What evidence exists for alternative ecological regimes in salinising wetlands?

1. Land clearing in Australia's southwest has led to widespread salinisation of aquatic ecosystems. Four different ecological regimes (clear, submerged macrophyte‐dominated; clear, benthic microbial community‐dominated; turbid, phytoplankton‐dominated; and turbid, sediment‐dominated) have previously been identified in the salinising wetlands of this region. 2. Monitoring data from seven saline wetlands over an 18‐month period were used to evaluate whether a continuum, simple threshold or alternative regimes conceptual model (sensu Hydrobiologia, 200/201, 1990, 367; Hydrobiologia, 200/201, 1990, 475) most appropriately represented transitions between these ecological regimes. We also aimed to identify whether factors other than salinity played a major role in defining ecological regimes or causing shifts between them. 3. Ordination of biological variables revealed a separation of benthic microbial community‐dominated from submerged macrophyte‐dominated sites and times. The mean salinities of these two groups were very similar, suggesting that a salinity threshold was not responsible for benthic microbial versus macrophyte dominance. No other environmental variable was found to have a strong, direct influence on the groupings. 4. The dynamics of regime change in saline wetlands appear not to be driven by any single variable, but by the combined effects of salinity and water regime on species life histories and competitive abilities. Macrophytes were powerful competitors, able to germinate and establish under a range of salinities, turbidities and water depths, and were favoured by seasonal drying. 5. Data from the seven wetlands indicated that the continuum, simple threshold and alternative regimes conceptual models did not appropriately represent transitions between ecological regimes in seasonally drying wetlands. Macrophyte and benthic microbial regimes occurred at overlapping salinity levels, excluding both the continuum and threshold models, and the regular occurrence of drying appeared to preclude the alternative regimes model. Drying prevented the development of strong positive feedback mechanisms, which might otherwise have maintained the benthic microbial community‐dominated regime. We hypothesise that an alternative regimes model might still be valid for salinising ecosystems holding permanent water.     

Migratory benthic fishes may induce regime shifts in a tropical floodplain pond

1. Alternative states are a widely recorded phenomenon in shallow lakes, which may shift between turbid‐ and clear‐water conditions. Here, we investigate whether such shifts in a tropical floodplain pond may be related to the effect of the flood pulse regime on the community structures of fish and macrophytes. 2. Using a long‐term data set, we demonstrate how benthic fish migration together with colonisation by submerged plants affected the transition from a turbid to a macrophyte‐dominated state in a floodplain pond without top‐down control. 3. In our study, the turbid state occurred mostly during low water phases and was largely characterised by high values for the biomass of benthic fish, chlorophyll‐a and total phosphorous. 4. During the period of rising water levels, the migration of benthic fish out of the pond occurs simultaneously with the establishment of submerged plants, while water turbidity decreases along with phytoplankton and nutrient concentrations, inducing a clear‐water phase. However, when submerged plants are absent and fish migration is low, a transient state is generated. 5. We suggest that, in contrast to temperate ponds and shallow lakes, where the main driving mechanisms establishing alternative states are related to cascading effects via the food chain, in tropical ponds and shallow lakes it is resuspension of sediments by benthic fish that plays the most significant role in establishing alternative states. However, the effect of the flood pulse regime plays an important role in the temporal dynamics of fish community structure by controlling benthic fish migration.     

Can different vegetative states in shallow coastal bays of the Baltic Sea be linked to internal nutrient levels and external nutrient load?

Hydrobiologia - Two different vegetative states, i.e. one clear water state dominated by benthic macrophytes and one turbid state dominated by phytoplankton, are commonly found in shallow lakes. In...     

