The effect of lake morphology on aquatic vegetation development and changes under the influence of eutrophication

Data on aquatic and emergent vegetation, morphology and water quality from 274 Polish lowland lakes surveyed in the years 1996–2009 were used to validate the preliminary typology of Polish lakes based on macrophytes and to indicate the environmental parameters which most significantly determine the vegetation patterns in lakes under various morphological conditions. In highly alkaline lowland lakes representing non-disturbed conditions the key determinants influencing the vegetation patterns were mean depth and the shape of the littoral. Three morphological lake types were distinguished: shallow (<3.5 m), deep, and additionally, within the latter, deep ribbon-shaped, with a clearly elongated base and steep bed slopes. The lake types varied in their vegetation patterns developed under non-disturbed conditions. In the shallow lakes, the share of the phytolittoral in the total lake area (%phytol) was the highest (40–100%, 72.3% on average) and the maximum colonisation depth (C_max) the lowest (3.2 m as the maximum) compared to the lakes from both deep types. In the ribbon-shaped deep lakes, %phytol and the plant coverage (%cover) were the lowest, the proportion of submerged vegetation was extraordinarily high (over 90%) and the emergent vegetation was extremely sparsely developed (<6%) compared to the lakes of the two other types.The alterations of aquatic vegetation resulting from the eutrophication process in distinguished morphological lake types were explored. Within the macrophyte variables tested, three groups of indicators were distinguished: (a) metrics performing best in selected lake types, i.e. the type-specific indicators (abundance metrics, %Pota), (b) metrics performing equally well in all the lake types, considered as the universal indicators (e.g. S_Chara, %Subm and %Emerg) and (c) metrics performing poorly in all the lake types, with generally limited applicability (most of the metrics on syntaxonomic richness). In the shallow lakes, %cover and %phytol performed notably better than in deep lakes, whereas C_max worked best in deep lakes and showed the strongest response in the deep regular-shaped lakes. Moreover, in deep regular-shaped lakes the number of communities of stoneworts and submerged plants (S_Chara and S_Subm), and in deep ribbon-shaped lakes the proportion of area inhabited by vascular plant communities (%Pota) performed exceptionally better than in the other two lake types. The most universal metrics, performing equally well in all the lake types, were the proportions of submerged (%Subm) and emergent (%Emerg) vegetation in the total phytolittoral area.     

Vegetation abundance in lowland flood plan lakes determined by surface area, age and connectivity

SUMMARY 1. We analysed the vegetation structure of 215 lakes in the flood plain of the river Lower Rhine in relation to environmental variables related to hydrological connectivity, lake morphometry, lake age and land use on adjacent land. 2. The frequency distribution of the cover of submerged macrophytes was not normal, implying that submerged macrophytes in any one lake were either scarce or abundant. 3. We observed clear water lakes with submerged macrophyte dominance over a wide range of total P concentration (0.020–0.40 mg total P L^?1). 4. Multiple logistic regression indicated that the probability of dominance by submerged macrophytes decreased markedly with the surface area, depth and age of the lakes. The surface area effect occurred independently of the depth. Further, there was a negative relationship between submerged macrophyte dominance and the long‐term annual duration of inundation by the river. 5. Nymphaeid cover showed a distinct optimum with respect to mean lake depth, being almost absent in lakes shallower than 0.5 m. In contrast to what was found for submerged plants, the probability of occurrence of nymphaeids increased with lake age. 6. The probability of helophyte occurrence increased with lake age, and decreased with the presence of trees, cattle grazing, surface area, use of manure and mean lake depth. 7. In all cases the critical level of one factor (e.g. mean lake depth) depended on other factors (e.g. surface area or age of lake). Thus, in the present study, small lakes tended to remain dominated by submerged macrophytes up to a greater depth than large lakes, and helophytes colonised smaller lakes in an earlier phase. 8. The effect of inundation by the river was modest. This could be because most of our lakes are rarely inundated during the growing season and experience only moderate current velocities while flooded. 9. The results have practical implications for future management of flood plains for conservation purposes. In new water bodies, macrophyte domination will be promoted if many small shallow lakes, rather than few large deep ones, are excavated.     

