by cyclopoid copepods and deeper littoral zones were dominated by Bosmina longirostris. In contrast, phytoplankton biomass entering the outlet was similar to that found within the lake and did not vary with lake outlet morphology. These effects were consistent across several sampling weeks and were not affected by surface zooplankton biomass changes associated with diurnal vertical migration in the lake centre. 4. A comparison with published river zooplankton data suggests that zooplankton are rapidly eliminated from shallow outlet streams (≤1 m deep) but persist in most deeper outlet rivers (≥2 m deep). Because the depth of an outlet river determines downstream zooplankton community development, the contribution of lakes to river plankton communities may be influenced by the location of each lake within the drainage basin. These findings suggest that lake and outflow physical structure influences connection strength between spatially successive habitats.     

Abrupt shift from clear to turbid state in a shallow eutrophic, biomanipulated lake

Monitoring data were used to assess causes behind a recent shift from a clear-water to a turbid-water state in Lake Major, a 10 ha shallow lake in Hungary. In 1999–2000, fish manipulation was conducted in this hypertrophic lake. Reduced fish stock resulted in clearing water and the development of a dense (>80% coverage) submerged vegetation in 2005. During the recent abrupt shift, which occurred in 2007, submerged vegetation subsequently declined after a two-year period of clear water and abundant vegetation. An intense decay of macrophytes within the lake produced a rapid transition between the clear- and turbid-water states. During the clear-water state in 2005–2006, the most important variables predominantly correlating with macrophyte cover were Secchi transparency, temperature and TN, while TN, temperature, Secchi depth and chlorophyll-a were the most significant variables during the turbid-water state in 2007. Nitrogen may play a significant role in the cover of submerged macrophytes when TP is moderate. We argue that several factors in concert are necessary to initiate a shift. Water temperature likely has contributed to triggering shift through inter-year-dependent changes in cover of macrophytes, with fish recruitment having key roles in the dynamics of shallow lakes. Handling editor: Luigi Naselli-Flores     

Establishing realistic restoration targets for nutrient-enriched shallow lakes: linking diatom ecology and palaeoecology at the Attenborough Ponds, U.K.

Eutrophication is a major problem for shallow lakes in the U.K. lowlands. Over the last few decades issues relating to the management and restoration of these lakes have come to the fore, including the need to develop methods for establishing realistic restoration targets. One group of lakes for which restoration is a key concern are the Attenborough Ponds in the English Midlands, an interconnected series of shallow, nutrient-enriched gravel-pit lakes. In November 1972, the highly polluted River Erewash was diverted into the Attenborough Ponds. To determine the ecological effects of this event and to establish restoration goals for this lake system, two complimentary studies were undertaken here; (i) monitoring of the ecology and sedimentary representation of diatom communities in a disturbed lake connected to the R. Erewash (Tween Pond: high nutrient concentrations, no submerged macrophytes) and a relatively undisturbed lake isolated from the R. Erewash (Clifton Pond: lower nutrient concentrations, abundant submerged macrophytes), and; (ii) examination of the sedimentary diatom record in a short sediment core collected from the chosen disturbed lake (Tween Pond). The species composition, seasonality, relative productivity and sedimentary representation of planktonic and periphytic diatom communities were very different in Tween and Clifton Ponds. In Tween Pond two major phases of planktonic diatom production/sedimentation were observed during March–May and July–September. By contrast, planktonic diatoms were restricted to a single spring peak in Clifton Pond and after May there was a switch to the dominance of periphytic diatoms associated with the development of submerged macrophytes. The 1972 diversion of the River Erewash was clearly reflected in the sedimentary diatom record from Tween Pond, by; (i) the abrupt shift to planktonic diatom dominance, and; (ii) increasing percentages of late summer–autumn associated planktonic diatom species. These changes suggest both significant nutrient enrichment and the switch from submerged macrophyte to phytoplankton dominance. Numerical matching of pre-1972 diatom assemblages with surface sediment assemblages in the gravel-pits using Principal Components Analysis (PCA) and a squared chord distance measure revealed no close analogues. Nevertheless, similarities between; (i) percentages of different diatom habitat and seasonality groups, and; (ii) pre-1972 and contemporary macrophyte survey data suggested that Clifton Pond is probably a good pre-diversion analogue for habitat structure (e.g. macrophyte biomass, composition and architecture) and phenology (e.g. diatom and macrophyte seasonality) in the Ponds. The practical value of combining space-for-time substitution and palaeoecological approaches in restoration ecology studies of shallow lakes is discussed.     

The vegetation of Nyumba ya Mungu reservoir, Tanzania

A study was made of the aquatic macrophyte ecology of Nyumba ya Mungu; a nine year old man-made lake in northern Tanzania. The vegetation is described with the aid of percentage cover histograms and standing crop estimates. Concurrent physical, chemical and biological studies helped in ecological interpretations. Distinct vegetation communities were found in the deltaic swamps of the two major inflow rivers and it is concluded that this was due to water chemical differences. The water of the more alkaline River Kikuletwa appeared to influence the greater part of the perimeter swamps as the species composition largely coincided with that reported for higher pH, higher conductivity African lakes. Typha domingensis swamp was predominant and extensive, whilst Cyperus alopecuroides and Paspalidium geminatum were also common emergent species. Floating and submerged plants were relatively unimportant in the lake with respect to cover. The major zonations of species appeared to be a function of water depth, while chemistry seemed the main factor controlling vegetation pattern. In conclusion the significance of interrelationships between the macrophytes and other aquatic organisms is discussed.     

