Benthic-planktonic coupling, regime shifts, and whole-lake primary production in shallow lakes

Alternative stable states in shallow lakes are typically characterized by submerged macrophyte (clear-water state) or phytoplankton (turbid state) dominance. However, a clear-water state may occur in eutrophic lakes even when macrophytes are absent. To test whether sediment algae could cause a regime shift in the absence of macrophytes, we developed a model of benthic (periphyton) and planktonic (phytoplankton) primary production using parameters derived from a shallow macrophyte-free lake that shifted from a turbid to a clear-water state following fish removal (biomanipulation). The model includes a negative feedback effect of periphyton on phosphorus (P) release from sediments. This in turn induces a positive feedback between phytoplankton production and P release. Scenarios incorporating a gradient of external P loading rates revealed that (1) periphyton and phytoplankton both contributed substantially to whole-lake production over a broad range of external P loading in a clear-water state; (2) during the clear-water state, the loss of benthic production was gradually replaced by phytoplankton production, leaving whole-lake production largely unchanged; (3) the responses of lakes to biomanipulation and increased external P loading were both dependent on lake morphometry; and (4) the capacity of periphyton to buffer the effects of increased external P loading and maintain a clear-water state was highly sensitive to relationships between light availability at the sediment surface and the of P release. Our model suggests a mechanism for the persistence of alternative states in shallow macrophyte-free lakes and demonstrates that regime shifts may trigger profound changes in ecosystem structure and function.     

Regime shift in a coastal marine ecosystem

We demonstrate changes in ecosystem stable states in a coastal lagoon that are consistent with what a regime shift would hypothesize. In the nutrient-stressed Ringk?bing Fjord, Denmark, a small change in one variable (salinity) facilitated by a change in sluice management, caused a sudden regime shift from a bottom-up controlled turbid state, into a top-down controlled clear-water state. The change in dominating pathway of organic matter production from pelagic turnover to benthic-pelagic coupling was facilitated by new recruitment and growth of existing suspension-feeding clams, Mya arenaria. With the invasion of clams, benthic grazing became the key feature of the biological structure. Phytoplankton composition and zooplankton abundance were also affected by the change in biological structure. The small, but sudden, increase in salinity caused by the change in sluice management led to a dramatic reduction in biomass and coverage of benthic vegetation and thus affected herbivorous waterbird populations. In recent years, plant coverage has been increasing, as can be expected with increased water transparency. The regime shift has some major implications for coastal water management and revealed some conflicts between different types of nature and environmental protection legislation.     

Evaluating the trophic status of a shallow polluted lake,lake Ioannina,Greece

Lake Ioannina in western Greece is a shallow lake, charged by springs as well as untreated discharges of the nearby urban area. Physicochemical and biological parameters have been evaluated verifying the deterioration of the lake aquatic environment. The water quality and the trophic status of the lake are greatly altered during the last ten years while the total phosphorus critical loading is already at a dangerous state.     

罗非鱼-附着藻-沉水植物相互关系研究进展

在浅水富营养化湖泊中,沉水植被是决定湖泊清水态或混水态的关键因子。而附着藻对沉水植物强烈的遮阴作用以及对碳源、营养盐等资源强烈的竞争,成为限制沉水植物群落生长和发展的关键因子。罗非鱼作为一种杂食性鱼类,具有牧食附着藻的能力,其下行效应（top-down effect）可以在一定程度上减轻附着藻对沉水植物生长的这种不利影响。因此,作为我国南方水体中的优势种类,适当种群密度的罗非鱼在富营养化浅水湖泊生态修复过程中是可加以利用,并在一定程度上抑制了附着藻的生长和发展,有利于浅水湖泊的生态修复和管理。同时,罗非鱼也具有通过摄食、排泄等活动加速水体氮、磷营养盐再生,牧食浮游动物、沉水植物等不利的一面。因此,在综合考虑多种因素条件下,需要对罗非鱼-附着藻-沉水植物三者之间的相互关系进行深入研究,探讨生态系统对罗非鱼的响应,这对我国南方浅水富营养化湖泊的生态恢复与管理,尤其是沉水植被的重建与保护具有重要的理论意义和实践价值。     

Dynamic model of Lake Chozas (León, NW Spain)—Decrease in eco-exergy from clear to turbid phase due to introduction of exotic crayfish

We developed a dynamic model of the phosphorus cycle in Lake Chozas, a small shallow water body in León (NW Spain). The calibrated model simulated seasonal dynamics of phosphorus concentrations in major components of the lake's ecological network before and after 1997, the year when an invasive allochthonous crustacean, the Louisiana red swamp crayfish (Procambarus clarkii), was introduced into the lake. The shift from clean to turbid phase, due to grazing by crayfish on submerged vegetation, caused a gradual decrease in eco-exergy, reflecting an increase in entropy, related to breakdown of ecosystem internal equilibria. This case study verifies the hypothesis of Marchi et al. (2010) that, after an initial relatively stable state, the allochthonous species may cause an increase in entropy indicating perturbation of the ecosystem.     