Contrasting Zooplankton Assemblages in Two Oxbow Lakes with Low Transparencies and Narrow Emergent Macrophyte Belts (Krapina River,Croatia)

The aim of this study was to examine the combined effect of water transparency and narrow macrophyte belts on zooplankton assemblages in two oxbow lakes (Krapina River, Croatia). Samples were collected in open water and among helophytes in the littoral zone from April until September 2008. Rotifers were the most abundant group of zooplankton in both lakes, and dominated in the Krapina oxbow lake 1 (KO1). Lake KO1 had significantly lower transparency, lower percentage macrophyte cover and higher chlorophyll a concentration than Krapina oxbow lake 2 (KO2). In lake KO1, variation in the horizontal distribution of cladocerans and rotifers in terms of their abundance seemed to be determined by competition between Bosmina longirostris and Keratella cochlearis, initiated by oscillation in transparency and detritus availability. In lake KO2, with higher transparency and higher percentage macrophyte cover, the abundance of small‐ and large‐bodied cladocerans increased in the littoral zone simultaneously with higher transparency, suggesting fish predation. Results of this study indicated that small differences in transparencies between the two lakes caused significant differences in horizontal distribution of the zooplankton assemblage. Even narrow helophyte belts offered a refuge to zooplankton, although lower transparencies reduced the effectiveness of macrophytes as a refuge from predators. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Ecological sensitivity of marl lakes to nutrient enrichment: evidence from Hawes Water,UK

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Drainage ditching at the catchment scale affects water quality and macrophyte occurrence in Swedish lakes

1. We have limited knowledge of the effects of land use in general and of drainage ditching in particular on macrophyte communities in lakes. I quantified catchment land use, including drainage ditching, as well as water quality and the number of macrophyte species in 17 Swedish lakes in summer 2005. 2. Land use within 1 km of the studied lakes was analysed in a geographic information system. The following variables were included: areas of forests, mires, agricultural land and urbanization, length of drainage ditches (classified according to the use of the land they drained), and shortest distance from lake to an urban area. To extract and analyse general trends in the data sets, redundancy analysis was used. 3. Water quality was explained mainly by three land‐use related variables: the lengths of agricultural, forest and mire ditches. The length of agricultural ditches was positively correlated with lake water conductivity, total dissolved solids, Ca, N and total organic carbon (TOC). The lengths of forest and mire ditches were positively correlated with lake water characteristics, especially TOC. 4. The number of species representing different macrophyte life forms was explained by three environmental variables: conductivity, and lengths of forest and agricultural ditches. The numbers of isoetids, nymphaeids, elodeids and total obligate hydrophytes were negatively correlated with length of forest ditches. In contrast, the number of lemnids and helophytes was positively correlated with conductivity and length of agricultural ditches. Furthermore, the number of isoetids was exponentially related (negatively) to lengths of agricultural and forest ditches, indicating a threshold response to drainage ditch length. 5. The results suggest that effects on water quality and macrophyte communities result from drainage ditching in the lake catchments rather than from land use itself. Given the total area of drainage‐ditched land worldwide (millions of ha in Scandinavia alone), drainage ditching should be considered when evaluating environmental impacts on lake water quality and macrophyte occurrence and when determining reference conditions for catchment land use.     

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.     

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.     

Disentangling natural and anthropogenic drivers of changes in a shallow lake using palaeolimnology and historical archives

Shallow lakes are susceptible to catastrophic regime shifts characterised by the presence or absence or macrophytes. However, the long-term controls on macrophyte succession in shallow lakes are incompletely understood. To investigate this, we analysed multiple sediment proxies in Lake Rotcze (Eastern Poland), a small, shallow and densely macrophyte-covered lake to (1) reconstruct the ‘reference conditions’ (sensu WFD) and development of the lake in recent centuries, (2) compare historical evidence with the sedimentary record, and (3) identify the natural and anthropogenic drivers of macrophyte succession. Before the twentieth century, conditions in the lake may be referred to as ‘reference conditions’. Subsequently forest clearance in the catchment resulted in lower water transparency, but concurrent catchment drainage lowered water levels and increased macrophyte development. Since 1950 elevated nutrient supply and climatically driven increases in water levels led to the deterioration of water transparency and partial macrophyte withdrawal. At the end of the twentieth century lake-level drawdown led to low phytoplankton biomass and clear water creating a novel ecosystem where macrophytes invade the whole lake. These patterns suggest that both natural and anthropogenically induced water level fluctuations have been critical drivers of macrophyte development.     

