Influence of hydrophyte abundance on the spatial distribution of zooplankton in selected lakes in Greece

Submerged hydrophyte vegetation consists of a highly important biotic component of maintaining lake ecosystems towards a “clear water” ecological status. Aquatic macrophytes are well known to play a significant multidimensional role in lakes by competing with phytoplankton growth, stabilising sediment and offering refuge to fishes, macro-invertebrates and littoral zooplankton, amongst others. Zooplanktons that are associated with macrophyte beds, in particular, may act as a positive feedback mechanism that contributes to maintaining a clear-water state. Although there are several studies investigating the relationships between macrophytes and zooplankton in European lakes, few have yet been carried out in Greek lakes. Seasonal field sampling was conducted from spring 2006 to autumn 2008 in four lakes of northwestern Greece. Zooplankton samples were collected from within hydrophyte beds in each lake to estimate their relative abundance and species density. Hydrophyte abundance and composition was recorded on a five-point scale. Moreover, water samples were analysed to determine nutrient and chlorophyll-a concentration. Pearson correlations between zooplankton density and key physicochemical variables were conducted to distinguish significant abiotic variables related with major zooplankton groups. Kruskal–Wallis non-parametric analysis was used to test for significant differences in zooplankton composition and environmental variables amongst the five hydrophyte abundance classes. In addition, Canonical correspondence analysis was used to distinguish possible correlations amongst the macrophyte and zooplankton species. Zooplankton density was significantly higher in dense macrophyte vegetation. Small-sized species (e.g. Rotifera) dominated the zooplankton community, indicating the eutrophic nature of the lakes. Large Cladocera were present in low abundance and were mostly littoral. The current research contributes to a better understanding of relationships between biotic groups in selected Greek lakes.     

Ecological status assessment of European lakes: a comparison of metrics for phytoplankton, macrophytes, benthic invertebrates and fish

Data on phytoplankton, macrophytes, benthic invertebrates and fish from more than 2000 lakes in 22 European countries were used to develop and test metrics for assessing the ecological status of European lakes as required by the Water Framework Directive. The strongest and most sensitive of the 11 metrics responding to eutrophication pressure were phytoplankton chlorophyll a, a taxonomic composition trophic index and a functional traits index, the macrophyte intercalibration taxonomic composition metric and a Nordic lake fish index. Intermediate response was found for a cyanobacterial bloom intensity index (Cyano), the Ellenberg macrophyte index and a multimetric index for benthic invertebrates. The latter also responded to hydromorphological pressure. The metrics provide information on primary and secondary impacts of eutrophication in the pelagic and the littoral zone of lakes. Several of these metrics were used as common metrics in the intercalibration of national assessment systems or have been incorporated directly into the national systems. New biological metrics have been developed to assess hydromorphological pressures, based on aquatic macrophyte responses to water level fluctuations, and on macroinvertebrate responses to morphological modifications of lake shorelines. These metrics thus enable the quantification of biological impacts of hydromorphological pressures in lakes.     

Seasonal fluctuations in macrophyte cover and water transparency of four brown-water lakes: implications for crustacean zooplankton in littoral and pelagic habitats

The dynamics of crustacean zooplankton in the littoral and pelagic zones of four forest lakes having variable water qualities (colour range 130–340 mg Pt l⁻¹, Secchi depth 70–160 cm) were studied. The biomass of zooplankton was higher in the littoral zone than in the pelagic zone only in the lake having the highest transparency. In the three other lakes, biomass was significantly higher in the pelagic zone than in the littoral zone. In the two lakes with highest transparency, the littoral biomass of cladocerans significantly followed the development of macrophyte vegetation, and cladoceran biomass reached the maximum value at the time of highest macrophyte coverage. In lakes with lowest transparency, littoral zooplankton biomass developed independently of macrophyte density and decreased when macrophyte beds were densest. The seasonal development of the littoral copepod biomass did not follow the development of macrophytes in any of the lakes. The mean size of cladocerans in the pelagic zone decreased with increasing Secchi depth of the lake, whereas in the littoral zone no such phenomenon was detected. Seasonally, when water transparency increased temporarily in two of the lakes, the mean size of cladocerans in the pelagic zone decreased steeply. For copepods, no relationship between water transparency and body size was observed. The results suggested that in humic lakes the importance of the littoral zone as a refuge decreases with decreasing transparency of the water and that low water transparency protects cladocerans from fish predation. All the observed between-lake differences could not be explained by fish predation, but were probably attributed to the presence of chaoborid larvae with variable densities. Feeding efficiency of chaoborids is not affected by visibility and thus they can obscure the relationship between water quality, fish density, and the structure of crustacean zooplankton assemblages. Handling editor: S. I. Dodson     

