Lake responses to reduced nutrient loading – an analysis of contemporary long-term data from 35 case studies

1. This synthesis examines 35 long‐term (5–35 years, mean: 16 years) lake re‐oligotrophication studies. It covers lakes ranging from shallow (mean depth <5 m and/or polymictic) to deep (mean depth up to 177 m), oligotrophic to hypertrophic (summer mean total phosphorus concentration from 7.5 to 3500 μg L^?1 before loading reduction), subtropical to temperate (latitude: 28–65°), and lowland to upland (altitude: 0–481 m). Shallow north‐temperate lakes were most abundant. 2. Reduction of external total phosphorus (TP) loading resulted in lower in‐lake TP concentration, lower chlorophyll a (chl a) concentration and higher Secchi depth in most lakes. Internal loading delayed the recovery, but in most lakes a new equilibrium for TP was reached after 10–15 years, which was only marginally influenced by the hydraulic retention time of the lakes. With decreasing TP concentration, the concentration of soluble reactive phosphorus (SRP) also declined substantially. 3. Decreases (if any) in total nitrogen (TN) loading were lower than for TP in most lakes. As a result, the TN : TP ratio in lake water increased in 80% of the lakes. In lakes where the TN loading was reduced, the annual mean in‐lake TN concentration responded rapidly. Concentrations largely followed predictions derived from an empirical model developed earlier for Danish lakes, which includes external TN loading, hydraulic retention time and mean depth as explanatory variables. 4. Phytoplankton clearly responded to reduced nutrient loading, mainly reflecting declining TP concentrations. Declines in phytoplankton biomass were accompanied by shifts in community structure. In deep lakes, chrysophytes and dinophytes assumed greater importance at the expense of cyanobacteria. Diatoms, cryptophytes and chrysophytes became more dominant in shallow lakes, while no significant change was seen for cyanobacteria. 5. The observed declines in phytoplankton biomass and chl a may have been further augmented by enhanced zooplankton grazing, as indicated by increases in the zooplankton : phytoplankton biomass ratio and declines in the chl a : TP ratio at a summer mean TP concentration of <100–150 μg L^?1. This effect was strongest in shallow lakes. This implies potentially higher rates of zooplankton grazing and may be ascribed to the observed large changes in fish community structure and biomass with decreasing TP contribution. In 82% of the lakes for which data on fish are available, fish biomass declined with TP. The percentage of piscivores increased in 80% of those lakes and often a shift occurred towards dominance by fish species characteristic of less eutrophic waters. 6. Data on macrophytes were available only for a small subsample of lakes. In several of those lakes, abundance, coverage, plant volume inhabited or depth distribution of submerged macrophytes increased during oligotrophication, but in others no changes were observed despite greater water clarity. 7. Recovery of lakes after nutrient loading reduction may be confounded by concomitant environmental changes such as global warming. However, effects of global change are likely to run counter to reductions in nutrient loading rather than reinforcing re‐oligotrophication.     

Nutrient ratios and phytoplankton community structure in the large, shallow, eutrophic, subtropical Lakes Okeechobee (Florida, USA) and Taihu (China)

Analysis of ten- and four-year datasets for the large, shallow, subtropical, and eutrophic Lakes Okeechobee (USA) and Taihu (China), respectively, suggest that resource-ratio explanations for cyanobacteria dominance may not apply to these two lakes. Datasets were examined to identify relationships between nutrient ratios [total nitrogen (TN):total phosphorus (TP) and ammonium (NH₄ ⁺):oxidized N (NO _x )] and phytoplankton community structure (as proportions of cyanobacteria and diatoms to total phytoplankton biomass). Datasets were pooled by sampling month, averaged lake-wide, and analyzed with linear regression. In Okeechobee, the cyanobacteria proportion increased and the diatom proportion decreased with increasing TN:TP. In Taihu, cyanobacteria decreased with increasing TN:TP, but the opposite trend observed for diatoms was marginally significant. Okeechobee cyanobacteria increased and diatoms decreased with increasing NH₄ ⁺:NO _x , but no significant relationships between phytoplankton and NH₄ ⁺:NO _x were observed in Taihu. Both lakes had significant relationships between phytoplankton community structure and total nutrients, but these relationships were the opposite of those expected. Relationships between phytoplankton community structure and water quality parameters from the previous month resulted in improved relationships, suggesting a predictive capability. Statistical analysis of the entire datasets (not pooled) supported these and additional relationships with other parameters, including temperature and water clarity.     

