Ecological threshold responses in European lakes and their applicability for the Water Framework Directive (WFD) implementation: synthesis of lakes results from the REBECCA project

The objective of this synthesis is to present the key messages and draw the main conclusions from the work on lakes in the REBECCA project, pointing out their links to theoretical ecology and their applicability for the WFD implementation. Type-specific results were obtained from analyses of large pan-European datasets for phytoplankton, macrophytes, macroinvertebrates and fish, and indicators and relationships showing the impact of eutrophication or acidification on these biological elements were constructed. The thresholds identified in many of the response curves are well suited for setting ecological status class boundaries and can be applied in the intercalibration of classification systems. Good indicators for phytoplankton (chrysophytes, cyanobacteria) and macrophytes (isoetids and charaphytes) responses to eutrophication were identified, and the level of eutrophication pressure needed to reach the thresholds for these indicators was quantified. Several existing metrics developed for macrophytes had low comparability and need further harmonisation to be useful for intercalibration of classification systems. For macroinvertebrates, a number of metrics developed for rivers turned out to be less useful to describe lake responses to eutrophication and acidification, whereas other species based indicators were more promising. All the biological elements showed different responses in different lake types according to alkalinity and humic substances, and also partly according to depth. Better harmonisation of monitoring methods is needed to achieve better precision in the dose–response curves. Future research should include impacts of hydromorphological pressures and climate change, as well as predictions of timelags involved in responses to reduction of pressures.     

Morphological alterations of lake shores in Europe: A multimetric ecological assessment approach using benthic macroinvertebrates

Besides pollution, lakes are affected by human alterations of lake-shore morphology. However, ecological effects of such alterations have rarely been studied systematically. Hence, we developed tools to assess the ecological effects of anthropogenic morphological alterations on European lake-shores based on pressure-specific response patterns of littoral macroinvertebrate community composition. Littoral invertebrates were sampled from 51 lakes in seven European countries. Sampling covered a range of natural to heavily morphologically degraded sites including natural shorelines, recreational beaches, ripraps and retaining walls. Biological data were supplemented by standardized morphological data that were collected via a Lake Habitat Survey (LHS) protocol and subsequently used to develop a morphological stressor index. Two biotic multimetric indices were developed based on habitat-specific samples (Littoral Invertebrate Multimetric based on HAbitat samples, LIMHA) and composite samples (Littoral Invertebrate Multimetric based on COmposite samples, LIMCO) through correlations with the morphological stressor index. Similarity analyses showed strong spatial differences in macroinvertebrate community composition between four main geographical regions, i.e. Western, Northern, Central and Southern Europe. The morphological stressor index as well as LIMCO and LIMHA have been developed for each geographical region specifically, thereby optimizing correlations of LIMCO and LIMHA with the respective morphological stressor index. The metric composition of LIMCO and LIMHA and their correlation coefficients with the morphological stressor index are comparable to existing national and regional methods that assess morphological lakeshore degradation via macroinvertebrate communities. Hence, LIMCO and LIMHA indices constitute a new stressor-specific assessment tool that enables comparable lake morphology assessment across Europe, as it has been developed involving a uniform methodology followed by regionalized optimization. These tools fulfil the standards of the EU Water Framework Directive and thus may complement existing assessment approaches used in lake monitoring focusing solely on lake eutrophication so far.     

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.     

Efficient sampling methodologies for lake littoral invertebrates in compliance with the European Water Framework Directive

Lake shores are characterised by a high natural variability, which is increasingly threatened by a multitude of anthropogenic disturbances including morphological alterations to the littoral zone. The European Water Framework Directive (EU WFD) calls for the assessment of lake ecological status by monitoring biological quality elements including benthic macroinvertebrates. To identify cost- and time-efficient sampling strategies for routine lake monitoring, we sampled littoral invertebrates in 32 lakes located in different geographical regions in Europe. We compared the efficiency of two sampling methodologies, defined as habitat-specific and pooled composite sampling protocols. Benthic samples were collected from unmodified and morphologically altered shorelines. Variability within macroinvertebrate communities did not differ significantly between sampling protocols across alteration types, lake types and geographical regions. Community composition showed no significant differences between field composite samples and artificially generated composite samples, and correlation coefficients between macroinvertebrate metrics calculated with both methods and a predefined morphological stressor index were similar. We conclude that proportional composite sampling represents a time- and cost-efficient method for routine lake monitoring as requested under the EU WFD, and may be applied across various European geographical regions.     

