Ecosystem-level effects of re-oligotrophication and N:P imbalances in rivers and estuaries on a global scale

Trends and ecological consequences of phosphorus (P) decline and increasing nitrogen (N) to phosphorus (N:P) ratios in rivers and estuaries are reviewed and discussed. Results suggest that re-oligotrophication is a dominant trend in rivers and estuaries of high-income countries in the last two–three decades, while in low-income countries widespread eutrophication occurs. The decline in P is well documented in hundreds of rivers of United States and the European Union, but the biotic response of rivers and estuaries besides phytoplankton decline such as trends in phytoplankton composition, changes in primary production, ecosystem shifts, cascading effects, changes in ecosystem metabolism, etc., have not been sufficiently monitored and investigated, neither the effects of N:P imbalance. N:P imbalance has significant ecological effects that need to be further investigated. There is a growing number of cases in which phytoplankton biomass have been shown to decrease due to re-oligotrophication, but the potential regime shift from phytoplankton to macrophyte dominance described in shallow lakes has been documented only in a few rivers and estuaries yet. The main reasons why regime shifts are rarely described in rivers and estuaries are, from one hand the scarcity of data on macrophyte cover trends, and from the other hand physical factors such as peak flows or high turbidity that could prevent a general spread of submerged macrophytes as observed in shallow lakes. Moreover, re-oligotrophication effects on rivers may be different compared to lakes (e.g., lower dominance of macrophytes) or estuaries (e.g., limitation of primary production by N instead of P) or may be dependent on river/estuary type. We conclude that river and estuary re-oligotrophication effects are complex, diverse and still little known, and in some cases are equivalent to those described in shallow lakes, but the regime shift is more likely to occur in mid to high-order rivers and shallow estuaries.     

Oligotrophication as a result of planktivorous fish removal with rotenone in the small,eutrophic, Lake Mosvatn,Norway

In September 1987 the shallow, eutrophic, Lake Mosvatn was treated with rotenone to eliminate planktivorous fish (mainly whitefish,Coregonus lavaretus, L.), and the effects were studied. The first summer after treatment the zooplankton community changed markedly from rotifer dominance and few grazers, to a community with few rotifers and many grazers. Accordingly there was a fivefold increase in the biomass ofDaphnia galeata. Adult females ofD. galeata approximately doubled in weight. The decrease in rotifer biomass was probably mainly due to a loss of food by competition with the daphnids. The phytoplankton community was also markedly affected. Prior to treatment Secchi depth was 1.7 m and Chl-a 23μg l⁻¹ in the summer. After treatment there was an increase in the proportion of small and gelatinous algae and the mean chlorophyll concentration fell to 7μg Chl-a l⁻¹. Secchi depth increased to>2.3 m (bottom-sight most of the season). After the treatment there were also fewer cyanobacterial blooms. This seems to be related to oligotrophication caused indirectly by increased grazing by the zooplankton. Total nutrient concentrations were affected. Prior to treatment the mean summer concentration of total phosphate was 44μg P l⁻¹. This decreased to 29μg P l⁻¹ in the first summer and 23μg P l⁻¹ the second summer after the treatment. Total nitrogen decreased from 0.68 mg N l⁻¹ before treatment to 0.32 mg N l⁻¹ the first summer after the treatment. The phosphate loading was not reduced, therefor it can be concluded that the fish removal provided a biomanipulation which caused the more oligotrophic conditions.     

Cladoceran community responses to biomanipulation and re‐oligotrophication in Lake Vesijärvi,Finland, as inferred from remains in annually laminated sediment