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).     

Factors influencing macrophyte metrics in Estonian coastal lakes in the light of ecological status assessment

Macrophytes and factors affecting their distribution were studied in 19 coastal lakes of Estonia. The aim of the study was to determine the factors influencing the distribution of macrophytes in coastal lakes and to assess the suitability of valid macrophyte metrics. Our hypothesis was that in coastal lakes most of the macrophyte distribution patterns are caused by lake-specific variables. Morphological, physico-chemical and catchment area characteristics of the lakes varied greatly. Lakes were in different development stages—lakes nearest to the Baltic Sea were younger and more influenced by brackish water and the furthest lakes were older with more freshwater. All that variability was reflected in macrophyte parameters. Factor analysis of environmental indices divided them into three groups—catchment area, morphometric and water chemistry factors. The first factor may be considered as a pressure and the other two as lake-type-specific factors. Lake catchment area parameters had an influence on Bolboschoenus maritimus, Chara tomentosa and Typha latifolia abundance. Morphometric parameters had an influence on the depth distribution of macrophytes and water chemistry factors on the abundance of helophytes. Current indicator species showed more variability associated with lake-specific factors than with changes in status or the influence of pressures.     

Seasonal dynamics of macrophyte abundance in two regulated streams

This study analysed seasonal dynamics of macrophyte abundance in two perennial lowland regulated streams (Stream 1 and 2) in the Danube basin (Slovakia). Assessments of macrophyte abundance and environmental characteristics were accomplished 7 times during the vegetation period in 2005 within a 100 m long section. Statistically significant differences in total abundance of macrophytes as well as an abundance of macrophyte groups (hydrophytes, amphiphytes, helophytes) and Potamogeton nodosus were detected among months within the vegetation period. Abundance fluctuations for individual macrophyte groups and species were moderate in Stream 1 and much stronger in Stream 2. Only amphiphytes showed bimodal temporal distribution in Stream 1, but the abundance of this group was low, reflecting more or less random occurrence of species in vegetation period. Multiple linear regression revealed that water depths and air temperature are the most significant environmental variables affecting the seasonal pattern of total as well as dominating group abundance in Stream 1 and 2, respectively. In all cases, abundances are significantly influenced by the abundance of the respective group in the preceding month. Culmination time differed between streams for all macrophyte groups except helophytes. Total abundance culminated 0.57 month later in the Stream 1 compared to Stream 2.     

Effects of habitat conditions at hatching time on growth history of offspring European anchovy, <Emphasis Type="Italic">Engraulis encrasicolus</Emphasis>, in the Central Mediterranean Sea

Species distributions are structured by regional and local determinants, which operate at multiple spatial and temporal scales. The purpose of our work was to distinguish the relative roles of local variables, climate, geographical location and post glaciation condition (i.e., delineation between supra- and subaquatic lakes during the post-glacial Ancylus Lake) in explaining variation in macrophyte community composition of all taxa, helophytes and hydrophytes. In addition, we investigated how these four explanatory variable groups affected macrophyte strategy groups based on Grime’s classification. Using partial linear regression and variation partitioning, we found that macrophyte communities are primarily filtered by local determinants together with regional characteristics at the studied spatial scale. We further evidenced that post glaciation condition indirectly influenced on local water quality variables, which in turn directly contributed to the macrophyte communities. We thus suggest that regional determinants interact with local-scale abiotic factors in explaining macrophyte community patterns and examining only regional or local factors is not sufficient for understanding how aquatic macrophyte communities are structured locally and regionally.     

The influence of <Emphasis Type="Italic">Ceratophyllum demersum</Emphasis> L. and <Emphasis Type="Italic">Stratiotes aloides</Emphasis> L. on richness and diversity of aquatic vegetation in the lakes of mid-eastern Poland

