The sensitivity of phytoplankton in Loch Leven (U.K.) to changes in nutrient load and water temperature

1. Loch Leven is a shallow, eutrophic lake in Scotland, U.K. It has experienced much change over the 30 years that it has been studied; this has primarily been due to reduced nutrient loads to the lake through active catchment management. Its recovery has been slow and, therefore, we used a phytoplankton community model (PROTECH) to test its sensitivity to changing nutrient loads and water temperature.
2. PROTECH was initialized to simulate the observed phytoplankton community in 1995 and was then repeatedly run through a combination of step-wise changes in water temperature and nutrient load (two treatments were simulated for nutrient load: one changing both nitrate and phosphorus, and one changing just phosphorus). The effect on total chlorophyll- a concentration, cyanobacteria abundance and phytoplankton diversity was examined.
3. Whilst changes in temperature had little effect, variations in the nutrient load produced a range of responses. Increasing only the phosphorus load caused a large increase in Anabaena abundance and total chlorophyll- a concentration. However, the opposite response was recorded when nitrate load was changed as well, with Anabaena increasing its biomass under reduced nutrient load scenarios.
4. The key factor determining the type of response appeared to be nitrogen availability. Anabaena , a nitrogen fixer, could exploit the phosphorus resource of Loch Leven under limiting nitrogen conditions, allowing it to dominate under most of the scenarios tested apart from those supplying extra nitrogen to the lake. The model predictions agree with the observed data, which show that Anabaena continues to dominate the summer phytoplankton bloom in Loch Leven despite the considerable reduction in phosphorus supply from the catchment. This research provides a possible explanation for this.     

Changes in the fauna and flora of a nutrient enriched lake

Summary Changes in the flora and fauna at Loch Leven are described. Species associated with oligotrophic conditions such as charr and desmids have declined or disappeared, and prolonged, dense algal blooms, particularly Cyanophyceae, have been a feature of recent years. Much of the macrophytic vegetation has disappeared and there have been changes in the composition of the zooplankton and benthos, several abundant species disappearing (Table II). Possible reasons for these changes are given.Zusammenfassung Die Veränderungen in der Pflanzen-und Tierwelt von Loch Leven werden beschrieben. Gattungen welche gewöhnlich in oligotrophen Gewässern gefunden werden, wie Salvelinus und Desmidiaceae, sind verringert oder ganz verschwunden. In den letzten Jahren sind Cyanophyceae immer haüfiger hervorgetreten. Viele höhere Pflanzen sind verschwunden und es hat sich vieles geändert in der Komposition des Zooplanktons und Benthos. Mehrere wichtige Gattungen dieser Gruppen verschwinden allmählich. Mögliche Gründe für diese Veränderungen werden gegeben.     

Diet composition and prey preference of tench, Tinca tinca (L.), common bream, Abramis brama (L.), perch, Perca fluviatilis L. and roach, Rutilus rutilus (L.), in two contrasting gravel pit lakes: potential trophic overlap with wildfowl

The abundance of benthic macroinvertebrate taxa and the relative abundance of zooplankton taxa were compared with the diet of bream, tench. perch and roach from two gravel pit lakes during May-July 1986 and 1987. Significant food preferences were demonstrated between species and between lakes. Chironomid pupae dominated the macroinvertebrate diet of perch, roach, ate predominantly either Spirogyra sp. (St Peters Lake) or Daphniu hyalina (Main Lake).
The Main Lake, bream fed largely upon chironomid larvae and the bivalve Sphuerium but in St Peters Lake they positively selected a variety of less abundant benthic invertebrate taxa. Bream switched from benthos to zooplankton in the Main Lake in 1986. Tench ate large numbers of Aselhs and showed positive selection of various macroinvertebrate prey in St Peters but ate D. hjulim in Main Lake. Tropic overlap for chironomids and other macroinvertebrate prey was demonstrated between perch, bream and tench, and potentially with wildfowl which used the gravel pits for breeding and wintering.     

Can lake restoration by fish removal improve the status of profundal macroinvertebrate assemblages?