Contrasting impacts of invasive engineers on freshwater ecosystems: an experiment and meta-analysis

Invasion by common carp (Cyprinus carpio) and red swamp crayfish (Procambarus clarkii) in shallow lakes have been followed by stable-state changes from a macrophyte-dominated clear water state to a phytoplankton-dominated turbid water state. Both invasive carp and crayfish are, therefore, possible drivers for catastrophic regime shifts. Despite these two species having been introduced into ecosystems world-wide, their relative significance on regime shifts remains largely unexplored. We compared the ecological impacts of carp and crayfish on submerged macrophytes, water quality, phytoplankton, nutrient dynamics, zooplankton and benthic macroinvertebrates by combining an enclosure experiment and a meta-analysis. The experiment was designed to examine how water quality and biological variables responded to increasing carp or crayfish biomass. We found that even at a low biomass, carp had large and positive impacts on suspended solids, phytoplankton and nutrients and negative impacts on benthic macroinvertebrates. In contrast, crayfish had a strong negative impact on submerged macrophytes. The impacts of crayfish on macrophytes were significantly greater than those of carp. The meta-analysis showed that both carp and crayfish have significant effects on submerged macrophytes, phytoplankton, nutrient dynamics and benthic macroinvertebrates, while zooplankton are affected by carp but not crayfish. It also indicated that crayfish have significantly greater impacts on macrophytes relative to carp. Overall, the meta-analysis largely supported the results of the experiment. Taken as a whole, our results show that both carp and crayfish have profound effects on community composition and ecosystem processes through combined consequences of bioturbation, excretion, consumption and non-consumptive destruction. However, key variables (e.g. macrophytes) relating to stable-state changes responded differently to increasing carp or crayfish biomass, indicating that they have differential ecosystem impacts.     

Thresholds and Stability of Alternative Regimes in Shallow Prairie–Parkland Lakes of Central North America

Numerous studies have demonstrated alternative regimes in shallow lake ecosystems around the world, with one state dominated by submerged macrophytes and the other by phytoplankton. However, the stability of each regime, and thresholds at which lakes shift to the alternative regime, are poorly known. We used a cross-sectional analysis of 72 shallow lakes located in prairie and parkland areas of Minnesota, USA, during 2005 and 2006 to assess the occurrence of alternative regimes and shifts between them. Cluster analysis revealed two distinct groups of lakes characterized not only by different macrophyte abundance and chlorophyll a levels but also by different total phosphorus–chlorophyll a relationships. Thirty-nine lakes were macrophyte- and 23 lakes phytoplankton-dominated in both years, whereas 10 sites shifted sharply between those regimes. We failed to detect a universal shifting threshold in terms of chlorophyll a or total phosphorus. However, 95% of the lakes with chlorophyll a concentrations less than 22 μg l⁻¹ were in a clear-water regime, whereas 95% of the lakes with chlorophyll a higher than 31 μg l⁻¹ were in a turbid regime. Total phosphorus less than 62 μg l⁻¹ was an accurate predictor of lakes in a stable clear-water regime, whereas a large change in biomass of planktivores and benthivores between years was the only variable weakly related to regime shifts. Our results support the theoretical prediction that regime thresholds vary among lakes. We recommend that lake managers focus on improving resilience of clear regimes in shallow lakes by reducing nutrient loading, rather than attempting to identify and manage complex triggers of regime shifts. Author contributions KDZ, MAH, BRH, and MLK all contributed to the design of the study, performed the research, analyzed data, and helped write the article.     

Community structure and diel migration of zooplankton in shallow brackish lakes: role of salinity and predators