Incorporating invasive weeds into a plant indicator method (LakeSPI) to assess lake ecological condition

The use of lake macrophytes for ecological quality assessments usually seeks to indicate the degree of anthropogenic impact, but few of these schemes implicitly consider impacts of alien weeds. LakeSPI (submerged plant indicators) uses indicators of habitat degradation for macrophytes but also incorporates the degree of impact from alien weeds. Application of LakeSPI to 195 New Zealand lakes provided a dataset to examine how component metrics responded over gradients of anthropogenic pressures, and consider whether weed invasion was merely a ‘passenger’ of habitat degradation, or represented an additional pressure. As expected, metrics measuring depth, and diversity of native vegetation negatively correlated with independent measures of lake eutrophy and were also relatively well explained (69–78% variation) by multiple regression with lake and catchment attributes that included proxies for anthropogenic pressure. In contrast, metrics for invasive impact were largely de-coupled from eutrophication, and poorly explained (31%) by the multiple regression. The response of native vegetation metrics to invasive impact measures varied, with the strongest detected interaction relating to native displacement by increased weed occupation of the vegetated zone. Interactions between invasion and lake trophic status were also discerned, with the extent of weed occupation having a more substantial outcome for the presence of charophyte meadows in low productivity lakes than in more productive lakes. These results suggest weed invasion should be considered as an additional source of anthropogenic pressure, and incorporated in macrophyte bioassessment schemes for a more complete differentiation of lake ecological condition.     

Relations between trophic state indicators and plant biomass in Florida lakes

We collected quantitative data on macrophyte abundance and water quality in 319 mostly shallow, polymictic, Florida lakes to look for relationships between trophic state indicators and the biomasses of plankton algae, periphyton, and macrophytes. The lakes ranged from oligotrophic to hypereutrophic with total algal chlorophylls ranging from 1 to 241 mg m^–3. There were strong positive correlations between planktonic chlorophylls and total phosphorus and total nitrogen, but there were weak inverse relationships between the densities of periphyton and the trophic state indicators total phosphorus, total nitrogen and algal chlorophyll and a positive relationship with Secchi depth. There was no predictable relationship between the abundance of emergent, floating-leaved, and submersed aquatic vegetation and the trophic state indicators. It was only at the highest levels of nutrient concentrations that submersed macrophytes were predictably absent and the lakes were algal dominated. Below these levels, macrophyte abundance could be high or low. The phosphorus–chlorophyll and phosphorus–Secchi depth relationships were not influenced by the amounts of aquatic vegetation present indicating that the role of macrophytes in clearing lakes may be primarily to reduce nutrient concentrations for a given level of loading. Rather than nutrient concentrations controlling macrophyte abundance, it seems that macrophytes acted to modify nutrient concentrations.     

Biological indicators track differential responses of pelagic and littoral areas to nutrient load reductions in German lakes

External nutrient loading was reduced over the past decades as a measure for improving the water quality of eutrophic lakes in western Europe, and has since been accelerated by the adoption of the European Water Framework Directive (WFD) in 2000 (EC, 2000). A variety of eutrophication-related metrics have indicated that the response of biological communities to this decreased nutrient loading has been diverse. Phytoplankton, a major component of the pelagic community, often responded rapidly, whereas a significant delay was observed for submerged macrophytes colonizing littoral areas. In this study we tested whether assessment methods developed for phytoplankton and macrophytes in lakes during Germany's implementation of the WFD reflect this differential response. An assessment of 263 German lakes confirmed that a lower ecological state was recorded when based on the biological quality element (BQE) for macrophytes than the BQE for phytoplankton during the investigated period (2003–2012). On average, lakes had a moderate ecological status for both phytoplankton and macrophyte BQEs, but differences of up to three classes were observed in single cases. Long-term data were available for five lowland lakes subject to strong reductions in phosphorus loading. Their phytoplankton-based assessments indicated a constant improvement of the ecological status in parallel to decreasing water phosphorus concentrations. In contrast, macrophyte-based assessments indicated a 10–20 year delay in their ecological recovery following nutrient load reduction. This delay was confirmed by detailed data on the temporal development of macrophyte species diversity and maximum colonization depths of two lakes after nutrient load reduction. We conclude that the available WFD assessment methods for phytoplankton and macrophyte BQEs are suitable to track the differential response of pelagic and littoral areas to nutrient load reductions in German lakes.     

