Long Transients Near the Ghost of a Stable State in Eutrophic Shallow Lakes with Fluctuating Water Levels

Alternative stable states in shallow lakes have received much attention over the past decades, but less is known about transient dynamics of such lakes in the face of stochastic perturbations such as incidental extremes in water levels driven by climatic variability. Here, we report on the ecosystem dynamics of 70 lakes in the floodplains of the Lower Rhine in The Netherlands from 1999 to 2004. In any particular year, most lakes were either in a macrophyte-dominated clear state or in a contrasting state with turbid water and sparse submerged macrophyte cover. Macrophyte dominance was positively related to the occurrence of drawdown, and negatively to lake surface area and mean depth. We did not find a relation with nutrient levels. Remarkably, shifts between the two contrasting states were common, and episodes of low water levels appear to be an important external driver. A dry period before our study and the exceptionally dry summer of 2003 caused widespread drawdown of floodplain lakes, resulting in establishment of submerged macrophytes in the next year upon refill. In the 4 years without drawdown, many lakes returned to a macrophyte-poor turbid state. Although some lakes turned turbid again quickly, others took several years to shift into the turbid state. A model analysis suggests that such prolonged transient vegetated states may be explained by the fact that the system dynamics slow down in the vicinity of the “almost stable” macrophyte-dominated state. Such a “ghost” of an equilibrium causes the system to stick around that state relatively long before slipping into the only true stable state. Our results support the idea that transient dynamics rather than equilibrium may be the key to understanding the overall state of some ecosystems. A practical implication of our findings is that artificial stabilization of the water level in shallow lakes may have been an important factor aggravating the permanent loss of submerged macrophytes due to cultural eutrophication.     

Hydrology-Driven Regime Shifts in a Shallow Tropical Lake

Shifts between alternative stable states have become a focus of research in temperate shallow lakes. Here we show that sharp transitions between a clear, macrophyte-dominated state and a turbid state without submerged plants can also occur in tropical floodplain lakes, albeit driven by a largely different set of mechanisms. We show how a shallow lake in the Pantanal becomes covered by an exploding population of the submerged macrophyte Egeria najas Planchon as the water level rises during the annual high-water period. Water clarity increases spectacularly in this period due to flushing with river water that has lost most of its suspended matter during its slow flow over the flooded vegetated plains. A few months later when the water level drops again, the submerged plant beds die and decompose rapidly, triggering a phase of increasing turbidity. During this period an increase in dissolved organic matter, suspended matter, and phytoplankton biomass results in a sharp deterioration in water clarity. The concomitant water level decrease largely counteracts the effects on the underwater light climate, so that the amount of light at the bottom may not differ in comparison with the high-water period. Therefore, changes in light climate seem unlikely to be the sole driver of the vegetation shifts, and other mechanisms may prevent recovery of the submerged vegetation until the next high-water episode. Also, contrary to what is found in temperate lakes, there is no evidence for top-down control of phytoplankton biomass associated with the macrophyte-dominated state in our tropical lake. Author Contributions  Simoni Maria Loverde-Oliveira, Vera Lúcia Moraes Huszar—conceived the study, Simoni Maria Loverde-Oliveira—performed research and analyzed data, Simoni Maria Loverde-Oliveira, Vera Lúcia Moraes Huszar, Nestor Mazzeo, Marten Scheffer—wrote the paper.     

The restoration of Lake Apopka in relation to alternative stable states: an alternative view to that of Bachmann et al. (1999)