Diel horizontal migration (DHM), where zooplankton moves towards macrophytes during daytime to avoid planktivorous fish, has been reported as a common migration pattern of zooplankton in shallow temperate freshwater lakes. However, in shallow eutrophic brackish lakes, macrophytes seem not to have the same refuge effect, as these lakes may remain turbid even at relatively high macrophyte abundances. To investigate the extent to which macrophytes serve as a refuge for zooplankton at different salinities, we introduced artificial plants mimicking submerged macrophytes in the littoral zone of four shallow lakes, with salinities ranging from almost freshwater (0.3) to oligohaline waters (3.8). Furthermore, we examined the effects of different salinities on the community structure. Diel samples of zooplankton were taken from artificial plants, from areas where macrophytes had been removed (intermediate areas) and, in two of the lakes, also in open water. Fish and macroinvertebrates were sampled amongst the artificial plants and in intermediate areas to investigate their influence on zooplankton migration. Our results indicated that diel vertical migration (DVM) was the most frequent migration pattern of zooplankton groups, suggesting that submerged macrophytes were a poor refuge against predation at all salinities under study. Presumably, this pattern was the result of the relatively high densities of small planktivorous fish and macroinvertebrate predators within the submerged plants. In addition, we found major differences in the composition of zooplankton, fish and macroinvertebrate communities at the different salinities and species richness and diversity of zooplankton decreased with increasing salinity. At low salinities both planktonic/free-swimming and benthic/plant-associated cladocerans occurred, whilst only benthic ones occurred at the highest salinity. The low zooplankton biomass and overall smaller-bodied zooplankton specimens may result in a lower grazing capacity on phytoplankton, and enhance the turbid state in nutrient rich shallow brackish lakes.     

Relative importance of periphyton and phytoplankton in turbid and clear vegetated shallow lakes from the Pampa Plain (Argentina): a comparative experimental study

We analyzed experimentally the relative contribution of phytoplankton and periphyton in two shallow lakes from the Pampa Plain (Argentina) that represent opposite scenarios according to the alternative states hypothesis for shallow lakes: a clear lake with submerged macrophytes, and a turbid lake with high phytoplankton biomass. To study the temporal changes of both microalgal communities under such contrasting conditions, we placed enclosures in the littoral zone of each lake, including natural phytoplankton and artificial substrata, half previously colonized by periphyton until a mature stage and half clean to analyze periphyton colonization. In the clear vegetated shallow lake, periphyton chlorophyll a concentrations were 3–6 times higher than those of the phytoplankton community. In contrast, phytoplankton chlorophyll a concentrations were 76–1,325 times higher than those of periphyton in the turbid lake. Here, under light limitation conditions, the colonization of the periphyton was significantly lower than in the clear lake. Our results indicate that in turbid shallow lakes, the light limitation caused by phytoplankton determines a low periphyton biomass dominated by heterotrophic components. In clear vegetated shallow lakes, where nitrogen limitation probably occurs, periphyton may develop higher biomass, most likely due to their higher efficiency in nutrient recycling.     

Effects of elevated turbidity on shallow lake fish communities

Synopsis We compared the fish communities of two shallow lakes in the lower Waikato River basin, North Island, New Zealand, to determine the effects of elevated suspended solids (SS) and collapse of submerged macrophytes. Lake Waahi was turbid (20–40 g m^-3 SS) and devoid of submerged macrophytes whereas Lake Whangape was clearer (5 g m^-3 SS) and dominated by submerged macrophytes. The lakes had similar fish species richness and had nine major species in common; representing eight families including Anguillidae, Retropinnidae, Galaxiidae, Eleotridae, Mugilidae, Ictaluridae, Poeciliidae, and Cyprinidae (two species). The only major fish that was absent from Lake Waahi was a lacustrine form of the common smelt, Retropinna retropinna, which disappeared after the lake became turbid in the late 1970s. CPUE, condition, and size of most species in Lake Waahi were similar to, or greater than, those in Lake Whangape. Lake Whangape clearly exceeded Lake Waahi only for CPUE of two species. Within Lake Whangape two species displayed significantly greater condition, and one species greater size, in a turbid arm of the lake than in the main basin. Apart from lacustrine Retropinna retropinna, the fish in these lakes appear well adapted to cope with, or to avoid, the direct toxic effects of suspended and settleable solids on sensitive early developmental stages. In Lake Waahi loss of cover and food provided by submerged macrophytes appears to have been compensated for by increased turbidity and an associated increase in the biomass of the mysid, Tenagomysis chiltoni (a major prey item).     