Variable response of functional macrophyte groups to lake characteristics,land use,and space: implications for bioassessment

We investigated how land use at multiple scales affects functional macrophytes groups and ecological status index in the boreal region. We employed a variance partitioning analysis to quantify the relative role of lake characteristics, multiple-scaled land use (catchment, buffer zones of 100, 300 and 500 m), and space in explaining the composition and richness of functional macrophyte groups (emergent and submerged macrophytes and hydrophytes) and ecological status of macrophytes in 110 Finnish lakes. Partial redundancy analysis (community composition) and partial linear regression (richness and status index) revealed that macrophyte community composition, richness, and status index were mostly explained by the pure effect of lake characteristics, which dominated over space for most macrophyte variables. Land use adjacent to shoreline had a higher effect on emergent macrophytes and status index compared to the land use of the whole catchment. Our findings suggest that emergent macrophytes can indicate changes in water quality and hydro-morphology originated from the close vicinity of the littoral zone. Ecological quality assessment based on emergent macrophytes only is probably not sufficient, but including emergent species in the assessments is recommended, especially in the species-poor boreal region.     

Effects of substrate and shading on the growth of two submerged macrophytes

Excessive nutrient loading may cause a shift from submerged macrophyte dominance to free-floating macrophyte dominance. Tolerance and persistence of submerged plants in response to shade may be key characteristics in determining when/if such a shift occurs in shallow eutrophic lakes. This study examines how the cover of floating macrophyte (Lemna minor) and shade of dark mesh affect the growth and photosynthetic efficiency of two submerged plants (Vallisneria natans and Myriophyllum spicatum) on different nutrient substrates. We found that low- and mid-cover intensities generally enhanced the leaf/shoot growth of both submerged plants under all cover and substrate types. The relative growth rates (RGR) were slightly enhanced under the treatment of Lemna with low- and mid-intensity cover on both nutrient-rich substrates. The leaf/shoot growth and RGR of both submerged macrophytes generally increased more under Lemna cover than mesh cover. The photosynthetic efficiency (F _v/F _m value) typically increased with the duration of treatment and the cover densities. In addition, these two macrophytes with contrasting growth forms have markedly different growth and survival strategies in response to covers. These results strengthen the hypothesis that submerged plants can successfully develop under a low-intensity cover of floating vegetation on nutrient-rich substrate.     

The influence of water transparency on the distribution and abundance of macrophytes among lakes of the Mackenzie Delta, Western Canadian Arctic

SUMMARY 1. Macrophyte abundance and distribution was assessed in a chain of six interconnected lakes (all with the same flooding frequency) in the Arctic, where increasing distance from the Mackenzie River channel resulted in a gradient of water transparency (‘chain‐set’ lakes), and in a group of 26 spatially discrete lakes where increasing frequency and duration of lake flooding with river water (controlled by sill height) also resulted in a transparency gradient (‘sill‐set’ lakes). 2. Among the chain‐set lakes, above‐ground macrophyte biomass increased from 0 to 1000 g m^?2 with increasing water transparency. Among the sill‐set lakes, the transparency gradient among the lakes was less well defined and the relations with biomass were more varied. A decrease in flooding was associated with increasing water transparency and an increasing biomass of macrophytes from about 0 to over 2000 g m^?2. For a specific flood frequency, however, the effect of flooding was much greater when lakes were directly connected to a river channel than when floodwaters flowed first through an intervening lake. Among infrequently flooded lakes the effect of flooding on water transparency and biomass was negligible. 3. Among relatively clear lakes in both sets of lakes, biomass increased with increasing water transparency and decreasing lake depth. Among relatively turbid lakes, however, biomass increased with the combined effect of increasing water colour (decreasing water transparency) and increasing lake water depth. The increases in biomass with increasing water colour (coloured dissolved organic matter) and increasing depth, which together result in reduced light at the bed, may be explained by reduced exposure to ultra violet light. 4. An average light attenuation of 1.3 m^?1 (Secchi depth about 1 m) over the growing season appears to represent a threshold water transparency which, in combination with water depths early in the growing season, is consistent with a light supply on the bed required for growth of the common macrophytes in lakes of the Mackenzie Delta. However, a comparison with other systems indicates that macrophytes among lakes of the Mackenzie Delta grow deeper, for a given level of transparency, than is reported in lakes at lower latitude, despite the lower sun angles and increased reflectivity of water surfaces in the arctic. 5. A complete accounting of water transparency (at PAR and UV wavelengths), lake depth, summer sun angle and duration of sunlight may be necessary to explain patterns of macrophyte growth among lakes across a full range of latitudes.     

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.     

Colonization and succession of submerged macrophytes in shallow Lake Væng during the first five years following fish manipulation

Colonization of submerged macrophytes and changes in species composition were studied in shallow Lake Væng during the first five years (1987–91) following fish manipulation in 1986–1988 and a resultant significant improvement in lake water transparency. No submerged macrophytes were present in the lake from 1981–1986, during which time the summer mean Secchi depth ranged from 0.6 and 0.8 m. From 1987 to 1990, Secchi depth increased from 0.9 m to 1.8 m and macrophyte coverage consequently increased (1 % of the lake area in 1987, 2% in 1988, 50% in 1989, 80% in 1990 and 90% in 1991). At the same time, the macrophytes became taller, and the weedbeds more dense. The macrophytes colonized from the exposed and deeper part of the lake towards the sheltered and more shallow part of the lake, a colonization pattern that was confirmed by transplantation experiments. The delay in colonization of the shallow parts may be caused by waterfowl grazing. The vegetation was initially dominated by Potamogeton crispus L., but there was a gradual change during 1988–1989 and Elodea canadensis Michx became exclusively dominant in 1990–1991.