1. We studied the role of zooplankton in biomanipulation and the subsequent recovery phase in the Enonselkä basin of Lake Vesijärvi, using subfossil cladocerans in annually laminated sediment. Measures to restore the Enonselkä basin included reduction in external nutrient loading and mass removal of plankti‐ and benthivorous fish. Water clarity increased and the lake changed from a eutrophic to a mesotrophic state. However, some signs of increased turbidity were observed after 5–10 years of successful recovery. 2. Annual laminae in a freeze core sample were identified and sliced, based on the seasonal succession of diatoms. Cladoceran remains and rotifer eggs were counted, and Daphnia ephippia and Eubosmina and Bosmina ephippia and carapaces were measured. Annual changes in pelagic species composition were studied with principal component analysis. Individual species abundance, size measurements and various cladoceran‐based indices or ratios (commonly used to reconstruct changes in trophic state and fish predation) were tested for change between four distinct periods: I (1985–1988) dense fish stocks, poor water quality; II (1989–1992) fish removal; III (1993–1997) low fish density, improved water quality; IV (1998–2002) slightly increased fish density and poorer water quality. 3. After the removal of fish, the mean size of Daphnia ephippia and Eubosmina crassicornis ephippia and carapaces increased significantly. In contrast, the percentage of Daphnia did not increase. When based on ephippia, the ratio Daphnia/(Daphnia + E. crassicornis) increased, but the interpretation was obscured by the tolerance of fish predation by small Daphnia and by the fact that bosminids were the preferred food of roach. Moreover, ephippial production by E. crassicornis decreased in recent years. 4. The abundance of Diaphanosoma brachyurum and Limnosida frontosa increased significantly after the fish population was reduced, while that of Ceriodaphnia and rotifers decreased. 5. The expanding littoral vegetation along with improved water clarity was clearly reflected in the concentration of littoral species in the deep sediment core. The species diversity index for the entire subfossil community also increased. 6. The period of faltering recovery was characterised by greater interannual variability and an increased percentage of rotifers. Nevertheless, the mean sizes of Daphnia ephippia and E. crassicornis ephippia and carapaces indicated a low density of fish. The deteriorating water quality was apparently related to multiple stressors in the catchment after rehabilitation, such as intensified lakeshore building, as well as to exceptional weather conditions, challenging the management methods in use.     

Relative importance of phosphorus,invasive mussels and climate for patterns in chlorophyll a and primary production in Lakes Michigan and Huron

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Recent histories of six productive lakes in the Irish Ecoregion based on multiproxy palaeolimnological evidence

Palaeolimnological data from six mesotrophic, eutrophic and hypertrophic lakes in the Irish Ecoregion, in the form of microfossil (cladocera, diatoms and pollen) and sediment chemistry data from radiometrically dated sediment cores, were used to reconstruct past variations in lake water quality and catchment conditions. Basal sediments from sediment cores from the six sites ranged in age from the late 18th century to the early 20th century. A weighted averaging partial least squares regression model was developed to reconstruct past epilimnetic total phosphorus concentrations. The results indicate that all but one of the study sites currently are in a far more productive state compared with the beginning of the sediment core record and that those same five lakes have experienced accelerated enrichment post c. 1980. Two of the sites demonstrated long-term enrichment, in one case beginning in the late 19th century, while both eutrophication and oligotrophication have occurred at three sites. The results highlight the difficulties in applying a general temporal end-point for reference conditions and demonstrate that productive lakes in the Irish Ecoregion have complex, locally specific and often long histories of enrichment. These may not be responsive to reduced external loadings of phosphorus and, as a result, restoration could prove particularly challenging. The results also provide evidence of the ways in which palaeolimnological techniques can assist implementation of the EU Water Framework Directive. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Warming and oligotrophication cause shifts in freshwater phytoplankton communities

While there is a lot of data on interactive effects of eutrophication and warming, to date, we lack data to generate reliable predictions concerning possible effects of nutrient decrease and temperature increase on community composition and functional responses. In recent years, a wide‐ranging trend of nutrient decrease (re‐oligotrophication) was reported for freshwater systems. Small lakes and ponds, in particular, show rapid responses to anthropogenic pressures and became model systems to investigate single as well as synergistic effects of warming and fertilization in situ and in experiments. Therefore, we set up an experiment to investigate the single as well as the interactive effects of nutrient reduction and gradual temperature increase on a natural freshwater phytoplankton community, using an experimental indoor mesocosm setup. Biomass production initially increased with warming but decreased with nutrient depletion. If nutrient supply was constant, biomass increased further, especially under warming conditions. Under low nutrient supply, we found a sharp transition from initially positive effects of warming to negative effects when resources became scarce. Warming reduced phytoplankton richness and evenness, whereas nutrient reduction at ambient temperature had positive effects on diversity. Our results indicate that temperature effects on freshwater systems will be altered by nutrient availability. These interactive effects of energy increase and resource decrease have major impacts on biodiversity and ecosystem function and thus need to be considered in environmental management plans.     