Precipitation-Induced Alternative Regime Switches in Shallow Lakes of the Boreal Plains (Alberta,Canada)

This study focused on unraveling the natural mechanism for the frequent shifts in alternative regimes in pristine shallow lakes of the Boreal Plains, Alberta, Canada. The lakes tend to be clear and dominated by submerged aquatic vegetation (SAV) or turbid and dominated by phytoplankton. We report on the inter-annual response of 23 lakes from 2001 to 2007. We explore the effect of fluctuations in annual precipitation on the lake response including water depth, total phosphorus (TP) concentration, turbidity, phytoplankton biomass, SAV biomass, and the proportion of clear and turbid lakes. The regime switches appear driven by the transient dynamics of phytoplankton, and dilution of nutrients, phytoplankton biomass, and turbidity during wet years, and evapoconcentration during dry years. Increased precipitation was correlated with decreased phytoplankton biomass, TP concentration, chloride concentration, and turbidity. In 2005, the wettest year, no phytoplankton-dominated lakes were observed. During the driest year (2002), the phytoplankton-dominant regime (>18 μg chl-a L^?1) occurred in 22% of lakes, which was higher than the study period average. SAV biomass was not directly affected by precipitation, but was negatively associated with phytoplankton biomass and positively associated with the previous year’s SAV growth. SAV biomass was carried over from year-to-year, and the occurrence of SAV-dominated (>25% cover) lakes was significantly higher in 2007 (90%) following 3 years of high precipitation levels.     

Phytoplankton and Epipelon Responses to Clear and Turbid Phases in a Seepage Lake (Buenos Aires,Argentina)

Annual changes in the algal density and concentrations of chlorophyll a, total phosphorus, and organic matter were analyzed in water and sediments at four sites characterized by the presence or absence of submerged and emergent macrophytes, during turbid‐ and clear‐water conditions to determine the contribution of the algal components of the plankton and the epipelon and to identify the most typical species in each community. Three states were recognized: one turbid and two clear, with different submerged macrophyte cover. The peaks of phytoplankton and epipelon occurred in the turbid phase, whereas the highest proportion of true epipelic algae in sediments was reached in the second clear phase. The Oscillatoriaceae dominated during the turbid phase in the water and throughout the entire year within the sediments. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Plant community structure determines primary productivity in shallow,eutrophic lakes

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Ecological regime shifts in salinised wetland systems. II. Factors affecting the dominance of benthic microbial communities

This paper is the second in a pair investigating potential mechanisms for ecological regime change in salinising wetlands. The first paper in this series focused on the responses of the salt-tolerant submerged macrophyte community to salinity. In this second paper, we investigated some of the environmental conditions required for initiation and dominance of benthic microbial communities using a combination of experimental and observational data. Two experiments were carried out. One investigated the importance of prior establishment of benthic microbial communities on their ability to maintain prevalence over macrophyte colonisation (‘persistence’ experiment), while the other investigated hydrology and its effect on sediment perturbation, potential nutrient release and subsequent benthic microbial community establishment (‘flooding’ experiment). The ‘persistence’ experiment measured the biomass of benthic microbial communities and emergence of macrophytes from sediments kept either wet or dry for 4 weeks then flooded at a range of salinities. Benthic microbial biomass was similar across all of the salinities tested (15, 45 and 70 ppt), with a slight increase at higher salinities, suggesting that none of these limited benthic microbial community development. Pre-wetting of sediments usually increased benthic microbial community biomass and reduced macrophyte germination, but the latter was attributed to the presence of anoxic sediments rather than the increased benthic microbial community biomass. Germinating macrophytes emerged through benthic microbial communities or dense heterotrophic bacterial blooms, demonstrating that they could become dominant even when another community was already established. Field data supported these results, suggesting that the development of benthic microbial communities is not limited by salinity alone, but includes other factors, such as the water regime. In the ‘flooding’ experiment, the largest differences in nutrient concentrations ultimately lay between the pre-wet and pre-dry treatments (due to the greater release of nutrients and development of anoxia in the latter) rather than those subjected to fast versus slow flooding. In response to this, highest benthic microbial community biomass was in treatments with pre-wet sediment, corresponding with lower phytoplankton biomass.     