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Identifying from recent sediment records the effects of nutrients and climate on diatom dynamics in Loch Leven

1. Changes in nutrients and climate have occurred over approximately the same timescales in many European lake catchments. Here, we attempt to interpret the sedimentary diatom record of a large shallow lake, Loch Leven, in relation to these pressures using information gained from analysis of long‐term data sets of water quality, climate and planktonic diatoms. 2. The core data indicate the enrichment of Loch Leven starting in c. 1800–1850, most likely from agricultural practices in the catchment, with a more marked phase since c. 1940–1950 caused by increased phosphorus inputs from sewage treatment works, land drainage and a woollen mill. 3. While the recent diatom plankton remains are dominated by taxa associated with nutrient‐rich conditions, an increase in Aulacoseira subarctica relative to Stephanodiscus taxa since the mid‐1980s suggests that reductions in external catchment sources of nutrients (since 1985) may have resulted in partial recovery. This observation accords well with the long‐term monitoring series of water chemistry and phytoplankton. 4. On a decadal‐centennial scale, the eutrophication signal in the sediment record outweighs any evidence of climate as a control on the diatom community. However, at an inter‐annual scale, while the diatom data exhibit high variability, there are several changes in species composition in the recent fossil record that may be attributed to climatic controls. 5. The study highlights the value of a palaeolimnological approach, particularly when coupled with long‐term data sets, for developing our understanding of environmental change at a range of temporal scales. The diatom record in the sediment can be used effectively to track recovery from eutrophication, but requires greater understanding of contemporary ecology to fully interpret climate impacts. 6. The study illustrates the complexity of ecosystem response to synchronous changes in nutrients and climate, and the difficulty of disentangling the effects of these multiple, interacting pressures.     

Changes in aquatic macrophyte communities in Loch Leven: evidence of recovery from eutrophication?

This article assesses changes in the macrophyte community of Loch Leven over a period of 100 years. Evidence is presented that shows that these changes are associated with eutrophication and with subsequent recovery from eutrophication when anthropogenic nutrient inputs to the loch were reduced. This study uses macrophyte survey data from 1905, 1966, 1972, 1975, 1986, 1993, 1999 and 2008. In each of the four most modern surveys, the loch was divided into 19 sectors, each with at least one transect ranging from the shallowest to the deepest occurrence of macrophytes. From these data, a range of indicators of recovery were derived at the whole lake scale: the relative abundance of taxa, taxon richness and evenness. All of these metrics showed an improvement since 1972. Species richness, measured at the scales of survey sector and individual samples, also appeared to have increased in recent years. All of these measures, coupled with ordination of the presence/absence composition data from all survey years, indicate that the macrophyte community in the loch is recovering towards the state that was recorded in 1905.     

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.     

Combining limnological and palaeolimnological data to disentangle the effects of nutrient pollution and climate change on lake ecosystems: problems and potential

1. As long‐term observational lake records continue to lengthen, the historical overlap with lake sediment records grows, providing increasing opportunities for placing the contemporary ecological status of lakes in a temporal perspective. 2. Comparisons between long‐term data sets and sediment records, however, require lake sediments to be accurately dated and for sediment accumulation rates to be sufficiently rapid to allow precise matching with observational data. 3. The critical role of the sediment record in this context is its value in tracking the changing impact of human activity on a lake from a pre‐disturbance reference through to the present day. 4. Here, we use data from a range of lakes across Europe presented as case studies in this Special Section. The seven sites considered all possess both long‐term observational records and high‐quality sediment records. Our objective is to assess whether recent climate change is having an impact on their trophic status and in particular whether that impact can be disentangled from the changes associated with nutrient pollution. 5. The palaeo‐data show clear evidence for the beginning of nutrient pollution varying from the mid‐nineteenth century at Loch Leven to the early and middle twentieth century at other sites. The monitoring data show different degrees of recovery when judged against the palaeo‐reference. 6. The reason for limited recovery is attributed to continuing high nutrient concentrations related to an increase in diffuse nutrient loading or to internal P recycling, but there is some evidence that climate change may be playing a role in offsetting recovery at some sites. If this is the case, then lake ecosystems suffering from eutrophication may not necessarily return to their pre‐eutrophication reference status despite the measures that have been taken to reduce external nutrient loading. 7. The extent to which future warming might further limit such recovery can be evaluated only by continued monitoring combined with the use of palaeo‐records that set the pre‐eutrophication reference.     