Use of thermodynamic indices as ecological indicators of the development state of lake ecosystems: 2. Exergy and specific exergy indices

The effectiveness of exergy and specific exergy indices as ecological indicators of the trophic state of lake ecosystems is here tested on a small homogeneous set of shallow lakes which, in spite of their similar nutrient concentrations, morphology and hydrology, show a different trophic state and structure, species composition and abundance. The findings reveal that exergy and specific exergy indices have good negative correlation with phytoplankton biomass and Carlson's trophic state index (TSI) and strong positive correlation to water transparency (the relationship between exergy and eutrophication is clearer if the exergy refers to surface units, rather than volume units) and, hence, that they may be used as ecological indicators of the trophic state of lake ecosystems. The relationship between the responses of the thermodynamic approach and other conventional trophic classification methods (Vollenweider's eutrophication model based on phosphorus loading, the Hillbrich-Ilkowska method and the Vollenweider–OECD classification criterion) previously applied to Lake Trasimeno, was also investigated. The decreasing trend of exergy and specific exergy indices with eutrophication increase appears to be essentially due to the change in species composition and trophic structure, rather than to a different trophic potentiality of the ecosystems investigated. Concerning the identification of the environmental factors responsible for exergy and specific exergy trends, the coherence of the correlation structure between water depth, TSI, exergy and specific exergy indices, suggests that the lake's mean water depth plays a significant role in determining the changes in trophic structure and state (and consequently in exergetic indices) within the set of lakes examined and emphasises the importance of lake morphology in the development and ageing of lake ecosystems.     

Linking species- and ecosystem-level impacts of climate change in lakes with a complex and a minimal model

To study the interaction between species- and ecosystem-level impacts of climate change, we focus on the question of how climate-induced shifts in key species affect the positive feedback loops that lock shallow lakes either in a transparent, macrophyte-dominated state or, alternatively, in a turbid, phytoplankton-dominated state. We hypothesize that climate warming will weaken the resilience of the macrophyte-dominated clear state. For the turbid state, we hypothesize that climate warming and climate-induced eutrophication will increase the dominance of cyanobacteria. Climate change will also affect shallow lakes through a changing hydrology and through climate change-induced eutrophication. We study these phenomena using two models, the full ecosystem model PCLake and a minimal dynamic model of lake phosphorus dynamics. Quantitative predictions with the complex model show that changes in nutrient loading, hydraulic loading and climate warming can all lead to shifts in ecosystem state. The minimal model helped in interpreting the non-linear behaviour of the complex model. The main output parameters of interest for water quality managers are the critical nutrient loading at which the system will switch from clear to turbid and the much lower critical nutrient loading – due to hysteresis – at which the system switches back. Another important output parameter is the chlorophyll-a level in the turbid state. For each of these three output parameters we performed a sensitivity analysis to further understand the dynamics of the complex model PCLake. This analysis showed that our model results are most sensitive to changes in temperature-dependence of cyanobacteria, planktivorous fish and zooplankton. We argue that by combining models at various levels of complexity and looking at multiple aspects of climate changes simultaneously we can develop an integrated view of the potential impact of climate change on freshwater ecosystems.     

Sediment resuspension by wind in a shallow lake of Esteros del Iberá (Argentina): a model based on turbidimetry

A model based on empirical relationships is used to study frequency and magnitude of the sediment resuspension by wind-induced waves. The model has been developed for Laguna Galarza, a mesotrophic round-shaped shallow lake located in Esteros del Iberá wetland. Given the logistic and accessibility difficulties of this macrosystem, the installation of automated field stations facilitated continuous data acquisition. Using the wave theory, a daily spatial model of resuspension was built from simultaneous time series of hourly measurements of infrared nephelometric turbidity, wind speed and wind direction. The model was used to predict total suspended solids in another lake of the wetland (Laguna Iberá) showing a good agreement with observed field values, even although Laguna Iberá has a more irregular contour and a eutrophic state. Finally, we apply the model to discuss the ecological impacts of resuspension on the distribution of the littoral communities and to characterize the composition of the particulate suspended matter of the limnetic ecosystem. The model was useful to simulate the possible implications of the recent alterations of the wetland water level on the resuspension regime of the open water bodies.     