太湖浮游植物优势种长期演化与富营养化进程的关系

利用1991年至2002年每月一次的监测资料,系统分析了浮游植物优势种和生物量的周年变化情况。同时,总氮、总磷和浮游植物叶绿素a含量等相关资料也被用于解释太湖富营养化演化与浮游植物的关系。结果显示,太湖总氮、总磷、叶绿素a和生物量均呈自梅梁湾底至湖心的逐步递减趋势。在20世纪80年代末太湖刚开始富营养化时,浮游植物优势种群从硅藻转变为蓝藻。之后,浮游植物优势种群一直是蓝藻,但各年的浮游植物总生物量有变化。总氮、总磷、叶绿素a和生物量的年均值持续增长至1996年,其后有逐步下降的趋势,究其原因可能和当地政府在太湖流域的控制排污行动有关。微囊藻在太湖的占优是太湖富营养化的标志之一。研究结果说明浮游植物在大型浅水湖泊中可以作为反映富营养化进程的生态指标。     

Seasonal Patterns of Nitrogen and Phosphorus Limitation in Four German Lakes and the Predictability of Limitation Status from Ambient Nutrient Concentrations

To identify the seasonal pattern of nitrogen (N) and phosphorus (P) limitation of phytoplankton in four different lakes, biweekly experiments were conducted from the end of March to September 2011. Lake water samples were enriched with N, P or both nutrients and incubated under two different light intensities. Chlorophyll a fluorescence (Chla) was measured and a model selection procedure was used to assign bioassay outcomes to different limitation categories. N and P were both limiting at some point. For the shallow lakes there was a trend from P limitation in spring to N or light limitation later in the year, while the deep lake remained predominantly P limited. To determine the ability of in-lake N:P ratios to predict the relative strength of N vs. P limitation, three separate regression models were fit with the log-transformed ratio of Chla of the P and N treatments (Response ratio = RR) as the response variable and those of ambient total phosphorus:total nitrogen (TN:TP), dissolved inorganic nitrogen:soluble reactive phosphorus (DIN:SRP), TN:SRP and DIN:TP mass ratios as predictors. All four N:P ratios had significant positive relationships with RR, such that high N:P ratios were associated with P limitation and low N:P ratios with N limitation. The TN:TP and DIN:TP ratios performed better than the DIN:SRP and TN:SRP in terms of misclassification rate and the DIN:TP ratio had the highest R² value. Nitrogen limitation was predictable, frequent and persistent, suggesting that nitrogen reduction could play a role in water quality management. However, there is still uncertainty about the efficacy of N restriction to control populations of N₂ fixing cyanobacteria.     

TN : TP ratio and planktivorous fish do not affect nutrient-chlorophyll relationships in shallow lakes

1. In previous work, phytoplankton regulation in freshwater lakes has been associated with many factors. Among these, the ratio of total nitrogen to total phosphorus (TN : TP) has been widely proposed as an index to identify whether phytoplankton are N‐ or P‐limited. From another point of view, it has been suggested that planktivorous fish can be used to control phytoplankton. 2. Large‐scale investigations of phytoplankton biomass [measured as chlorophyll a, (chl‐a)] were carried out in 45 mid‐lower Yangtze shallow lakes to test hypotheses concerning nutrient limitation (assessed with TN : TP ratios) and phytoplankton control by planktivorous fish. 3. Regression analyses indicated that TP was the primary regulating factor and TN the second regulating factor for both annual and summer phytoplankton chl‐a. In separate nutrient–chl‐a regression analyses for lakes of different TN : TP ratios, TP was also superior to TN in predicting chl‐a at all particular TN : TP ranges and over the entire TN : TP spectrum. Further analyses found that chl‐a : TP was not influenced by TN : TP, while chl‐a : TN was positively and highly correlated to TP : TN. 4. Based on these results, and others in the literature, we argue that the TN : TP ratio is inappropriate as an index to identify limiting nutrients. It is almost impossible to specify a ‘cut‐off’ TN : TP ratio to identify a limiting nutrient for a multi‐species community because optimal N : P ratios vary greatly among phytoplankton species. 5. Lakes with yields of planktivorous fish (silver and bighead carp, the species native to China) >100 kg ha^?1 had significantly higher chl‐a and lower Secchi depth than those with yields <100 kg ha^?1. TP–chl‐a and TP–Secchi depth relationships are not significantly different between lakes with yields >100 kg ha^?1 or <100 kg ha^?1. These results indicate that the fish failed to decrease chl‐a yield or enhance Z_SD. Therefore, silver carp and bighead carp are not recommended as a biotic agent for phytoplankton control in lake management if the goal is to control the entire phytoplankton and to enhance water quality.     