The use of aquatic vegetation in lake assessment: testing the sensitivity of macrophyte metrics to anthropogenic pressures and water quality

The relationships between aquatic vegetation, anthropogenic pressures and water quality in 83 Polish lowland lakes were analysed in order to select the best responding macrophyte metrics to be used in ecological status assessment. Several metrics describing the syntaxonomic composition, abundance and spatial structure of macrophytes were tested in three morphological lake types (deep regular-shaped, deep ribbon-shaped and shallow) separately by using the Spearman rank correlation coefficients. Among all the parameters tested, only some anthropogenic pressure parameters (the percentage share of urban areas and forests in a catchment, the pollution load from point sources) and all the water quality indicators (TP, TN, Chla, SD) were significantly correlated with most of the macrophyte metrics, although for different lake types the strength and significance of relationships varied substantially. The macrophyte metrics which best responded to anthropogenic pressures and water quality changes were: (i) in deep regular-shaped lakes: the percentage share of Chara phytocenoses and rush phytocenoses in the total phytolittoral area, the maximum depth of plant distribution and the colonisation index; (ii) in deep ribbon-shaped lakes: the hydrophytes/helophytes area ratio, the percentage share of submerged and rush phytocenoses in the total phytolittoral area; (iii) in shallow lakes: the percentage share of Chara phytocenoses and rush phytocenoses in the total phytolittoral area, and the hydrophytes/helophytes area ratio. These metrics can then be used when elaborating new or refining existing macrophyte-based methods of ecological status assessment.     

Similarity between contemporary vegetation and plant remains in the surface sediment in Mediterranean lakes

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European river plant communities: the importance of organic pollution and the usefulness of existing macrophyte metrics

The macrophyte surveys undertaken as part of the EU-funded STAR project are a unique resource allowing aquatic plant communities to be studied at a Pan-European scale (211 stream sites with macrophytes in 14 countries). Using this dataset, we examined the influence of organic pollution in relation to other environmental correlates of river plant community variation across Europe. We examined the relationships between several existing macrophyte metrics and nutrient enrichment, and we also explored the possibility of developing a pan-European macrophyte-based assessment system. We showed that trophic (nutrient) status is an important driver of aquatic plant communities in European rivers. We found that while most existing macrophyte metrics are useful, none can be applied at a pan-European scale in their current form. Our attempt to redesign the Mean Trophic Rank (MTR) index by the addition of further species, and the re-scoring of existing species, resulted in a considerable improvement in the relationship between MTR scores and nutrient variables. We conclude that an enlarged core group of macrophyte species can form part of an improved pan-European macrophyte-based bioassessment system, although regional modifications may be required to adequately describe the nutrient status of certain stream types. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Submerged macrophyte vegetation and the European Water Framework Directive: assessment of status and trends in shallow,alkaline lakes in the Netherlands

The submerged macrophyte vegetation of lakes created after enclosing former estuaries, situated in the central and south-western part of the Netherlands, has been monitored annually from 1992 onwards. Between 1992 and 2004, pronounced changes in overall cover and species composition of the submerged vegetation have occurred, resulting from changes of water quality and morphology in the lakes. In most cases vegetation cover and species diversity increased or remained stable, with the exception of two lakes in the southwest part of the country. Abundance and species composition were assessed according to the requirements of the EU Water Framework Directive, using the assessment procedure proposed to assess macrophytes in natural water bodies in the Netherlands. The assessment procedure included calculation of the ‘ecological quality ratio’ (EQR) for each of eleven water bodies in each of 13 years, based on transect monitoring data. The EQR indicating Good Ecological Status for Macrophytes was achieved in only three of the lakes. The consequences of hydromorphological modifications, and measures necessary to achieve the desired condition are discussed. Nutrient concentrations should be reduced further, while additional management measures are necessary to improve conditions for macrophytes.     

Assessing the condition of lake habitats: a test of methods for surveying aquatic macrophyte communities

The European Union requires member states to monitor the conservation status of water bodies designated under the Habitats and Species Directive. In the UK, macrophytes were identified as useful indicators of conservation status but it was recognised that sampling methods required assessment before nationwide survey work began. At eight lakes, which varied in size (0.9–2797 ha) and trophic status (oligotrophic–eutrophic), we tested a method which uses a combination of three techniques (boat transects, wader transects and perimeter strandline searches) to characterise repeat sectors (six or fewer). The combination of techniques used at a particular lake depended on the accessibility of each sector, the safety of applying the method, expediency and target flora. Species richness and abundance were recorded. By applying indicator metrics/rules to data produced by the method it was possible to categorise the conservation and trophic status of the lakes. Power analysis revealed that the data were statistically robust and could potentially be used to track significant, lake-specific changes in macrophyte flora through time. This analysis also revealed a decreasing return in statistical certainty with increased sampling effort (number of sectors sampled). Little additional certainty was added when more than 4 sectors were sampled per lake. The wader transects proved the most practical means of collecting quantitative data while perimeter surveys were an efficient means of adding useful supplementary semi-quantitative data. Boat surveys, although useful in some situations, particularly in assessing deeper water vegetation, were more time consuming than wader surveys. The potential applications and limitations of these methods are discussed and further tests suggested.     