Littoral benthic macroinvertebrates of mountain lakes in the Tatra Mountains (Slovakia,Poland)

Littoral benthic macroinvertebrates of 45 mountain lakes in the Tatra Mountains were sampled using a semi-quantitative method in September 2000. A total of 32,852 specimens were identified to 93 taxa belonging to 14 higher taxonomic groups. Multivariate statistics (CCA, RDA) and nine biotic metrics (AQEM/STAR) were used to explain relationships between macroinvertebrate assemblages and environmental variables. Up to 57% of the ecological position of littoral macroinvertebrate assemblages were explained by variance of environmental variables divided into chemical, trophic, physical, catchment and location. Five types of Tatra lakes were recognized using CCA: A — strongly acidified lakes (small catchment, low pH, high concentration of TP, DOC, highest amount of POM in littoral); B — alpine acidified lakes (low amount of POM, low values of biotic metrics); C — alpine non-acidified lakes (high value of diversity index, predominance of Diptera); D — subalpine acidified lakes (high values of biotic metrics: number of families, proportion of crenal and rhithral taxa/total taxa); E — subalpine non-acidified lakes (high values of biotic metrics: number of families, number of genera, BMWP score, number of taxa and abundance of EPT taxa). RDA was used to design five levels of macroinvertebrate taxa acidification tolerance. The Tatra Acidification Index (TAI) was established to assess the acidification status of the lakes in the Tatra Mts.     

The recovery of crustacean zooplankton from acidification depends on lake type

Acidification has harmed freshwater ecosystems in Northern Europe since the early 1900s. Stricter regulations aimed at decreasing acidic emissions have improved surface-water chemistry since the late 1980s but the recovery of biotic communities has not been consistent. Generally, the recovery of flora and fauna has been documented only for a few lakes or regions and large-scale assessments of long-term dynamics of biotic communities due to improved water quality are still lacking. This study investigates a large biomonitoring dataset of pelagic and littoral crustacean zooplankton (Cladocera and Copepoda) from 142 acid-sensitive lakes in Norway spanning 24 years (1997–2020). The aims were to assess the changes in zooplankton communities through time, compare patterns of changes across lake types (defined based on calcium and humic content), and identify correlations between abiotic and biological variables. Our results indicate chemical and biological recovery after acidification, as shown by a general increase in pH, acid neutralizing capacity, changes in community composition and increases in the total number of species, number of acid-sensitive species and functional richness through time. However, the zooplankton responses differ across lake types. This indicates that the concentration of calcium (or alkalinity) and total organic carbon (or humic substances) are important factors for the recovery. Therefore, assessment methods and management tools should be adapted to the diverse lake types. Long-term monitoring of freshwater ecosystems is needed to fully comprehend the recovery dynamics of biotic communities from acidification.     

Long-Term CO2 Variability in Two Shallow Tropical Lakes Experiencing Episodic Eutrophication and Acidification Events

Here we report the long-term (13-year) dynamics of surface pCO₂ and its response to episodic eutrophication and acidification events in two contrasting tropical coastal lakes, one clear-water and the other humic. A short-term nutrient addition experiment was also conducted in mesocosms in the humic lake where in situ eutrophication was moderate. Our objective was to elucidate the response of pCO₂ to interannual changes in key limnological conditions, such as nutrient concentrations and pH. The humic waters showed a median pCO₂ almost ninefold higher across the 13-year study than the clear waters, supporting pCO₂ values about tenfold above atmospheric equilibrium. Eutrophication of the clear-water lake resulted in a decrease in pCO₂ to median values below atmospheric equilibrium, producing a strong sink for atmospheric CO₂. In contrast, pCO₂ increased by over tenfold in both lakes during the acidification phase, resulting in very large CO₂ emissions to the atmosphere. Experimental nutrient additions in the humic lake showed a strong persistence of high pCO₂. The extreme variability in pCO₂ observed here might be a characteristic of tropical lakes and may have important consequences for regional carbon budgets.     