Lake‐type‐specific seasonal patterns of nutrient limitation in German lakes,with target nitrogen and phosphorus concentrations for good ecological status

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Deriving nutrient targets to prevent excessive cyanobacterial densities in U.S. lakes and reservoirs

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Long-term nutrient trends and harmful cyanobacterial bloom potential in hypertrophic Lake Taihu,China

Rapid economic development in China’s Lake Taihu basin during the past four decades has accelerated nitrogen (N) and phosphorus (P) loadings to the lake. This has caused a shift from mesotrophic to hypertrophic conditions, symptomized by harmful cyanobacterial blooms (CyanoHABs). The relationships between phytoplankton biomass as chlorophyll a (Chla) and nutrients as total nitrogen (TN) and total phosphorus (TP) were analyzed using historical data from 1992 to 2012 to link the response of CyanoHAB potential to long-term nutrient changes. Over the twenty year study period, annual mean Chla showed significantly positive correlations with both annual mean TN and TP (P < 0.001), reflecting a strong phytoplankton biomass response to changes in nutrient inputs to the lake. However, phytoplankton biomass responded slowly to annual changes in TN after 2002. There was not a well-defined or significant relationship between spring TN and summertime Chla. The loss of a significant fraction of spring N loading due to denitrification likely weakened this relationship. Bioavailability of both N and P during the summer plays a key role in sustaining cyanobacterial blooms. The frequency of occurrence of bloom level Chla (>20 μg L^?1) was compared to TN and TP to determine nutrient-bloom thresholds. A decline in bloom risk is expected if TN remains below 1.0 mg L^?1 and TP below 0.08 mg L^?1.     

Nutrient-phytoplankton relationships in a tropical meromictic soda lake

Seasonal variation through one year in total nitrogen (TN), total phosphorus (TP), phytoplankton biomass, phytoplankton species composition and other environmental factors were examined in Lake Sonachi, a tropical meromictic soda lake. Mean concentrations of TN and TP were 11 000 µg N l^-1 and 100 µg P l^-1, respectively. Maximum concentrations of TN and TP occurred in the monimolimnion. Phytoplankton biomass ranged from 350 to 1260 mg m^-3. Synechococcus bacillaris, a small coccoid cyanophyte, dominated the phytoplankton. The mean chlorophyll a concentration of 37 mg · m^-3 was a modest value when compared with those of other tropical soda lakes. High TN:TP ratios indicated phosphorus limitation in the lake.     

Nutrient limitation of phytoplankton communities in Subarctic lakes and ponds in Wapusk National Park, Canada

We determined the limiting nutrient of phytoplankton in 21 lakes and ponds in Wapusk National Park, Canada, using nutrient enrichment bioassays to assess the response of natural phytoplankton communities to nitrogen and phosphorus additions. The goal was to determine whether these Subarctic lakes and ponds were nutrient (N or P) limited, and to improve the ability to predict future impacts of increased nutrient loading associated with climate change. We found that 38% of lakes were not limited by nitrogen or phosphorus, 26% were co-limited by N and P, 26% were P-limited and 13% were N-limited. TN/TP, DIN/TP and NO₃ ⁻/TP ratios from each lake were compared to the Redfield ratio to predict the limiting nutrient; however, these predictors only agreed with 29% of the bioassay results, suggesting that nutrient ratios do not provide a true measure of nutrient limitation within this region. The N-limited lakes had significantly different phytoplankton community composition with more chrysophytes and Anabaena sp. compared to all other lakes. N and P limitation of phytoplankton communities within Wapusk National Park lakes and ponds suggests that increased phytoplankton biomass may result in response to increased nutrient loading associated with environmental change.     