Impact of introduced carp (Cyprinus carpio)in subtropical shallow ponds in Central Mexico

In Mexico, as in many other subtropical and tropical countries, there has been a recent trend towards stocking non-native carp (Cyprinus carpio) in lakes and ponds as a source of food in rural areas. However, the results of a study in a series of small(1–8 ha.), shallow (<2 m), semi-natural ponds in Acambay, a high altitude valley in the basin of the Lerma river in the volcanic belt in central west Mexico, illustrate that the stocking of carp over a threshold value may have a detrimental ecological impact at several trophic levels. Ponds with carp tended to be turbid with high levels of suspended solids, and with few rooted macrophytes and epibenthic invertebrates. In contrast, ponds without carp had clear water and abundant rooted macrophytes and associated invertebrates, particularly gastropod molluscs. The direct uprooting of macrophytes by benthic foraging carp appeared to be the most important mechanism in switching the ponds from a clear macrophyte-dominated to a turbid state. The subtropical study ponds thus appear to confirm the alternative stable-state hypothesis developed intemperate lakes, although the importance of benthic rather than pelagic interactions was emphasised. The implications of stocking carp for native fauna of high intrinsic conservation value and as a food supply for local people are outlined. This revised version was published online in July 2006 with corrections to the Cover Date.     

Periphyton-macroinvertebrate interactions in light and fish manipulated enclosures in a clear and a turbid shallow lake

In a clear and a turbid freshwater lake the biomasses of phytoplankton, periphytic algae and periphytonassociated macrograzers were followed in enclosures with and without fish (Rutilus rutilus) and four light levels (100%, 55%, 7% and < 1% of incoming light), respectively. Fish and light affected the biomass of primary producers and the benthic grazers in both lakes. The biomass of primary producers was generally higher in the turbid than the clear lake, and in both lakes fish positively affected the biomass, while shading reduced it. Total biomass of benthic grazing invertebrates was higher in the clear than in the turbid lake and the lakes were dominated by snails and chironomids + ostracods, respectively. While light had no effect on the biomass of grazers in the clear lake, snail breeding was delayed in the most shaded enclosures and presence of fish reduced the number of snails and the total biomass of grazers. In the turbid lake ostracod abundance was not influenced by light, but was higher in fish-free enclosures. Density of chironomids correlated positively with periphyton biomass in summer, while fish had no effect. Generally, light-mediated regulation of primary producers was stronger in the turbid than in the clear lake, but the regulation did not nambiguously influence the primary consumers. However, regulation by fish of the benthic grazer community was stronger in the clear than in the turbid lake, and in both lakes strong top-down effects on periphyton were seen. The results indicate that if present-day climate in Denmark in the future is found in coastal areas at higher latitudes, the effect of lower light during winter in such areas will be highest in clear lakes, with typically lower fish biomass and higher invertebrate grazer density.     

Abrupt regime shifts in space and time along rivers and connected lake systems

Regime shifts between clear and turbid water states are commonly found in shallow lakes. These shifts are attributed to a positive feedback between water clarity and submerged macrophytes (underwater plants). Altering the retention time of the water may influence these interactions and thus potentially reduce the probability of alternative stable states. Here we assessed the effect of water retention time on the occurrence of alternative states in water quality of flushed lakes, chains of lakes and rivers using a spatially explicit simple model. Our results indicate that increased flushing of lakes rapidly decreases the range of parameters with alternative stable states up to their total disappearance at a flushing rate of about 50% the algal growth rate. Similarly, in a chain of lakes or in rivers with low flowing velocity, our model predicts that alternative stable states can only occur for systems with a high retention time. Despite the lack of hysteresis at lower water retention times, we predict that abrupt changes between clear and turbid states are still possible both in time and in space. Over a wide range of parameters, the equilibrium state of the chain of lakes shows a steep gradient of vegetation cover. Further, the transient dynamics of the model often include rapid shifts in time. For example, a local regime shift that occurs upstream may propagate through the whole lake chain or river due to a domino effect. All results of the simple model could qualitatively be reproduced with a more mechanistic model. The abrupt rather than gradual response of submerged macrophytes to reduced turbidity levels still makes river systems rather resilient to management measure. The importance of the initial turbidity and the observed domino effect suggest that restoration measures should start upstream and that these measures should eventually trigger regime shifts downstream.     