Bachmann et al. (1999) postulated that wind energy initiated, and has maintained, high turbidity in hypertrophic (mean chlorophyll a = 92 g l^–1) Lake Apopka, Florida (mean depth = 1.6 m; area = 12500 ha). They asserted that the turbid condition was initiated by a hurricane in late 1947 that destroyed submersed plant beds and that high turbidity has since been maintained by wind-driven resuspension of fluid sediments. In their view, there has been sufficient light for re-establishment of submersed plants over about 38% of the lake bottom, but plant growth has been precluded by the fluid character of the sediments. They concluded that the restoration program of the St. Johns River Water Management District, which includes reduction of the phosphorus (P) loading rate, will not restore water clarity or submersed vegetation. An alternative explanation for Lake Apopka's turbid state is that it was initiated, and has been maintained, by excessive P loading that led to algal blooms and elimination of submersed vegetation through light limitation. The transition to the turbid state was contemporaneous with drainage of 7300 ha of the floodplain wetland to create polders for farming, beginning in the early 1940s. Lake P budgets indicate that drainage of the farms caused a seven-fold increase in the P loading rate (0.08 g TP m^–2 yr^–1 to 0.55 g TP m^–2 yr^–1). Paleolimnological analysis of lake sediments also indicates an increase in the P loading rate in mid-century, concomitant with the decline in submersed vegetation and the increase in phytoplankton abundance. After the increase in P loading, wind disturbance may have accelerated the transition to the turbid state; but, before the increase in P loading, wind disturbance was insufficient to elicit the turbid state, as evidenced by the stability of the clear-water state in the face of 14 hurricanes and 41 tropical storms from 1881 to 1946. Measurements of photosynthetically active radiation (PAR) indicate that light limitation has inhibited submersed plant growth except on the shallowest 5% of the lake bottom. Further, the correlation between the diffuse attentuation coefficient (K _PAR) and chlorophyll a (CHLA) indicates that light limitation would be removed over about 82% of the lake bottom with a reduction in CHLA from 92 g l^–1 to 25 g l^–1. Recently, following a 40% reduction in the P loading rate, the mean total P (TP) concentration, mean CHLA, and total suspended solids fell by about 30% while mean Secchi depth increased by more than 20%. Submersed plant beds appeared in areas devoid of macrophytes for nearly 50 years. These improvements, during a period with no change in mean wind speeds measured at Lake Apopka, provide the strongest evidence that the turbid state has been maintained by excessive P loading and that the current restoration program, which combines P load reduction with planting and removal of planktivorous fish, will be effective.     

Long‐term dynamics of submerged macrophytes and algae in a small and shallow,eutrophic lake: implications for the stability of macrophyte‐dominance

1. Submerged macrophyte and phytoplankton components of eutrophic, shallow lakes have frequently undergone dynamic changes in composition and abundance with important consequences for lake functioning and stability. However, because of a paucity of long‐term survey data, we know little regarding the nature, direction and sequencing of such changes over decadal–centennial or longer timescales. 2. To circumvent this problem, we analysed multiple (n = 5) chronologically correlated sediment cores for plant macro‐remains and a single core for pollen and diatoms from one small, shallow, English lake (Felbrigg Hall Lake, Norfolk, U.K.), documenting 250 years of change to macrophyte and algal communities. 3. All five cores showed broadly similar stratigraphic changes in macrophyte remains with three distinct phases of macrophyte development: Myriophyllum–Chara–Potamogeton (c. pre‐1900), to Ceratophyllum–Chara–Potamogeton (c. 1900–1960) and finally to Zannichellia–Potamogeton (c. post‐1960). Macrophyte species richness declined from at least 10 species pre‐1900 to just four species at the present day. Additionally, in the final Zannichellia–Potamogeton phase, a directional shift between epi‐benthic and phytoplankton‐based primary production was indicated by the diatom data. 4. Based on macrophyte–seasonality relationships established for the region, concomitant with the final shift to Zannichellia–Potamogeton, we infer a reduction in the seasonal duration of plant dominance (plant‐covered period). Furthermore, we hypothesise that this change in species composition resulted in a situation whereby macrophyte populations were seasonally ‘sandwiched’ between two phytoplankton peaks in spring and late summer as observed in the contemporary lake. 5. We suggest that eutrophication‐induced reductions in macrophyte species richness, especially if the number of plant‐seasonal strategies is reduced, may constrict the plant growing season. In turn, this may render a shallow lake increasingly vulnerable to seasonal invasions of phytoplankton resulting in further species losses in the plant community. Thus, as part of a slow (over perhaps 10–100s of years) and self‐perpetuating process, macrophytes may be gradually pushed out by phytoplankton without the need for a perturbation as required in the alternative stable states model of plant loss.     