Benthic macroinvertebrate fauna of two alpine lakes over the last century: The value of historical data for interpreting environmental changes

Data on benthic macroinvertebrate assemblages of two alpine lakes in the Tatra Mts (Slovakia) collected in 1914, 1933, 1979–1982, 1993–1997, and 2000 were collated and analysed in an attempt to define their relationship to major environmental events affecting these alpine lakes over the last century. The oldest data were considered an important background before the onset of acidification in one of the lakes in the 1950s, while the most current contain possible information on biological recovery. Results show that data from the 1910s are insufficient to characterize the macroinvertebrate fauna. Deep zone assemblages of both studied lakes have remained stable since the 1930s. Changes in the density of dominant species over time were found in the acidified lake, suggesting a connection between an increase in phosphorus and chlorophyll-a concentration. The composition of the littoral assemblages in the acidified lake in the 1930s indicates that the lake was not strongly acidified at that time. A stable composition since the 1980s reflects the ongoing acid stress. Incomplete species data on the non-acidified lake did not allow us to detect possible changes in the littoral fauna related to acidification. Single findings of species indicating a recovery process need longer-term data to confirm such a trend. Results from this study suggest that historical datasets consist of valuable information that can supplement palaeolimnological analyses in the reconstruction of lake ontogeny.     

Palaeoenvironmental Changes Inferred from Biological Remains in Short Lake Sediment Cores from the Central Alps and Dolomites

In this paper we review research on eutrophication, acidification and climate change based on studies of lake sediments in the Alps. Studies on fossil diatoms, chrysophytes, plant pigments, Cladocera and carbonaceous particles in a number of high altitude lakes in Italy and Switzerland have been used to track environmental changes. We present an original study on fossil Cladocera in sediment cores from 29 lakes that have shown changes in biodiversity from the pre-industrial period (ca. pre AD 1850) to the present. These data show that altitude, ca. 60 years of acidification impacts and fish stocking are the most important factors that affect their distribution and abundance. We review further case studies from two morphometrically and chemically different lakes (Tovel and Paione Superiore). Their lake sediment records span 400 and 150 years, respectively; multi-core, multi-proxy analyses show their pH, trophic and climatic evolution over time.     

Nutrient optima and tolerances of benthic invertebrates,the effects of taxonomic resolution and testing of selected metrics in lakes using an extensive European data base

A large dataset from 1,077 lakes in Finland, Ireland, Sweden and the United Kingdom was collated to analyse the relationship between nutrient status and occurrence of different taxa, as well as between total phosphorus or chlorophyll and commonly used macroinvertebrate metrics developed for river assessment. We found that most taxa were associated with mesotrophic conditions (sensu OECD). Species associated with oligotrophic status included Baetis rhodani, Gammarus lacustris and plecopteran larvae, a group commonly associated with low nutrient status also in rivers. Species tolerant of eutrophic conditions were the chironomid larvae (Chironomus plumosus and Cryptochironomus defectus); and two species of tubificids (Psammoryctides barbatus and Potamothrix hammoniensis). For a number of taxa the associations of benthic invertebrates with nutrient state reported in the literature were not supported by analysis of the REBECCA data. The analysis indicated a variable response of littoral macroinvertebrates to eutrophication pressure when using common metrics developed for macroinvertebrates in rivers. Several metrics showed significantly different responses in lakes with different alkalinity, justifying the use of alkalinity for typing water bodies. These significant responses suggest that benthic invertebrates may be a useful component for classification of ecological status in lakes. The low amount of variance explained by the regressions (<30%), however, suggests that further harmonisation of sampling methods, as well as statistically more robust assessment tools are needed to increase the comparability of datasets and to improve the precision in the dose–response relationships.     

Assessment of eutrophication pressure on lakes using littoral invertebrates

Until the E.U. Water Framework Directive listed benthic invertebrates as a biotic element to be used for ecological classification of lakes, techniques for the assessment of the response of littoral invertebrates to anthropogenic pressures were extremely limited compared with those of rivers and lake profundal zones. We describe here the development of an ecological classification model based on changes of littoral invertebrate assemblages across a gradient of eutrophication, which is the most widespread anthropogenic pressure on lakes across Europe. The model comprises three derived parameters, two of which were developed from taxon-specific optima along a total phosphorus gradient calculated using canonical correspondence analysis, and the third based on invertebrate abundance. Combining the parameter metrics, we can estimate the ecological quality ratio (EQR), relative to those from paleolimnologically-confirmed reference lakes. The model was tested using independent samples collected from both hard and soft substrata and across two seasons from 45 lakes, comprising three alkalinity groups (n = 15 in each), and across gradients in water column total phosphorus concentrations. For hard substrata, EQRs were related consistently and highly significantly to water column concentrations of total phosphorus, accounting for the majority of the variance in every alkalinity group. For samples taken from soft substrata, a significant relationship was found only for high alkalinity lakes, accounting for a moderate proportion of the variability in water column total phosphorus concentrations. Our results compare highly favourably with those from other aquatic ecological assessment methods, irrespective of the faunal or floral group upon which they are based, demonstrating that littoral invertebrate assemblages can provide a statistically robust prediction of nutrient status when samples are collected from hard substrata. While the method was developed specifically to assess nutrient pressures on littoral invertebrates, many lakes are subject to multiple pressures. The development of classification models that incorporate multiple pressures presents a particularly significant challenge for the implementation of the Water Framework Directive, requiring both reliable identification of minimally-impacted reference states and incorporation of pressures that are unlikely to interact in predictable ways.     