Ecological impact of introduced fish in high altitude lakes: a case of study from the European Alps

Fish introduction is a major threat to alpine lake biota leading to the loss of native species and to the degeneration of natural food-webs. This study provides an extensive investigation on the impact of the introduced fish Salvelinus fontinalis on the native communities of alpine lakes in the Gran Paradiso National Park. We compared the macroinvertebrate and zooplankton communities of six stocked and nine fishless lakes with a repeated sampling approach during the summers 2006–2009. The impact of fish presence on alpine lake fauna is often mediated by the strong seasonality governing these ecosystems, and it dramatically affects the faunal assemblage of littoral macroinvertebrates and the size, structure, and composition of the pelagic zooplankton community with a strong selective predation of the more visible taxa. Direct ecological impacts include a decrease or extinction of non-burrower macroinvertebrates and of large zooplankton species, while small zooplankton species and burrower macroinvertebrates were indirectly advantaged by fish presence. Due to the existence of a compensation between rotifers and crustaceans, fish presence does not affect total zooplankton biomass and diversity even if fish are a factor of ecological exclusion for large crustaceans. These compensatory mechanisms are a key process surrounding the impact of introduced fish in alpine lakes.     

The recent diatom history of Loch Leven, Kinross

Diatoms in the top metre of lake sediment have been studied for indications or recent changes in the environment of Loch Leven. The diatoms indicate a eutrophic, alkaline lake with Melosira italica ssp. subarctica, Tabellaria Flocculosa and Stephanodiscus astraea the dominant taxa at the base of the 1-m section. There is then an appeance of Melosira ambigua and Stephanodiscus hantzschii suggesting a richer environment. Post A.D. 1816 changes in taxa suggest possible correlation with the lowering of lake level in 1830 and the dominance of Diatoma elongatum, Cyclotella pseudostelligera, Stephanodiscus astraea var. minutula, S. tenuis, S. hantzschii, and Synedra rumpens and var. soctica in zone 4 agrees with present-day plankton records.     

Long-term recovery of macroinvertebrate biota in grossly polluted streams: Re-colonisation as a constraint to ecological quality

To meet targets imposed by the European Water Framework Directive (2000/60/EC) it is vital that measures to improve the status of rivers are both effective and economically viable. Achievement of such aims needs robust understanding of biological responses to changes in water quality vis-à-vis mechanisms of and constraints to the colonization of previously polluted sites. This study therefore examined the long-term chemical and biological changes in historically polluted rivers to elucidate the responses of macroinvertebrate biota to improvements in chemical water quality. For three historically polluted sites in the English Midlands, data from surveys over a period of ca. 50 years were analysed. Ammonia (NH₃) and 5-day biochemical oxygen demand (BOD₅) were used as chemical water quality indicators. Variations in the ecological recovery of the study sites were assessed using an average pollution sensitivity score (Average Score Per Taxon) and the number of taxa present (usually to family level) present in hand-net samples. Ecological recovery varied widely and was influenced by the intensity and spatial extent of the pollution and the proximity of available sources of potential colonisers. At the site most isolated from potential sources of colonizing taxa, no clean-water macroinvertebrate taxa were recorded 30 years after the major sources of pollution ceased. Where clean-water colonisers were more readily available, significant improvements in ecological quality followed within 2–5 years of the improvements in chemical quality. Macroinvertebrate communities and hence monitoring data may thus be indicative of long past conditions or of biological isolation rather than contemporaneous chemical conditions. Combined chemical and biological data were used to explore a generic model for predicting recovery rates and success. Neither BOD₅ nor NH₃ were found to provide a consistent and meaningful prediction of either average pollution tolerance of macroinvertebrate taxa or of the number of taxa present. Long-term relationships between macroinvertebrate variables and chemical water quality variables, however, were non-linear, suggesting that water quality thresholds may have to be exceeded before biological recovery can occur. Even when chemical water quality has been improved substantially, the apparent ecological status of macroinvertebrate communities may not reflect reduced pollution levels attained until adequate time to allow for re-colonisation (possibly decades) has elapsed.     