Integrated model projections of climate change impacts on a North American lake

Climate change is likely to impact terrestrial and aquatic ecosystems via numerous physical and biological mechanisms. This study outlines a framework for projecting potential impacts of climate change on lakes using linked environmental models. Impacts of climate drivers on catchment hydrology and thermal balance in Onondaga Lake (New York State) are simulated using mechanistic models HSPF and UFILS4. Outputs from these models are fed into a lake ecosystem model, developed in AQUATOX. Watershed simulations project increases in the magnitude of peak flows and consequent increases in catchment nutrient export as the magnitude of extreme precipitation events increases. This occurs concurrently with a decrease in annual stream discharge as a result of increased evapotranspiration. Simulated lake water temperatures increase by as much as 5 °C during the 2040-2069 time period, accompanied by a prolonging of the duration of summer stratification. Projected changes include shifts in the timing of nutrient cycling between lake sediments and water column. Plankton taxa projected to thrive under climate change include green algae and Bosmina longirostris. Responses for species at higher trophic levels are mixed. Benthic macroinvertebrates may either prosper (zebra mussels) or decline (chironomids), while fish (e.g., gizzard shad) exhibit high seasonal variability without any clear trend.     

Food web structure and trophic interactions of the tropical highland lake Hayq, Ethiopia

We generated a mass-balance model to figure out the food web structure and trophic interactions of the major functional groups of the Ethiopian highland Lake Hayq. Moreover, the study lay down a baseline data for future ecosystem-based investigations and management activities. Extensive data collection has been taken place between October 2007 and May 2009. Ecotrophic efficiency (EE) of several functional groups including phytoplankton (0.8) and detritus (0.85) was high indicating the utilization of the groups within the system. However, the EE of Mesocyclops (0.03) and Thermocyclops (0.30) was very low implying these resources were rather a ‘sink’ in the trophic hierarchy. Flows based on aggregated trophic level sensu Lindeman revealed the importance of both phytoplankton and detritus to higher trophic levels. The computed average transfer efficiency of 11.5% for the first four trophic levels was within the range for highly efficient African lakes. The primary production to respiration (P/R) ratio (1.05) of Lake Hayq indicates the maturity of the ecosystem. We also modeled the food-web by excluding Tilapia and reduced phytoplankton biomass to get insight into the mass balance before Tilapia was introduced. The analysis resulted in a lower system omnivory index (SOI = 0.016) and a reduced P/R ratio (0.13) that described the lake as immature ecosystem, suggesting the introduction of Tilapia might have contributed to the maturity of the lake. Tilapia in Lake Hayq filled an ecological empty niche of pelagic planktivores, and contributed for the better transfer efficiency observed from primary production to fish yield.     

Warming modifies trophic cascades and eutrophication in experimental freshwater communities

Climate warming is occurring in concert with other anthropogenic changes to ecosystems. However, it is unknown whether and how warming alters the importance of top-down vs. bottom-up control over community productivity and variability. We performed a 16-month factorial experimental manipulation of warming, nutrient enrichment, and predator presence in replicated freshwater pond mesocosms to test their independent and interactive impacts. Warming strengthened trophic cascades from fish to primary producers, and it decreased the impact of eutrophication on the mean and temporal variation of phytoplankton biomass. These impacts varied seasonally, with higher temperatures leading to stronger trophic cascades in winter and weaker algae blooms under eutrophication in summer. Our results suggest that higher temperatures may shift the control of primary production in freshwater ponds toward stronger top-down and weaker bottom-up effects. The dampened temporal variability of algal biomass under eutrophication at higher temperatures suggests that warming may stabilize some ecosystem processes.     

Identifying 120 years of decline in ecosystem structure and maturity of Great South Bay, New York using the Ecopath modelling approach

Mass balanced models yield valuable information regarding ecological function and delivery of ecosystem services, but often rely on data collected well before many species were reduced to fractions of their original abundance. Lagoonal systems, such as Great South Bay (GSB), NY, sit on the interface of terrestrial and marine ecosystems and are prone to anthropogenic stressors but proximity to land also makes the presence of data regarding historic populations and structure more likely. To quantify over a century of ecosystem change, Ecopath models were developed for GSB at each of four time periods where commercial and scientific data exist: 1880s, 1930s, 1980s and 2000s. The results indicated that the GSB has experienced a decline in ecosystem maturity, loss of top keystone predators, a decline in connectivity to the ocean though the reduction of migratory species and increasing dominance of low trophic level organisms. These changes undermine the delivery of ecosystem services, increase conflicts over limited resources and suggest that present day restoration targets fail to recognize appropriate baselines. We discuss the role of stochastic events, which result in state changes that could be defined as regime shifts, and ecosystem connectivity to the long-term stability of lagoonal systems.     