Cascading Trophic Interactions from Fish to Bacteria and Nutrients after Reduced Sewage Loading: An 18-Year Study of a Shallow Hypertrophic Lake

The effects of major reductions in organic matter, total phosphorus (TP), and total nitrogen (TN) loading on the chemical environment, trophic structure, and dynamics of the hypertrophic, shallow Lake Søbygård were followed for 18 years. After the reduction in organic matter loading in 1976, the lake initially shifted from a summer clear-water state, most likely reflecting high grazing pressure by large Daphniaspecies, to a turbid state with extremely high summer mean chlorophyll a (up to 1400 μg L^{? 1}), high pH (up to 10.2), and low zooplankton grazing. Subsequently, a more variable state with periodically high grazing rates on phytoplankton and bacteria was established. Changes in zooplankton abundance and grazing could be attributed to variations in cyprinid abundance due to a fish kill (probably as a consequence of oxygen depletion) and pH-induced variations in fish recruitment and fry survival. The results suggest strong cascading effects of fish on the abundance and size of zooplankton and phytoplankton and on phytoplankton production. A comparatively weak cascading effect on ciliates and bacterioplankton is suggested. Due to high internal loading, only minor changes were observed in lake-water TP after a reduction in external TP loading of approximately 80% in 1982; net retention of TP was still negative 13 years after the loading reduction, despite a short hydraulic retention time of a few weeks. TN, however, decreased proportionally to the TN-loading reduction in 1987, suggesting a fast N equilibration. Only minor improvement in the environmental state of the lake has been observed. We suggest that another decade will be required before the lake is in equilibrium with present external P loading.     

Long-term response of a shallow, moderately flushed lake to reduced external phosphorus and nitrogen loading

1. The responses of nutrient concentrations, plankton, macrophytes and macrozoobenthos to a reduction in external nutrient loading and to contemporary climatic change were studied in the shallow, moderately flushed Lake Müggelsee (Berlin, Germany). Weekly to biweekly data from 1979 to 2003 were compared with less frequently collected historical data. 2. A reduction of more than 50% in both total phosphorus (TP) and total nitrogen (TN) loading from the hypertrophic (1979–90) to the eutrophic period (1997–2003) was followed by an immediate decline in TN concentrations in the lake. TP concentrations only declined during winter and spring. During summer, phosphorus (P) release from the sediments was favoured by a drastic reduction in nitrate import. Therefore, Müggelsee acted as a net P source for 6 years after the external load reduction despite a mean water retention time of only 0.1–0.16 years. 3. Because of the likely limitation by P in spring and nitrogen (N) in summer, phytoplankton biovolume declined immediately after nutrient loading was reduced. The formerly dominant cyanobacteria (Oscillatoriales) Limnothrix redekei and Planktothrix agardhii disappeared, but the mean biovolume of the N₂‐fixing species Aphanizomenon flos‐aquae remained constant. 4. The abundance of Daphnia spp. in summer decreased by half, while that of cyclopoid copepod species increased. Abundances of benthic macroinvertebrates (mainly chironomids) decreased by about 80%. A resource control of both phytoplankton and zooplankton is indicated by significant positive correlations between nutrient concentrations and phytoplankton biovolume and between phytoplankton and zooplankton biomass. 5. Water transparency in spring increased after nutrient reduction and resulted in re‐colonisation of the lake by Potamogeton pectinatus. However, this process was severely hampered by periphyton shading and grazing by waterfowl and fish. 6. Water temperatures in Müggelsee have increased in winter, early spring and summer since 1979. The earlier development of the phytoplankton spring bloom was associated with shorter periods with ice cover, while direct temperature effects were responsible for the earlier development of the daphnid maximum in spring.     

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.     

Zooplankton biomass rarely improves predictions of chlorophyll concentration in Canadian Shield lakes that vary in pH

To determine the frequency with which zooplankton influence chlorophyll a (Chla) levels, we explored annually-averaged data from oligotrophic and mesotrophic lakes that differed in morphometry, total phosphorus (TP) concentrations, and zooplankton community composition due to pH. The data were divided into two sets according to the type of filter used to collect chlorophyll. Residuals of the Chla: TP regressions were not related to lake morphometry, TN content, water clarity or pH. In the first data set there were no consistent relationships between residuals in Chla and twelve grazer biomass variables for 37 of the 38 lakes. The single exception had a very large population of Daphnia dubiaand low concentrations of Chla for its TP. In the second data set, 3 of 25 lakes had exceptionally low Chla concentrations for their TP. These lakes were acidic (pH < 6) and had very large biomasses of Holopedium gibberumcorrelated with negative Chla residuals, indicating significant grazing. At pH > 6, Daphnia spp. strongly influenced the significant correlations. We conclude that zooplankton contribute to the prediction of Chla beyond that possible by TP alone in acidic and non-acidic Canadian Shield Lakes, but evidence for strong suppression of chlorophyll by grazers was relatively rare (4 of 63 cases) on annual time steps.     