Spatial patterns in aquatic vegetation composition and environmental covariates along chains of lakes in the Kokemäenjoki watershed (S. Finland)

The diversity and community structure of macrophyte vegetation was studied in 50 boreal lakes forming several upper reaches of lake chains around Lammi, southern Finland. Water chemical parameters and morphometry of the basins were included in a multivariate analysis. Floating-leaved vegetation was the dominant growth form, followed by emergent plants. In downstream lakes, the dominance of floating-leaved macrophytes declined, and emergent species increased in abundance. Species richness was highest in larger lakes, with a wider range of littoral habitats than smaller lakes. Electrical conductivity (range 18–151 mS cm⁻¹, 25 °C) of the water correlated well with patterns in diversity among lakes, but this was not the case for nutrient concentrations. As a whole, morphometrical characteristics of lake basins showed better correlations with vegetation structure than any of the measured chemical parameters. The macrophyte vegetation of neighbouring lake chains differed considerably, depending on the surrounding landscape properties, water quality of the lakes and immigration history of plant species.     

Detritus and nutrient dynamics in the shore zone of lakes: a review

The importance of detritus varies greatly among shore zones of lakes, but in a large majority of these regions detrital pathways prevail. Aside from a great spatial and seasonal variability, macrophytes and bottom sediments appear to be dominant stores of nutrients in these habitats. Macrophytes hold a central position in nutrient cycling in the shore-littoral lake zones. They are the main source of autochthonous detritus as they prevail in the total biomass of littoral organisms, and they are only rarely available as direct food of consumers. Various processes and interactions determine the role of macrophytes in nutrients dynamics. These are: the intensity of nutrient uptake and translocation, release of nutrients by healthy plants and from decomposing plants, exchange of elements between macrophytes and their periphyton, as well as interception of seston by macrophyte stands. Particular plant species differ in their time of dying and susceptibility to decomposition. The changes in decomposing material (size structure of particles and nutrient content) mean that detritus in various stages of decomposition differs in its role in trophic dynamics of shore-littoral lake zones. Several types of shore regions as regards detritus sources and retention level are discussed.     

Bird abundance and species richness on Florida lakes: influence of trophic status,lake morphology,and aquatic macrophytes

Data from 46 Florida lakes were used to examine relationships between bird abundance (numbers and biomass) and species richness, and lake trophic status, lake morphology and aquatic macrophyte abundance. Average annual bird numbers ranged from 7 to 800 birds km^–2 and bird biomass ranged from 1 to 465 kg km^–2. Total species richness ranged from 1 to 30 species per lake. Annual average bird numbers and biomass were positively correlated to lake trophic status as assessed by total phosphorus (r = 0.61), total nitrogen (r = 0.60) and chlorophyll a (r = 0.56) concentrations. Species richness was positively correlated to lake area (r = 0.86) and trophic status (r = 0.64 for total phosphorus concentrations). The percentage of the total annual phosphorus load contributed to 14 Florida lakes by bird populations was low averaging 2.4%. Bird populations using Florida lakes, therefore, do not significantly impact the trophic status of the lakes under natural situations, but lake trophic status is a major factor influencing bird abundance and species richness on lakes. Bird abundance and species richness were not significantly correlated to other lake morphology or aquatic macrophyte parameters after the effects of lake area and trophic status were accounted for using stepwise multiple regression. The lack of significant relations between annual average bird abundance and species richness and macrophyte abundance seems to be related to changes in bird species composition. Bird abundance and species richness remain relatively stable as macrophyte abundance increases, but birds that use open-water habitats (e.g., double-crested cormorant, Phalacrocorax auritus) are replaced by species that use macrophyte communities (e.g., ring-necked duck, Aythya collaris).     

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.     