Estimating the critical phosphorus loading of shallow lakes with the ecosystem model PCLake: Sensitivity, calibration and uncertainty

There is a vast body of knowledge that eutrophication of lakes may cause algal blooms. Among lakes, shallow lakes are peculiar systems in that they typically can be in one of two contrasting (equilibrium) states that are self-stabilizing: a ‘clear’ state with submerged macrophytes or a ‘turbid’ state dominated by phytoplankton. Eutrophication may cause a switch from the clear to the turbid state, if the P loading exceeds a critical value. The ecological processes governing this switch are covered by the ecosystem model PCLake, a dynamic model of nutrient cycling and the biota in shallow lakes. Here we present an extensive analysis of the model, using a three-step procedure. (1) A sensitivity analysis revealed the key parameters for the model output. (2) These parameters were calibrated on the combined data on total phosphorus, chlorophyll-a, macrophytes cover and Secchi depth in over 40 lakes. This was done by a Bayesian procedure, giving a weight to each parameter setting based on its likelihood. (3) These weights were used for an uncertainty analysis, applied to the switchpoints (critical phosphorus loading levels) calculated by the model. The model was most sensitive to changes in water depth, P and N loading, retention time and lake size as external input factors, and to zooplankton growth rate, settling rates and maximum growth rates of phytoplankton and macrophytes as process parameters. The results for the ‘best run’ showed an acceptable agreement between model and data and classified nearly all lakes to which the model was applied correctly as either ‘clear’ (macrophyte-dominated) or ‘turbid’ (phytoplankton-dominated). The critical loading levels for a standard lake showed about a factor two uncertainty due to the variation in the posterior parameter distribution. This study calculates in one coherent analysis uncertainties in critical phosphorus loading, a parameter that is of great importance to water quality managers.     

On the sudden disappearance of Egeria densa from a Ramsar wetland site of Southern Chile: A climatic event trigger model

Contemporary shallow lakes theory proposes that these ecosystems may experience abrupt regime shifts due to small changes in controlling variables or triggers. So far, these triggers have been related mostly to nutrients as the immediate driver. During May 2004 the río Cruces wetland, a Ramsar site located in Southern Chile, underwent a major regime shift, from a clear water state, vastly dominated by the invasive macrophyte Egeria densa, to a turbid water state. In this article we show, through the analysis of long-term meteorological data that late fall 2004 was anomalous due to the presence of a high-pressure cell that persisted most of the month of May over Southern Chile. This climatic event caused an almost complete absence of precipitations and lower temperatures during this period, including several freezing nights. Eco-physiological experiments showed that 6 h exposure to desiccation kill the macrophyte. We developed a simple-biology dynamic model, under Stella Research 9.1, to show that the climatic anomaly of May 2004, plus the increased sedimentation of the wetland's floodplains, and the associated response of E. densa, explains its sudden disappearance from río Cruces wetland.     

Increased spatial variance accompanies reorganization of two continental shelf ecosystems

Phase transitions between alternate stable states in marine ecosystems lead to disruptive changes in ecosystem services, especially fisheries productivity. We used trawl survey data spanning phase transitions in the North Pacific (Gulf of Alaska) and the North Atlantic (Scotian Shelf) to test for increases in ecosystem variability that might provide early warning of such transitions. In both time series, elevated spatial variability in a measure of community composition (ratio of cod [Gadus sp.] abundance to prey abundance) accompanied transitions between ecosystem states, and variability was negatively correlated with distance from the ecosystem transition point. In the Gulf of Alaska, where the phase transition was apparently the result of a sudden perturbation (climate regime shift), variance increased one year before the transition in mean state occurred. On the Scotian Shelf, where ecosystem reorganization was the result of persistent overfishing, a significant increase in variance occurred three years before the transition in mean state was detected. However, we could not reject the alternate explanation that increased variance may also have simply been inherent to the final stable state in that ecosystem. Increased variance has been previously observed around transition points in models, but rarely in real ecosystems, and our results demonstrate the possible management value in tracking the variance of key parameters in exploited ecosystems.     