Long-term trends in limnological characteristics in the Aurora trout lakes, Sudbury, Canada

Paleolimnological techniques were employed to document the limnological histories of the aurora trout lakes, located in the Sudbury region of Ontario. Two of these lakes are of special interest to fisheries managers, as they represent the only known native habitats of a rare strain of brook trout: the aurora trout. These lakes were limed as part of restoration efforts. Stratigraphic changes in diatom and chrysophyte assemblages from dated lake sediment cores indicate that all the lakes have been impacted by anthropogenic acidification, although the timing and the magnitude of acidification were different amongst the lakes. For example, Whirligig Lake was likely the most naturally acidic lake in the past, but it had further acidified since about 1960. This lake was limed in 1989 and then again in 1993. In Whitepine Lake, acidification started 1940; however, in the most recent sediments (1992), some recovery in lakewater acidity has occurred. In Little Whitepine Lake (a reference lake), acidification started earlier (1920) and the lakewater pH continued to decline until about 1990. This lake was limed in 1989. The chrysophyte paleoindicators suggest a recent recovery in this lake. The successful re-introduction of aurora trout in Whirligig and Whitepine lakes is undoubtedly related to the improved water quality through liming but, based on our paleolimnological indicators, the lakes' limnological characteristics (e.g. pH and metal concentrations) are still different from those present before atmospheric deposition of strong acids from the Sudbury smelters.     

Impacts of invasion by <Emphasis Type="Italic">Dreissena polymorpha</Emphasis> (Pallas, 1771) on the performance of macroinvertebrate assessment tools for eutrophication pressure in lakes

Aquatic ecosystems are experiencing increasing disturbance from multiple stressors caused by anthropogenic activities. The potential for multiple stressors to modify each others’ impacts is not well understood. Legislation such as the EU Water Framework Directive (WFD) requires the development of tools to assess human impacts in aquatic systems that incorporate ecological elements, such as macroinvertebrates. Nutrient enrichment and invasive species are major threats to freshwater systems. The invasive zebra mussel Dreissena polymorpha is a conspicuous invader in freshwater aquatic systems in Europe and North America, and has been linked to drastic changes in macroinvertebrate communities and lake ecology. In 31 lake sites varying in nutrient pressure and in the presence or absence of D. polymorpha we tested three ecological quality assessment tools based on macroinvertebrate assemblages (% Sensitive Taxa to Total Phosphorus (TP), TP Score and Indicator Taxa Metric) and two basic ecological metrics. There were highly significant changes in macroinvertebrate diversity, structure, and composition associated with the invasion by D. polymorpha. While the three metrics performed consistently well in non-invaded systems, they lost explanatory power for eutrophication pressure in invaded systems. Our results suggest that metrics may need to be developed separately for invaded and non-invaded systems, and that the interaction between alien species and nutrient enrichment requires further investigation.     

Rapid and unexpected effects of piscivore introduction on trophic position and diet of perch (Perca flavescens) in lakes recovering from acidification and metal contamination

1. Yellow perch (Perca flavescens) are often the only surviving fish species in acidified lakes. We studied four lakes along a gradient of recovery from acidification and that had different food web complexities. All had abundant yellow perch, two had low piscivore abundance, one had a well‐established piscivore population and one was manipulated by introducing piscivorous smallmouth bass (Micropterus dolomieu). We hypothesised that there would be strong effects on perch abundance, behaviour and diet induced by the presence of piscivores. 2. In the manipulated lake, the bass reduced yellow perch abundance by 75% over a 2‐year period. Concomitantly, perch use of the pelagic habitat fell from 48 to 40%. 3. In contrast to findings from less disturbed systems, yellow perch in the littoral zone of the manipulated lake did not strongly shift from zooplankton to benthic food sources after the arrival of piscivores. Diet analysis using stable carbon isotopes revealed a strong continued reliance on zooplankton in all lakes, independent of the degree of piscivory. The failure to switch to benthos in the refuge area of the littoral zone is most likely related to the depauperate benthos communities in these formerly acidified lakes. 4. Yellow perch in lakes recovering from acidification face a considerable ecological challenge as the necessary switch to benthic diet is hindered by a low abundance of benthos. The arrival of piscivores in these recovering lakes imposes further restrictions on perch access to food items. We infer that future recovery of perch populations (and higher trophic levels) will have to be preceded by the re‐establishment of diverse benthic macroinvertebrate communities in these lakes.     