The influence of flood duration on the surface soil properties and grazing management of karst wetlands (turloughs) in Ireland

This study tested the hypothesis that lake augmentation with well water impacts the distribution and abundance of aquatic plants in lakes. Water chemistry was measured from 14 wells, 14 augmented lakes, and 14 lakes without augmentation. Nine in-lake aquatic macrophyte abundance and species distribution metrics were measured in all lakes. Net photosynthetic rate (NPR) of nine submersed species was also measured in well and lake water. Augmentation increased alkalinity in receiving lakes, but total phosphorus was significantly lower, which resulted in lower chlorophyll and greater Secchi depths. Although measured NPR was higher for all plants incubated in well water, only one (emergent species richness) in-lake aquatic macrophyte metric was different in lakes with and without augmentation. Lake augmentation significantly changed water chemistry of receiving waters, but effects on aquatic macrophytes were minimal, suggesting that other environmental factors are limiting the distribution and abundance of macrophytes in the study lakes. The lower phosphorus levels in augmented lakes were unexpected because phosphorus concentrations in well water were significantly greater than in lakes with or without augmentation. Precipitation of calcium phosphate likely accounts for the reduced phosphorus levels in augmented lakes.     

A fish-based index for the assessment of the ecological quality of temperate lakes

A fish – based index for the assessment of the ecological quality of natural temperate lakes was developed, in accordance to the requirements of the Water Framework Directive (WFD) 2000/60/EC. As a case study, 11 natural lakes located at northern and western Greece were selected. Fish surveys were conducted during mid summer to mid autumn in 2010, 2011 and 2012 using Nordic gillnets and electrofishing. Environmental parameters and anthropogenic pressures were assessed for each lake. Fish species richness, abundance, trophic, reproductive and habitat functional guilds were used for extracting a set of 107 metrics, meeting the requirements of the WFD. All metrics were initially tested as candidates for the index development. A stepwise linear regression of each metric against environmental parameters (lake area, altitude, maximum depth, alkalinity) and anthropogenic pressures (drainage area covered by non-natural land uses – NNLC, water total phosphorus concentrations – TP, Lake Habitat Modification Score – LHMS) was initially conducted for ensuring pressure-response relationships. Reference conditions for each lake were estimated by the hindcasting procedure and the ecological quality for each lake was expressed as the ecological quality ratio (EQR) by a value ranging from 0 (poor quality) to 1 (excellent quality). Two fish fauna metrics, the relative numerical abundance of introduced species (Introduced_a) and the relative biomass of omnivorous species (OMNI_b) were finally extracted as the most significant, responding to LHMS and TP, respectively. The final index was expressed as the mean values of the EQRs of these two metrics. The multimetric fish index presented herein could serve as a tool for assessing the ecological quality of natural lakes at broad geographical scale and generally, in the Mediterranean temperate lakes with similar hydromorphological characteristics.     

西太湖水生植物时空变化

水生植物在浅水湖泊生态系统中具有十分重要的作用。根据中国科学院太湖湖泊生态系统研究站1989年以来的常规监测资料,将西太湖（除东太湖以外的湖区）划分为9个区,采用点截法（point intercept method）,于2002～2005年对各区水生植物的种类、生物量和空间分布情况进行了6次调查。结果表明：西太湖现有水生植物16种,分属于11科12属;水生植物总面积约10220hm^2,其中沉水植物分布面积约占64．58％;挺水植物约占0．29％;漂浮植物约占38．16％。各个种之间生物量差异显著,马来眼子菜、荇菜、芦苇的生物量在所有水生植物中居前3位。多样性分析表明,水生植物种类4a来未发生明显变化,但种类和生物量季节性差异较大。水生植物呈环状分布在距湖岸5km以内的水域和部分岛屿周围,东岸和南岸为水生植物的主要集中分布区域,分布区连续性好,且水草种类齐全。挺水植物种类单一,仅有芦苇（Phragmites communis）一种,分布区域多限于水深小于1．6m的湖岸;沉水植物共有8种,为水生植物的主要组成部分,马来眼子菜（Potamogeton malaianus）的分布频度最高,在西山岛周围水域逐年扩张,成为该区域的先锋种;漂浮植物3种,主要以荇菜（Nymphoides peltata）为主,在七都水域有逐渐扩张的趋势。马来眼子菜、芦苇、荇菜表现出对水环境较强的适应能力,目前为西太湖的3个优势种。20世纪50年代以来,西太湖水生植物种类减少了50种,其中水质下降是导致水生植物种类不断减少甚至消失的一个重要原因。围网养殖和不合理的捕捞方式也对局部水域的植物造成极大的破坏。水生植物生存环境日益严峻,种群单一化趋势日益明显。     

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.