Overwinter changes in mass and lipid content of Perca fluviatilis and Gymnocephalus cernuus

The body condition, lipid reserves and mortality of 0 and 1 year‐old perch Perca fluviatilis and ruffe Gymnocephalus cernuus , sampled during the winter in Lake Constance, Germany, were compared. Length‐frequency analyses did not provide evidence for overwinter mortality in either species. The fresh and dry mass of perch as well as their lipid contents decreased during winter, while ruffe were heavier and contained more lipid at the end of the winter. The superior performance of ruffe was mainly attributed to its sensory capabilities, which allowed it to ingest zoobenthos throughout the winter, while the zooplankton feeding of perch was constrained by low light levels. In lakes that undergo a process of re‐oligotrophication, this advantage of ruffe over perch may be even more pronounced, since lower food supply during the growth season and thus lower fish lipid content at the start of winter is probably better tolerated by ruffe than by perch.     

Combining palaeolimnological and limnological approaches in assessing lake ecosystem response to nutrient reduction

1. Palaeolimnological data and limnological time‐series data are highly complementary. Sediment records extend time‐scales, integrate subannual variability and expand the range of sites that can be studied, but they suffer from taphonomic biases and occasionally from uncertain chronology. Observational time‐series data, on the other hand, are highly resolved but are very limited in extent both in space and time. 2. Palaeolimnological and observational data‐sets need to be combined in oligotrophication research to establish (i) the past and present status of lakes needed to identify reference conditions; (ii) changes in ecosystem state; (iii) responses to nutrient reduction; and (iv) the potential role of other factors (e.g. additional stressors, climate change) that might confound predictions of future state.     

Do phytoplankton communities correctly track trophic changes? An assessment using directly measured and palaeolimnological data

1. Measurements of total phosphorus (TP) concentrations since 1975 and a 50‐year time series of phytoplankton biovolume and species composition from Lake Mondsee (Austria) were combined with palaeolimnological information on diatom composition and reconstructed TP‐levels to describe the response of phytoplankton communities to changing nutrient conditions. 2. Four phases were identified in the long‐term record. Phase I was the pre‐eutrophication period characterised by TP‐levels of about 6 μg L^?1 and diatom dominance. Phase II began in 1966 with an increase in TP concentration followed by the invasion of Planktothrix rubescens in 1968, characterising mesotrophic conditions. Phase III, from 1976 to 1979, had the highest annual mean TP concentrations (up to 36 μg L^?1) and phytoplankton biovolumes (3.57 mm³ L^?1), although reductions in external nutrient loading started in 1974. Phases II and III saw an expansion of species characteristic of higher nutrient levels as reflected in the diatom stratigraphy. Oligotrophication (phase IV) began in 1980 when annual average TP concentration, Secchi depth and algal biovolume began to decline, accompanied by increasing concentrations of soluble reactive silica. 3. The period from 1981 to 1986 was characterised by asynchronous trends. Annual mean and maximum total phytoplankton biovolume initially continued to increase after TP concentration began to decline. Reductions in phytoplankton biovolume were delayed by about 5 years. Several phytoplankton species differed in the timing of their responses to changing nutrient conditions. For example, while P. rubescens declined concomitantly with the decline in TP concentration, other species indicative of higher phosphorus concentrations, such as Tabellaria flocculosa var. asterionelloides, tended to increase further. 4. These data therefore do not support the hypotheses that a reduction in TP concentration is accompanied by (i) an immediate decline in total phytoplankton biovolume and (ii) persistence of the species composition characterising the phytoplankton community before nutrient reduction.