Quantifying uncertainties in biologically-based water quality assessment: A pan-European analysis of lake phytoplankton community metrics

Lake phytoplankton are adopted world-wide as a sensitive indicator of water quality. European environmental legislation, the EU Water Framework Directive (WFD), formalises this, requiring the use of phytoplankton to assess the ecological status of lakes and coastal waters. Here we provide a rigorous assessment of a number of proposed phytoplankton metrics for assessing the ecological quality of European lakes, specifically in response to nutrient enrichment, or eutrophication, the most widespread pressure affecting lakes. To be useful indicators, metrics must have a small measurement error relative to the eutrophication signal we want them to represent among lakes of different nutrient status. An understanding of variability in metric scores among different locations around a lake, or due to sampling and analytical variability can also identify how best this measurement error is minimised.To quantify metric variability, we analyse data from a multi-scale field campaign of 32 European lakes, resolving the extent to which seven phytoplankton metrics (including chlorophyll a, the most widely used metric of lake quality) vary among lakes, among sampling locations within a lake and through sample replication and processing. We also relate these metrics to environmental variables, including total phosphorus concentration as an indicator of eutrophication.For all seven metrics, 65–96% of the variance in metric scores was among lakes, much higher than variability occurring due to sampling/sample processing. Using multi-model inference, there was strong support for relationships between among-lake variation in three metrics and differences in total phosphorus concentrations. Three of the metrics were also related to mean lake depth. Variability among locations within a lake was minimal (<4%), with sub-samples and analysts accounting for much of the within-lake metric variance. This indicates that a single sampling location is representative and suggests that sub-sample replication and standardisation of analyst procedures should result in increased precision of ecological assessments based upon these metrics.For three phytoplankton metrics being used in the WFD: chlorophyll a concentration, the Phytoplankton Trophic Index (PTI) and cyanobacterial biovolume, >85% of the variance in metric scores was among-lakes and total phosphorus concentration was well supported as a predictor of this variation. Based upon this study, we can recommend that these three proposed metrics can be considered sufficiently robust for the ecological status assessment of European lakes in WFD monitoring schemes.     

Browning affects pelagic productivity in northern lakes by surface water warming and carbon fertilization

Global change impacts important environmental drivers for pelagic gross primary production (GPP) in northern lakes, such as temperature, light, nutrient, and inorganic carbon availability. Separate and/or synergistic impacts of these environmental drivers on pelagic GPP remain largely unresolved. Here, we assess key drivers of pelagic GPP by combining detailed depth profiles of summer pelagic GPP with environmental and climatic data across 45 small and shallow lakes across northern Sweden (20 boreal, 6 subarctic, and 19 arctic lakes). We found that across lakes summer pelagic GPP was strongest associated with lake water temperatures, lake carbon dioxide (CO₂) concentrations impacted by lake water pH, and further moderated by dissolved organic carbon (DOC) concentrations influencing light and nutrient conditions. We further used this dataset to assess the extent of additional DOC-induced warming of epilimnia (here named internal warming), which was especially pronounced in shallow lakes (decreasing 0.96°C for every decreasing m in average lake depth) and increased with higher concentrations of DOC. Additionally, the total pools and relative proportion of dissolved inorganic carbon and DOC, further influenced pelagic GPP with drivers differing slightly among the boreal, subarctic and Arctic biomes. Our study provides novel insights in that global change affects pelagic GPP in northern lakes not only by modifying the organic carbon cycle and light and nutrient conditions, but also through modifications of inorganic carbon supply and temperature. Considering the large-scale impacts and similarities of global warming, browning and recovery from acidification of lakes at higher latitudes throughout the northern hemisphere, these changes are likely to operate on a global scale.     