Long-term pattern of alternative stable states in two shallow eutrophic lakes

• 1 Lake Tåkern and Lake Krankesjön, two moderately eutrophic, shallow lakes in southern Sweden, have during the past few decades shifted several times between a clear-water state with abundant submerged vegetation and a turbid state with high phytoplankton densities.
• 2 Between 1985 and 1991, Lake Takern was in a clear state, whereas Lake Krankesjon shifted from a turbid to a clear state. During this shift, the area covered by submerged macrophytes expanded, followed by an increase in water transparency, plant-associated macroinvertebrates, and piscivorous fish. Nutrient concentrations, phytoplankton biomass and abundance of planktonic cladocerans decreased.
• 3 In both lakes, water level fluctuations were the most common factor causing shifts, affecting submerged macrophytes either through changes in light availability or through catastrophic events such as dry-out or mechanical damage by ice movement.
• 4 Our data give further support for the existence of two alternative stable states in shallow lakes maintained by self-stabilizing feedback mechanisms.

     

Fish manipulation as a lake restoration tool in shallow,eutrophic, temperate lakes 2: threshold levels,long-term stability and conclusions

In order to evaluate short-term and long-term effects of fish manipulation in shallow, eutrophic lakes, empirical studies on relationships between lake water concentration of total phosphorus (P) and the occurrence of phytoplankton, submerged macrophytes and fish in Danish lakes are combined with results from three whole-lake fish manipulation experiments. After removal of less than 80 per cent of the planktivorous fish stock a short-term trophic cascade was obtained in the nutrient regimes, where large cyanobacteria were not strongly dominant and persistent. In shallow Danish lakes cyanobacteria were the most often dominating phytoplankton class in the P-range between 200 and 1 000μg P l⁻¹. Long-term effects are suggested to be closely related to the ability of the lake to establish a permanent and wide distribution of submerged macrophytes and to create self-perpetuating increases in the ratio of piscivorous to planktivorous fish. The maximum depth at which submerged macrophytes occurred, decreased exponentially with increasing P concentration. Submerged macrophytes were absent in lakes>10 ha and with P levels above 250–300μg P l⁻¹, but still abundant in some lakes<3 ha at 650μg P l⁻¹. Lakes with high cover of submerged macrophytes showed higher transparencies than lakes with low cover aboveca. 50μg P l⁻¹. These results support the alternative stable state hypothesis (clear or turbid water stages). Planktivorous fish>10 cm numerically contributed more than 80 per cent of the total planktivorous and piscivorous fish (>10 cm) in the pelagical of lakes with concentrations above 100μg P l⁻¹. Below this threshold level the proportion of planktivores decreased markedly toca. 50 per cent at 22μg P l⁻¹. The extent of the shift in depth colonization of submerged macrophytes and fish stock composition in the three whole-lake fish manipulations follows closely the predictions from the relationships derived from the empirical study. We conclude that a long-term effect of a reduction in the density of planktivorous fish can be expected only when the external phosphorus loading is reduced to below 0.5–2.0 g m⁻² y⁻¹. This loading is equivalent to an in-lake summer concentration below 80–150μg P l⁻¹. Furthermore, fish manipulation as a restoration tool seems most efficient in shallow lakes.     