Benthic algal production across lake size gradients: interactions among morphometry, nutrients, and light

Attached algae play a minor role in conceptual and empirical models of lake ecosystem function but paradoxically form the energetic base of food webs that support a wide variety of fishes. To explore the apparent mismatch between perceived limits on contributions of periphyton to whole-lake primary production and its importance to consumers, we modeled the contribution of periphyton to whole-ecosystem primary production across lake size, shape, and nutrient gradients. The distribution of available benthic habitat for periphyton is influenced by the ratio of mean depth to maximum depth (DR = z/ z(max)). We modeled total phytoplankton production from water-column nutrient availability, z, and light. Periphyton production was a function of light-saturated photosynthesis (BPmax) and light availability at depth. The model demonstrated that depth ratio (DR) and light attenuation strongly determined the maximum possible contribution of benthic algae to lake production, and the benthic proportion of whole-lake primary production (BPf) declined with increasing nutrients. Shallow lakes (z < or =5 m) were insensitive to DR and were dominated by either benthic or pelagic primary productivity depending on trophic status. Moderately deep oligotrophic lakes had substantial contributions by benthic primary productivity at low depth ratios and when maximum benthic photosynthesis was moderate or high. Extremely large, deep lakes always had low fractional contributions of benthic primary production. An analysis of the world's largest lakes showed that the shapes of natural lakes shift increasingly toward lower depth ratios with increasing depth, maximizing the potential importance of littoral primary production in large-lake food webs. The repeatedly demonstrated importance of periphyton to lake food webs may reflect the combination of low depth ratios and high light penetration characteristic of large, oligotrophic lakes that in turn lead to substantial contributions of periphyton to autochthonous production.     

Trophic relationships in the recently impounded Bagré reservoir in Burkina Faso

The trophic dynamics of Bagré reservoir which has been recently impounded in Burkina Faso was based on the data collected during 1997–1998 period using the Ecopath model and software. Total fish biomass is 22.63 t km⁻² and mainly represents trophic levels (TLs) 2 and 3. The trophic food chain is relatively long and the overall transfer efficiency is quite low. Grazing foodweb based on primary producers is prominent in the reservoir ecosystem and detritus plays a less significant role. Seasonal and long-term variations in water quality have significant influences on the lower TLs clearly showing a bottom-up functioning of the ecosystem. Environmental degradations, such as siltation occurring in the lake, suggest possible risks in limiting ecosystem productivity.     

Nutrient cycling by fish supports relatively more primary production as lake productivity increases

Animals can be important in nutrient cycling in particular ecosystems, but few studies have examined how this importance varies along environmental gradients. In this study we quantified the nutrient cycling role of an abundant detritivorous fish species, the gizzard shad (Dorosoma cepedianum), in reservoir ecosystems along a gradient of ecosystem productivity. Gizzard shad feed mostly on sediment detritus and excrete sediment-derived nutrients into the water column, thereby mediating a cross-habitat translocation of nutrients to phytoplankton. We quantified nitrogen and phosphorus cycling (excretion) rates of gizzard shad, as well as nutrient demand by phytoplankton, in seven lakes over a four-year period (16 lake-years). The lakes span a gradient of watershed land use (the relative amounts of land used for agriculture vs. forest) and productivity. As the watersheds of these lakes became increasingly dominated by agricultural land, primary production rates, lake trophic state indicators (total phosphorus and chlorophyll concentrations), and nutrient flux through gizzard shad populations all increased. Nutrient cycling by gizzard shad supported a substantial proportion of primary production in these ecosystems, and this proportion increased as watershed agriculture (and ecosystem productivity) increased. In the four productive lakes with agricultural watersheds (>78% agricultural land), gizzard shad supported on average 51% of phytoplankton primary production (range 27-67%). In contrast, in the three relatively unproductive lakes in forested or mixed-land-use watersheds (>47% forest, <52% agricultural land), gizzard shad supported 18% of primary production (range 14-23%). Thus, along a gradient of forested to agricultural landscapes, both watershed nutrient inputs and nutrient translocation by gizzard shad increase, but our data indicate that the importance of nutrient translocation by gizzard shad increases more rapidly. Our results therefore support the hypothesis that watersheds and gizzard shad jointly regulate primary production in reservoir ecosystems.     

Hydrology and pollution of lake vistonis in Northern Greece

The basic objective of this study is the understanding of the hydrology and hydraulic characteristics of Lake Vistonis, a very shallow lake in Northern Greece, connected to the Aegean Sea through an irregular shallow channel. In this paper we estimate the pollutant residence time and the discharge of lake to the Aegean Sea as a function of lake elevation. Also we determine the seasonal variations of salinity of the lake.