Responses of insects,especially Chironomidae (Diptera), and mites to 130 years of acidification in a Scottish lake

About 130 years of anthropogenic acidification of Round Loch of Glenhead, SW Scotland, has resulted in successively decreased stability, diversity, productivity and survival rate of the non-biting midge (Diptera: Chironomidae) fauna. Similar trends have also been observed among mayflies, caddis-flies and water mites. The first effects of anthropogenic acidification on the insect and mite fauna, as evidenced by palaeolimnological analyses of²¹⁰Pb-dated sediment cores, occurred as early as around 1850, i.e. earlier than in any other lake hitherto studied. The drop in the lake's pH was first indicated by decreased stability and changes in species composition of chironomids and mayflies in the littoral zone. Major changes in the profundal chironomid fauna did not take place until about 1950, when mean lake pH dropped below 5.0. At the same time, the littoral insect fauna became even more unstable and the first significant elimination of species occurred. Comparison with insect fauna of other lakes suggests that a pH of less than 5 might be critical for the ecological conditions in many acidified lakes. None of the twelve most common chironomid species present prior to the acidification of the lake had disappeared after 120–130 years of considerable acidification, and they are all common in oligotrophic lakes with a pH of 6.5–7.0. This is in contrast to the effects of heavy acidification on other aquatic animal groups. Chironomids are probably more sensitive to lake trophic status than lake acidity.     

The effect of turbidity state and microhabitat on macroinvertebrate assemblages: a pilot study of six shallow lakes

Shallow lakes can occur in two alternative stable states, a clear-water state and a turbid state. This is associated with separate assemblages of fish, zooplankton and plants. Little is known about whether macroinvertebrate assemblages differ across both stable states. This study investigated this in a connected set of three turbid and three clear-water shallow lakes. To overcome confounding effects of differences in spatial structure of macrophytes in turbid and clear-water lakes, we sampled three microhabitats that occurred in both alternative stable states: open water, sago pondweed (Potamogeton pectinatus) and reed (Phragmites australis). Univariate analyses indicated no differences in the number of organisms, taxon richness or diversity between turbid and clear-water lakes. Multivariate analysis, however, showed significant differences in the macroinvertebrate community structure of both stable states. Nine taxa explained a significant amount of the variation between both lake types, of which seven preferred the clear-water lakes. The number of organisms and the taxon richness were higher in reed than in the other microhabitats, but diversity and evenness did not differ among the microhabitats. Multivariate analyses could separate all three microhabitats. Eight taxa, mainly detritus feeders and collector–gatherers, explained most of the variation in the data and preferred the reed microhabitat. The effects of stable state (6.8% explained variance) and microhabitat (13.1% explained variance) on the macroinvertebrate assemblages were largely independent from each other (1.5% shared variance). Although macroinvertebrates are not implemented in the initial theory of stable states, our results show clearly different assemblages across both stable states.     

Phosphorus transformations in the epilimnion of humic lakes: biological uptake of phosphate

SUMMARY. 1. The hypothesis that dissolved humic material (DHM) stimulates bacterial involvement in phosphorus transformations and may thus lead to decreased accessibility of phosphorus to algae was investigated by studying three small forest lakes in southern Finland representing a wide range of concentrations of DHM. 2. Other chemical differences between the three lakes were slight, although the most humic lake exhibited higher concentrations of total phosphorus and of molybdate-reactive phosphorus. Bacterial biomass did not differ significantly between the lakes, but algal biomass was significantly lower at higher DHM concentrations. Consequently the ratio of algal biomass to bacterial biomass was significantly lower in the most humic lake. 3. Uptake of phosphorus from added ³³PO₄ was partitioned between algal and bacterial size fractions by differential filtration. No significant variation between lakes was found in the proportion of particulate ³³P recovered from the algal fraction. 4. Turnover times for phosphate were significantly longer in the most humic lake and also showed lower variability. In general turnover times were long in comparison with values reported from many other lakes. Only briefly in mid summmer did turnover times in two of the lakes shorten to values which would indicate that demand for phosphate was outstripping supply. 5. Short-term storage of samples from the most humic lake stimulated biological incorporation of ³³P, but additions of nitrogen and iron had little effect on phosphate uptake. 6. In these small forest lakes it is probable that no single nutrient consistently limits plankton development. Since no evidence was found that DHM shifts the balance of plankton phosphate uptake away from algae towards bacteria, the influence of DHM on phosphorus transformations may rather be through chemical regulation of free phosphate availability.