The aim of this study was to estimate the influence of Ceratophyllum demersum L. and Stratiotes aloides L. on the species richness and phytocoenotic diversity of aquatic vegetation in lakes. The study was based on field investigations in the Łęczna-Włodawa Lake District located in mid-eastern Poland. The studied lakes (32) included both polymictic and dimictic reservoirs. Both in the polymictic and dimictic lakes, statistically significant correlation coefficients were obtained between the parameters that characterize Stratiotes aloides (the frequency in the phytosociological relevés, the percent share of Stratiotetum aloidis association in the phytolittoral) and those characteristic for species richness (the total number of hydromacrophytes, the number of charophytes, elodeids, nymphaeids and lemnids), phytocoenotic richness (the mean number of species in the phytosociological relevés) and phytocoenotic diversity (the Shannon–Wiener Index). The communities with a share of S. aloides are characterized by higher species richness. However, not a single case of statistically significant dependency was reported between the parameters that describe C. demersum (the frequency in the phytosociological relevés, the percent share Ceratophylletum demersi association in the phytolittoral) and the vegetation traits in both lake groups. Common occurrence of Stratiotes aloides might be considered to be evidence for good status of an aquatic ecosystem and a very good indicator of species richness and diversity of aquatic vegetation.     

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.     

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.     

Submerged vegetation development in two shallow, eutrophic lakes

Submerged macrophyte vegetation in two shallow lakes in the Netherlands, Lake Veluwemeer and Lake Wolderwijd, has been affected by eutrophication in the late 1960's and 1970's. Recent changes in the vegetation occurred in the period following lake restoration measures. Between 1987 and 1993, the dominance of Potamogeton pectinatus decreased, while Charophyte meadows expanded over the same time interval. The pattern of change of the dominant macrophyte species might result from changes in the underwater light climate. Seasonally persistent clear water patches associated with the Chara meadows have been observed in the last few years. The interaction between submerged macrophyte vegetation succession and water transparency in the lakes is discussed.     

Species richness and taxonomic distinctness of lake macrophytes along environmental gradients in two continents

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Measurements of uncertainty in macrophyte metrics used to assess European lake water quality

Uncertainty is an important factor in ecological assessment, and has important implications for the ecological classification and management of lakes. However, our knowledge of the effects of uncertainty in the assessment of different ecological indicators is limited. Here, we used data from a standardized campaign of aquatic plant surveys, in 28 lakes from 10 European countries, to assess variation in macrophyte metrics across a set of nested spatial scales: countries, lakes, sampling stations, replicate transects, and replicate samples at two depth-zones. Metrics investigated in each transect included taxa richness, maximum depth of colonisation and two indicators of trophic status: Ellenberg’s N and a metric based on phosphorus trophic status. Metrics were found to have a slightly stronger relationship to pressures when they were calculated on abundance data compared to presence/absence data. Eutrophication metrics based on helophytes were found not to be useful in assessing the effects of nutrient pressure. These metrics were also found to vary with the depth of sampling, with shallower taxa representing higher trophic status. This study demonstrates the complex spatial variability in macrophyte communities, the effect of this variability on the metrics, and the implications to water managers, especially in relation to survey design.     

Geographic patterns and environmental correlates of taxonomic and phylogenetic diversity of aquatic plants in China

Studies on large-scale geographic patterns of aquatic plant diversity can promote research on the generality of macroecological patterns in different ecosystems. Here, we compiled a checklist of 889 aquatic angiosperms in China, including 738 helophytes (emergent and marshy plants) and 151 hydrophytes (submerged, free-floating, and floating-leaved plants). We explore the geographic patterns and environmental correlates of aquatic plant diversity based on six metrics including species richness (SR), weighted endemism (WE), phylogenetic diversity (PD), phylogenetic endemism (PE), the standardized effect size of phylogenetic diversity (PDses), and the standardized effect size of mean phylogenetic distance (MPDses). Our results show that the diversity of aquatic plants in China is extremely uneven, with high diversity in southeastern China and low diversity in northwestern China, and the geographic patterns of taxonomic and PD are generally consistent. The pattern of helophytes differs from that of hydrophytes. Notably, the wavy-shaped pattern of aquatic plant diversity (especially SR and PD for hydrophytes) across the latitude observed in this study is not consistent with those previously observed for aquatic plants in other continents. Climatic variables and water environmental variables are the main drivers of aquatic plant diversity in China; however, the effects of individual variables differ between helophytes and hydrophytes. Water environmental variables have a greater impact on PDses and MPDses of hydrophytes than those of helophytes. Overall, our work provides insight into understanding the large-scale patterns of aquatic plant diversity and is a critical addition to previous studies on the macroecological pattern of terrestrial organisms.     

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.