A Whole-Lake Experiment to Determine the Effects of Winter Droughts on Shallow Lakes

Lake-level fluctuations are common in the North American Great Plains region, where large-scale climate systems (El Niño, the Pacific Decadal Oscillation) and periodic droughts cause substantial hydrologic variability in both summer and winter. To date, most such research has focused on the effects of summer droughts on prairie lake ecosystems; therefore, we studied the impact of water-level decline during winter on ecosystem structure and function. Specifically, we hypothesized that lower lake levels during winter would increase anoxia, freezing and scouring of benthos, fish kills, herbivory by zooplankton, and nutrient release from sediments. In addition, we tested the hypothesis that winter droughts may initiate a switch between alternative stable states (turbid, clear). Physical, chemical, and biological variables were monitored from 1996 to 2001 in both Wascana Lake, which experienced a 50% decline in lake level, and Buffalo Pound Lake, where water levels were constant. A combination of before-after-control-impact (BACI) and multivariate analyses showed that drawdown resulted in elevated NH₄-N concentrations following reinundation; otherwise there were few detectable effects on lake water chemistry (PO₄-P, NO₃-N, total dissolved nitrogen, total dissolved carbon) or pelagic food web structure (phytoplankton, zooplankton), and the experimental lake remained in a macrophyte-rich state. There was, however, a 2.5-fold increase in macrophyte abundance and a shift from a community dominated by Ceratophyllum demersum before drawdown to one composed of Potamogeton pectinatus after manipulation. Overall, the lack of substantial dewatering effects suggests that lakes of the northern Great Plains may be resilient to severe winter conditions, possibly because of the recruitment of fish from regional metapopulations during summer. Further, our results indicate that lower water levels during winter likely promote the buffer mechanisms that reinforce a macrophyte-rich, clear-water state in shallow prairie lakes.     

Effects of fish and nutrient additions on food-web stability in a charophyte-dominated lake

1. The response of major food‐web constituents to combinations of nutrient addition and zooplanktivorous fish abundance was tested during two subsequent years in the shallow charophyte‐dominated lake Naardermeer in the Netherlands, using in situ enclosures. 2. Treatment effects differed sharply between study years. In 1998, when the summer temperature was low (17–21 °C), high algal biomass only developed at high nutrient levels in the presence of fish, but with no major effect on Chara biomass. In 1999, when the summer temperature was relatively high (20–25 °C), algal blooms occurred at high nutrient levels regardless of fish abundance, and were associated with a drastic decline in Chara biomass. 3. Differences between years in temperature and initial zooplankton composition and biomass were likely to contribute to the varying relative importance of top‐down and bottom‐up effects in these enclosure experiments. 4. The results suggest that when nutrient loads are increased towards levels where the macrophyte‐dominated state is being destabilised, a ‘switch’ is more likely to occur in warm summers.     

Effects of waterfowl, large fish and periphyton on the spring growth of Potamogeton pectinatus L. in Lake Mogan, Turkey