Biodiversity dynamics of freshwater wetland ecosystems affected by secondary salinisation and seasonal hydrology variation: a model-based study

Freshwater wetlands worldwide are under threat from secondary salinisation and climate change. Given that many freshwater wetlands naturally have highly variable hydrology, it is important to understand the combined effects of salinity and water regime on wetland biodiversity. Here a mathematical model has been developed to explore the biodiversity dynamics of freshwater wetland ecosystems affected by secondary salinisation and seasonal hydrology variation. The model shows that seasonal hydrological change can drive the wetland ecosystem into a stable oscillatory state of biodiversity, with the same period as the wetting and drying cycle. The initial condition of a wetland mediates the ecological response of the wetland ecosystem to salinity and seasonal variability. There are two manifestations of stability that occur in relation to wetland biodiversity: monostability and bistability. In model simulations, some wetland ecosystems may respond to the effects of seasonal change quickly, while others may do so more slowly. In ‘slow response’ wetlands, seasonal variability has a weak impact on the ecosystem properties of stability, resilience, sensitivity and the species richness–mean salinity relationship. In contrast, ‘fast response’ wetlands are seasonally controlled heavily. Seasonal variability can play a critical role in determining ecosystem properties. Changes in the strength of seasonality can induce the transition between monostability and bistability. Seasonal variability may also reduce wetland resilience, exacerbating the risk of secondary salinisation. On the other hand, seasonal variability may provide opportunities for the restoration of salinised wetlands by increasing their sensitivity to management actions and facilitating recovery processes. Model simulations show that the response of the stable biodiversity oscillation to changing mean salinity is dependent on seasonality strength (primarily for fast response wetlands) and other wetland conditions. Generally, there are two types of wetland responses to changes in mean salinity: type 1 wetlands exhibit a graded response of species richness (a surrogate for biodiversity), whereas a hysteretic response occurs in type 2 wetlands. Species diversity displays critical behaviour: regime shifts in diversity occur at the thresholds of mean salinity, strength of seasonality or initial species diversity. The predictions are consistent with previously-published field observations in salinised freshwater wetlands. Handling editor: D. Hamilton     

Bimodal transparency as an indicator for alternative states in South American lakes

1. The alternative state theory claims that shallow lakes may have either clear water, and be dominated by submerged macrophytes, or turbid water and be dominated by phytoplankton. Most evidence for this theory comes from studies in temperate or boreal regions of Europe. Because of differences in the strength of trophic interactions, such as in the pressure of zooplankton grazing on phytoplankton, this influential theory might not apply elsewhere. 2. Here, we test the theory for South American lakes, combining field data and Landsat satellite data. We studied the frequency distribution of primary producers and water transparency, looking for potential bimodality separating clear and turbid lakes. A bimodal distribution might be observed if there are indeed alternative states, although would not itself be sufficient evidence for the theory. Possible shifts between alternative states were analysed by comparing satellite data from 1987 to 2005. 3. In our field data, there was a bimodal pattern in phytoplankton abundance and possibly in the abundance of submerged macrophytes, but not in water transparency. Analyses of the larger satellite data set revealed bimodality in lake transparency in 2005, but less so in 1987. In 1987, the lakes were generally clearer, and the transition to higher turbidity was more gradual than in 2005. The stronger bimodality in the more recent data, and the overall lower transparency, could have been caused by an increase in fertiliser use and subsequent eutrophication but also by differences in hydrology. Further, 1987 was much wetter than 2005, which could have caused dilution of suspended particles, leading to clearer water. 4. While a bimodal distribution in the abundance of primary producers and water clarity is not decisive evidence for or against the theory of alternative states, our data clearly fail to refute it.     

Loss of Submerged Macrophytes in Shallow Lakes in North‐Eastern Germany

During the 1950s, the submerged vegetation of shallow lakes in north‐eastern Germany was dominated by nutrient tolerant species, with Ceratophyllum demersum and Myriophyllum sp. being most common. Almost one third of 300 investigated lakes had already lost their submerged macrophytes at that time. Very shallow lakes showed either high or low macrophyte abundance. Increasing depth resulted in medium macrophyte abundances, which may contribute to the stabilisation of local or temporary clearwater states. Forty years later, the percentage of lakes without macrophytes had dramatically increased. Between 55 and 85% of the investigated lakes showed a low abundance. The decline was most pronounced in very shallow lakes. The majority of the investigated lakes showed summer TP concentrations below 100 μg L^–1, but no colonisation by submerged macrophytes, which indicates a resilience against re‐colonisation.