Nutrient–chlorophyll relationships in tropical–subtropical lakes: do temperate models fit?

In tropical lakes relatively little is known about the general relationship between nutrient concentration and phytoplankton biomass. Using data from 192 lakes from tropical and subtropical regions we examine the relationship between total P (TP) and chlorophyll (Chl). The lakes are all located between 30° S to 31° N include systems in Asia, Africa, and North and South America but are dominated by Brazilian (n=79) and subtropical N. American (n=67) systems. The systems vary in morphometry (mean depth and lake area), trophic state as well total N (TN) to␣total P (TP) ratios and light extinction. Despite a nearly 500-fold range in TP concentrations (2–970 μg P l⁻¹), there was a poorer relationship between log TP and log Chl (r ²=0.42) than is generally observed for temperate systems from either narrow or broad geographic regions. N limitation is not a likely explanation for the relatively weak TP–Chl relationship in the tropical–subtropical systems. Systems had high average TN:TP ratios and neither a multiple regression with log TP and log TN nor separating systems with high TN:TP (>17 by weight) improved the predictive power of the log TP–log Chl relationship.     

Does high nitrogen loading prevent clear‐water conditions in shallow lakes at moderately high phosphorus concentrations?

1. The effect of total nitrogen (TN) and phosphorus (TP) loading on trophic structure and water clarity was studied during summer in 24 field enclosures fixed in, and kept open to, the sediment in a shallow lake. The experiment involved a control treatment and five treatments to which nutrients were added: (i) high phosphorus, (ii) moderate nitrogen, (iii) high nitrogen, (iv) high phosphorus and moderate nitrogen and (v) high phosphorus and high nitrogen. To reduce zooplankton grazers, 1⁺ fish (Perca fluviatilis L.) were stocked in all enclosures at a density of 3.7 individuals m^?2. 2. With the addition of phosphorus, chlorophyll a and the total biovolume of phytoplankton rose significantly at moderate and high nitrogen. Cyanobacteria or chlorophytes dominated in all enclosures to which we added phosphorus as well as in the high nitrogen treatment, while cryptophytes dominated in the moderate nitrogen enclosures and the controls. 3. At the end of the experiment, the biomass of the submerged macrophytes Elodea canadensis and Potamogeton sp. was significantly lower in the dual treatments (TN, TP) than in single nutrient treatments and controls and the water clarity declined. The shift to a turbid state with low plant coverage occurred at TN >2 mg N L^?1 and TP >0.13–0.2 mg P L^?1. These results concur with a survey of Danish shallow lakes, showing that high macrophyte coverage occurred only when summer mean TN was below 2 mg N L^?1, irrespective of the concentration of TP, which ranged between 0.03 and 1.2 mg P L^?1. 4. Zooplankton biomass and the zooplankton : phytoplankton biomass ratio, and probably also the grazing pressure on phytoplankton, remained overall low in all treatments, reflecting the high fish abundance chosen for the experiment. We saw no response to nutrition addition in total zooplankton biomass, indicating that the loss of plants and a shift to the turbid state did not result from changes in zooplankton grazing. Shading by phytoplankton and periphyton was probably the key factor. 5. Nitrogen may play a far more important role than previously appreciated in the loss of submerged macrophytes at increased nutrient loading and for the delay in the re‐establishment of the nutrient loading reduction. We cannot yet specify, however, a threshold value for N that would cause a shift to a turbid state as it may vary with fish density and climatic conditions. However, the focus should be widened to use control of both N and P in the restoration of eutrophic shallow lakes.     

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

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

Phytoplankton composition modifies predator-driven life history evolution in Daphnia