The combined effects of treated sewage discharge and land use on rivers

Freshwater ecosystems are increasingly threatened by multiple anthropogenic stressors. Release of treated sewage effluent and pollution from agricultural or urban sources can independently reduce water quality with implications for ecological communities. However, our knowledge of the combined effects of these stressors is limited. We performed a field study to quantify the combined effect of treated sewage discharge and land use on nutrient concentrations, sewage fungus presence and communities of macroinvertebrates and benthic algae. Over three seasons in four rivers we found that a model which included an interaction between sewage pollution and time of the year (i.e. months) was the best predictor of nutrient concentrations and the abundance of algae and sewage fungus. Both macroinvertebrate and algae communities shifted downstream of sewage input. Specifically, more tolerant groups, such as cyanobacteria and oligochaetes, were more abundant. The EPT (Ephemeroptera, Plecoptera and Tricoptera) water quality score was best explained by an interaction between month and agriculture in the surrounding landscape. Overall, our results show that sewage discharge has a significant impact on water quality and benthic riverine communities, regardless of the surrounding land uses. Agricultural inputs, however, could be more important than treated sewage discharge in reducing the abundance of sensitive invertebrate taxa. We need both improvements to wastewater treatment processes and reductions in agricultural pollution to reduce threats to vulnerable freshwater communities.     

Changes in shallow lake functioning: response to climate change and nutrient reduction

Hydrobiologia - Nutrient, phytoplankton and zooplankton dynamics have been monitored intensively at Loch Leven for 34 years. The data collected reveal a decline in phosphorus concentrations,...     

A history of scientific research at Loch Leven,Kinross, Scotland

Loch Leven is a large, shallow lake in lowland Scotland, UK. Scientific research began here almost 200 years ago. Early research characterised the biodiversity and physical characteristics of the loch, providing an important historical background for future research. In the mid-1960s, this ad hoc approach was superseded by a more structured research programme under the umbrella of the International Biological Programme. This was the beginning of the Loch Leven long-term monitoring programme. Today, the results of these studies form one of the longest and most comprehensive limnological datasets for shallow freshwater lakes in the world, comprising more than 500 physical, chemical and biological variables collected at two-weekly intervals. To celebrate the 40th anniversary of the start of the long term monitoring programme, and to highlight the scientific investigations still being conducted at Loch Leven, the NERC Centre for Ecology & Hydrology (CEH) organised a symposium entitled “Loch Leven: 40 years of scientific research” in Kinross, Scotland, UK, on 11 December 2008. This examined the role of long-term monitoring in developing our understanding of the links between pollution, climate change and ecological responses in shallow lakes. This article introduces a series of papers summarising the scientific results presented at this meeting.     

Long-term effects of nutrient changes on rotifer communities in a subtropical lake

Limnology - External nutrient loading has a major influence on the abundance, composition and diversity of zooplankton communities. To clarify how the reduction of nutrient loading during lake...     

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.     

Extending the growth rate hypothesis to species development: Can stoichiometric traits help to explain the composition of macroinvertebrate communities?

Ecological stoichiometry seeks to understand the ecological consequences of elemental imbalances between consumers and their resources. Therein, the well-accepted growth rate hypothesis (GRH) states that organisms exhibiting rapid growth have higher phosphorus (P) demand – and thus lower C:P and N:P ratios – than slow growing ones, due to a higher allocation to P-rich rRNA. However, GRH has rarely been extended to other biological traits than growth, especially at the community level. In this study, we investigated whether macroinvertebrate stoichiometric traits (e.g. C:P and N:P ratios) can be linked to their development traits, and whether these stoichiometric traits are related to macroinvertebrate community assemblage under different nutrient conditions. We allocated more than 400 European taxa to different groups, defined using available information about three development-related traits: ‘life span', ‘voltinism' and ‘number of reproductive cycles per individual'. We sampled 18 invertebrate taxa in six streams exhibiting different levels of nutrient concentration and measured their stoichiometric traits. Further, we quantified invertebrate taxon abundances in these streams during an annual survey. Based on these data, we tested whether community composition regarding the developmental groups differs, depending on nutrient concentration. We found significant differences in the proportions of the developmental groups along a gradient of water N:P, in relation to their stoichiometric traits. Taxa with low C:P and N:P ratios were generally associated with faster development groups, and these taxa tended to occur at higher proportions in streams exhibiting low dissolved N:P ratios. In contrast, communities from P-poor, high dissolved N:P streams, were dominated by slowly developing taxa with high N:P ratios. Our results highlight that extending the GRH to species development rate might give some insights about the mechanisms by which nutrient concentrations in ecosystems influence consumers' community composition.     