It has been argued that waterfowl and fish may threaten growth of submerged macrophytes, especially in spring during the early growth phase when plant biomass is low. A small reduction of biomass at that time might delay growth or decrease subsequent productivity. We investigated the impact of waterfowl and large fish on the spring growth of fennel pondweed (Potamogeton pectinatusL.) by employing an exclosure experiment in the macrophyte-dominated clear-water Lake Mogan, Turkey. Birds and large fish were excluded from eight plots and both in situvegetation and macrophytes kept in pots were compared to eight open plots. Also, to investigate the effect of periphyton on plant growth it was removed from half of the pot plants. Exclusion of waterfowl and fish may decrease predation on macroinvertebrates, which in turn may affect periphyton, and macrophyte growth, why macroinvertebrates also were sampled. Waterfowl density was high (15–70 ind. of coot, Fulica atraL. ha^–1), abundance of submerged plants was also high with a surface coverage of 70–80%, and benthivorous fish were present, mainly tench, (Tinca tincaL.) and carp, (Cyprinus carpioL.). Exclusion of waterfowl and large fish did not significantly affect the spring growth of pondweed; neither plants growing in situnor kept in pots. Removal of periphyton from the plants in the pots did not favour growth. The density of macroinvertebrates was not affected by the exclusion of waterfowl and large fish, but it was positively related to aboveground biomass of fennel pondweed. We suggest that even if waterfowl and large fish are in high densities, their effect on fennel pondweed spring growth in lakes with abundant submerged vegetation, such as Lake Mogan, is low.     

Resilience of Alternative Stable States during the Recovery of Shallow Lakes from Eutrophication: Lake Veluwe as a Case Study

In this paper we analyze a long-term dataset on the recovery from eutrophication of Lake Veluwe (The Netherlands). Clear hysteresis was observed in a number of ecosystem variables: the route to recovery differed significantly from the route that led to loss of clear water. The macrophyte dominated state disappeared in the late 1960s at TP above 0.20 mg l⁻¹, whereas its return occurred at less than 0.10 mg TP l⁻¹. Several regime shifts resulting in the occurrence of three alternative stable states were observed over a period of 30 years. The turbid state showed resistance to change, despite a strong and prompt reduction in Chl-a following reduction of external P-loading. The most important component that determined hysteresis in the return to clear water was not internal P-loading, but a high level of nonalgal light attenuation (through sediment resuspension) maintained by the interaction between wind and benthivorous fish. Although Chara was able to re-colonize the most shallow parts of the lake, recovery stalled and for a number of years clear (above charophyte beds) and turbid (deeper parts of the lake) water co-existed, as a separate alternative state on route to full recovery. Lake-wide clear water was re-established after bream density had been reduced substantially. This allowed a return of zebra mussels to the lake, whose high filtration capacity helped in maintaining clear water. In this study, we were able to identify the main drivers of hysteresis and regime shifts, although formal demonstration of cause and effect was not possible on the basis of field data alone. We argue that resilience of the present clear water state of Lake Veluwe very much depends on sizable populations of a few keystone species, especially Chara (stoneworts) and Dreissena (zebra mussels), and that careful management of these species is equally important as control of nutrients. Lake management should strive to maintain and strengthen resilience of the ecosystem, and this should offer protection against a renewed collapse of the clear state.     

A Conceptual Model of Natural and Anthropogenic Drivers and Their Influence on the Prince William Sound,Alaska, Ecosystem

Prince William Sound (PWS) is a semi-enclosed fjord estuary on the coast of Alaska adjoining the northern Gulf of Alaska (GOA). PWS is highly productive and diverse, with primary productivity strongly coupled to nutrient dynamics driven by variability in the climate and oceanography of the GOA and North Pacific Ocean. The pelagic and nearshore primary productivity supports a complex and diverse trophic structure, including large populations of forage and large fish that support many species of marine birds and mammals. High intra-annual, inter-annual, and interdecadal variability in climatic and oceanographic processes as drives high variability in the biological populations. A risk-based conceptual ecosystem model (CEM) is presented describing the natural processes, anthropogenic drivers, and resultant stressors that affect PWS, including stressors caused by the Great Alaska Earthquake of 1964 and the Exxon Valdez oil spill of 1989. A trophodynamic model incorporating PWS valued ecosystem components is integrated into the CEM. By representing the relative strengths of driver/stressors/effects, the CEM graphically demonstrates the fundamental dynamics of the PWS ecosystem, the natural forces that control the ecological condition of the Sound, and the relative contribution of natural processes and human activities to the health of the ecosystem. The CEM illustrates the dominance of natural processes in shaping the structure and functioning of the GOA and PWS ecosystems.