Organisms experience competing selective pressures, which can obscure the mechanisms driving evolution. Daphnia ambigua is found in lakes where a predator, the alewife (Alosa pseudoharengus) either does (anadromous) or does not (landlocked) migrate between marine and freshwater. We previously reported an association between alewife variation and life history evolution in Daphnia. However, differences in alewife migration indirectly influence phytoplankton composition for Daphnia. In ‘anadromous lakes’, Daphnia are present in the spring and experience abundant high-quality green algae. Intense predation by young-of-the-year anadromous alewife quickly eliminates these Daphnia populations by early summer. Daphnia from ‘landlocked lakes’ and lakes without alewife (‘no alewife lakes’) are present during the spring and summer and are more likely to experience high concentrations of sub-optimal cyanobacteria during the summer. To explore links between predation, resources, and prey evolution, we reared third-generation laboratory-born Daphnia from all lake types on increasing cyanobacteria concentrations. We observed several significant ‘lake type × resource’ interactions whereby the differences among lake types depended upon cyanobacteria concentrations. Daphnia from anadromous lakes developed faster, were larger at maturation, produced more offspring, and had higher intrinsic rates of increase in the absence of cyanobacteria. Such trends disappeared or reversed as cyanobacteria concentration was increased because Daphnia from anadromous lakes were more strongly influenced by the presence of cyanobacteria. Our results argue that alewife migration and phytoplankton composition both play a role in Daphnia evolution.     

Does stocking of filter-feeding fish for production have a cascading effect on zooplankton and ecological state? A study of fourteen (sub)tropical Chinese reservoirs with contrasting nutrient concentrations

Stocking of filter-feeding fish is a common tool used in Chinese reservoirs to increase fish production because of low natural recruitment. Whether such stocking has important negative effects on zooplankton with cascading effects on phytoplankton is debated. We compared the zooplankton communities in fourteen reservoirs with different nutrient concentrations and fish densities. Both chlorophyll a (Chla) and fish catch were positively related with total phosphorus (TP), whereas zooplankton biomass did not show a similar relationship with TP. Zooplankton seemed to be influenced by fish as high fish catches coincided with a low proportion of calanoids of the total copepod biomass, a high proportion of rotifers of the total zooplankton biomass, a low zooplankton:phytoplankton biomass ratio, and the absence of Daphnia irrespective of TP concentration. Both zooplankton biomass and most of the zooplankton:phytoplankton biomass ratios were among the lowest reported in the literature for the nutrient range studied. Furthermore, the Chla:TP ratio was higher than what is typically observed in temperate lakes. We conclude that top-down control of zooplankton is of key importance in reservoirs in South China where frequent stocking of filter-feeding fish seems to contribute to poor water quality in the form of higher algal biomass and reduced clarity.     

Critical N:P ratio for cyanobacteria and N2-fixing species in the large shallow temperate lakes Peipsi and Võrtsjärv, North-East Europe

In the 1990s a sharp decrease in nitrogen loading occurred in Estonian rivers, bringing about a reduction of the nitrogen-to-phosphorus ratio (N:P ratio) in the large shallow lakes, Peipsi (3,555 km², mean depth 7.1 m) and Võrtsjärv (270 km², 2.8 m). The average mass ratio of total nitrogen (TN) and total phosphorus (TP) in Võrtsjärv (45) was about twice as high as that in Peipsi (22). In Peipsi, the N₂-fixing Gloeotrichia echinulata, Aphanizomenon flos-aquae and Anabaena species prevailed in the summer phytoplankton, while in Võrtsjärv the dominant cyanobacteria were Limnothrix planktonica, L. redekei and Planktolyngbya limnetica, which cannot fix N₂; the main N₂-fixing taxa Aphanizomenon skujae and Anabaena sp. seldom gained dominance. In May–October the critical TN:TP mass ratio, below which N₂-fixing cyanobacteria (Nfix) achieved high biomasses, was ～40 in Võrtsjärv and ～30 in Peipsi. The percentages of both total cyanobacteria (CY) and Nfix (CY% and Nfix%) in Peipsi achieved their maximum values at an N:P mass ratio at or below 20 for both TN:TP and Nmin:SRP. In Võrtsjärv, the TN:TP supporting a high Nfix% was between 30 and 40 and the N_min:SRP supporting this high percentage was in the same range as that in Peipsi (<20), though the maximum Nfix% values in Võrtsjärv (69%) were much lower than in Peipsi (96%). The Nmin:SRP ratio explained 77% of the variability in Nfix% in May–October. The temperature dependence of Nfix% approximated to the maximum function type, with an upper limiting value at a certain water temperature, and this was most distinct in May–October. The critical TN:TP ratios obtained from our study (roughly 30 for Peipsi and 40 for Võrtsjärv) are much higher than the Redfield N:P mass ratio routinely considered (7). Our results represent valuable guidelines for creating effective management strategies for large shallow lakes. They provide a basis for stressing the urgent need to decrease phosphorus loading and to keep the in-lake P concentration low, and not to implement nitrogen reduction measures without a simultaneous decrease of phosphorus concentration.