Relevance of abiotic criteria used in German lake typology for macroinvertebrate fauna

For lake characterisation, top-down typologies are mostly used throughout Europe, including type criteria such as climate, lake area, catchment geology and conductivity. In Germany, a lake typology was applied comprising ecoregion, calcium concentration, Schindler’s ratio, stratification type and residence time. However, the relevance of these criteria for the macroinvertebrate fauna has not been conclusively demonstrated till now. Benthic invertebrate community data and related environmental parameters of pristine or near-pristine lakes in Germany were analysed by multivariate analysis techniques to elucidate which environmental parameters are reflected by invertebrate composition. Moreover, benthic invertebrate data were transformed to metrics expressing ecological attributes and species richness (summarising functional composition, diversity and sensitivity measures). Multivariate statistics were used to test whether information relevant to ordination was lost and whether variation decreases using metrics which combine data with ecological attributes. Analysis of lake-type criteria revealed that ecoregions and prevailing substrates were characterized by different taxonomic compositions of macroinvertebrates. In addition, a relationship was found between community composition and lake size. Creating a novel bottom-up lake typology based on ecoregions, lake size and prevailing substrate gives better separation of distinct macroinvertebrate communities and a higher level of homogeneity within groups compared to top-down typology or single environmental parameters alone, both on species and metrics data. Despite some data variation due to methodological differences (e.g. different sampling and sorting techniques) and interannual and seasonal variation in the data set, NMDS ordination presented well-separated groups of bottom-up lake types. Lake types were more precisely separated by species data than by metric data in both top-down and bottom-up typology. However, as information loss from species lists to calculated metrics is marginal, type-specific benthic invertebrate assemblages are reflected both on the species level and on the metric level. Species and metric data are both suitable for data ordination, while single environmental parameters affecting macroinvertebrate composition can best be obtained using metrics.     

Maximum growing depth of macrophytes in Loch Leven,Scotland, United Kingdom,in relation to historical changes in estimated phosphorus loading

Eutrophication is common in shallow lakes in lowland areas. In their natural state, most shallow lakes would have clear water and a thriving aquatic plant community. However, eutrophication often causes turbid water, high algal productivity, and low species diversity and abundance of submerged macrophytes. A key indicator of the ecological state of lake ecosystems is the maximum growing depth (MGD) of aquatic plants. However, few studies have yet quantified the relationship between changes in external phosphorus (P) input to a lake and associated variation in MGD. This study examines the relationship between these variables in Loch Leven, a shallow, eutrophic loch in Scotland, UK. A baseline MGD value from 1905 and a series of more recent MGD values collected between 1972 and 2006 are compared with estimated P loads over a period of eutrophication and recovery. The results suggest a close relationship between changes in MGD of macrophytes and changes in the external P load to the loch. Variation in MGD reflected the ‘light history’ that submerged macrophytes had been exposed to over the 5-year period prior to sampling, rather than responding to short term, within year, variations in water clarity. This suggests that changes in macrophyte MGD may be a good indicator of overall, long term, changes in water quality that occur during the eutrophication and restoration of shallow lakes.     

Impacts of multiple anthropogenic stressors on stream macroinvertebrate community composition and functional diversity

Ensuring the provision of essential ecosystem services in systems affected by multiple stressors is a key challenge for theoretical and applied ecology. Trait‐based approaches have increasingly been used in multiple‐stressor research in freshwaters because they potentially provide a powerful method to explore the mechanisms underlying changes in populations and communities. Individual benthic macroinvertebrate traits associated with mobility, life history, morphology, and feeding habits are often used to determine how environmental drivers structure stream communities. However, to date multiple‐stressor research on stream invertebrates has focused more on taxonomic than on functional metrics. We conducted a fully crossed, 4‐factor experiment in 64 stream mesocosms fed by a pristine montane stream (21 days of colonization, 21 days of manipulations) and investigated the effects of nutrient enrichment, flow velocity reduction and sedimentation on invertebrate community, taxon, functional diversity and trait variables after 2 and 3 weeks of stressor exposure. 89% of the community structure metrics, 59% of the common taxa, 50% of functional diversity metrics, and 79% of functional traits responded to at least one stressor each. Deposited fine sediment and flow velocity reduction had the strongest impacts, affecting invertebrate abundances and diversity, and their effects translated into a reduction of functional redundancy. Stressor effects often varied between sampling occasions, further complicating the prediction of multiple‐stressor effects on communities. Overall, our study suggests that future research combining community, trait, and functional diversity assessments can improve our understanding of multiple‐stressor effects and their interactions in running waters.