Resistance to drought affects persistence of alternative regimes in shallow lakes of the Boreal Plains (Alberta,Canada)

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How important is the crustacean plankton for the maintenance of water clarity in shallow lakes with abundant submerged vegetation?

• 1 We measured the abundance and biomass of filter‐feeding microcrustacean zooplankton and calculated their grazing impact on phytoplankton biomass during summer in five shallow, mesotrophic to eutrophic lakes. For three of the lakes data exist both from years with dense submerged vegetation and low turbidity (the clearwater state), as well as from years characterised by sparse vegetation and high turbidity (the turbid state). In the other two lakes data are available only for clearwater conditions.
• 2 In all lakes conditions of dense vegetation and clear water coincided with a low abundance of crustacean plankton during summer. In the three lakes that shifted, the calculated biovolume ingested by crustacean plankton (filtering rate) was 3–11 times lower during clearwater conditions compared with turbid conditions. Because phytoplankton biomass was lower during clearwater conditions, however, daily grazing pressure from microcrustacea (expressed as percentage of phytoplankton biomass) did not differ between states. In three of the five lakes, grazers were estimated to take less than 10% of the phytoplankton biomass per day, indicating filtration by zooplankton was not the most important mechanism to maintain clearwater conditions.
• 3 High densities of Cladocera were found in three of the lakes within dense stands of Charophyta. However, these samples were dominated by plant‐associated taxa that even during the night were rarely found outside the vegetation. This indicates that plant‐associated zooplankton has no major influence on the maintenance of water clarity outside the vegetation.
• 4 Spring peak abundance of Cladocera was observed in three of the lakes. In two of these, where seasonal development was studied in both the clearwater and the turbid state, spring peaks were lower during the clearwater state.
• 5 Predation, low food availability or a combination of both may explain the low zooplankton densities. Phytoplankton may be limited by low phosphorus availability in the lakes dominated by Charophyta. Our results indicate that the importance of zooplankton grazing may have minor importance for the maintenance of the clearwater state in lakes with dense, well‐established submerged vegetation.

     

Total phosphorus thresholds for regime shifts are nearly equal in subtropical and temperate shallow lakes with moderate depths and areas

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Long-term trends and regional differences of phytoplankton in large Finnish lakes

Major nutrients (N and P) and phytoplankton from 19 large lakes from southern (61°) to northern (69°) Finland were analyzed to detect long-term trends and regional differences. The data sets from June, July and August cover the period from the early 1980s to the present. Altogether >700 phytoplankton and >4000 N, P and Chl a results were used for the study. In 40% of the lakes, the total phosphorus (TP) concentration decreased significantly and in >25% of the lakes a significant reduction was found in the total nitrogen (TN) concentration. At the same time, the phytoplankton biomass declined only in 15% of the lakes and the long-term trends in chlorophyll a more often increased than decreased. A clear gradient from southern to northern Finland and western to eastern Finland was found in the phytoplankton biomass. During the study period, the biomasses of cyanobacteria and centrales (diatoms) decreased whilst there was an increase in the biomass of pennales (diatoms) in one-third of the lakes. The proportion of chlorophytes in the total biomass also increased in >20% of the study lakes. In southern and western Finland, the total biomass and the contribution of cyanobacteria were higher. Centrales made a higher contribution to the total biomass in the north. Pennales and chlorophytes were less abundant and chrysophytes more abundant in the east. Differences in the community composition reflected the gradients in the total nutrients, and particularly in TP. The observations support the assumed role of phosphorus as the key limiting nutrient in large Finnish lakes irrespective of lake′s location. The N:P ratio proved to be a poor predictor of cyanobacteria occurrence in the study 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.     

Ontogenetic changes in the predator–prey interactions between threadsail filefish and moon jellyfish

The global climate change may lead to more extreme climate events such as severe flooding creating excessive pulse-loading of nutrients, including nitrogen (N), to freshwaters. We conducted a 3-month mesocosm study to investigate the responses of phytoplankton, zooplankton and Vallisneria spinulosa to different N loading patterns using weekly and monthly additions of in total 14 g N m^?2 month^?1 during the first 2 months. The monthly additions led to higher phytoplankton chlorophyll a and total phytoplankton biomass than at ambient conditions as well as lower leaf biomass and a smaller ramet number of V. spinulosa. Moreover, the biomass of cyanobacteria was higher during summer (August) in the monthly treatments than those with weekly or no additions. However, the biomass of plankton and macrophytes did not differ among the N treatments at the end of the experiment, 1 month after the termination of N addition. We conclude that by stimulating the growth of phytoplankton (cyanobacteria) and reducing the growth of submerged macrophytes, short-term extreme N loading may have significant effects on shallow nutrient-rich lakes and that the lakes may show fast recovery if they are not close to the threshold of a regime shift from a clear to a turbid state.     

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.     

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.     

Ambiguous climate impacts on competition between submerged macrophytes and phytoplankton in shallow lakes

1. Shallow lakes may switch from a state dominated by submerged macrophytes to a phytoplankton‐dominated state when a critical nutrient concentration is exceeded. We explore how climate change may affect this critical nutrient concentration by linking a graphical model to data from 83 lakes along a large climate gradient in South America. 2. The data indicate that in warmer climates, submerged macrophytes may tolerate more underwater shade than in cooler lakes. By contrast, the relationship between phytoplankton biomass [approximated by chlorophyll‐a (chl‐a) or biovolume] and nutrient concentrations did not change consistently along the climate gradient. In warmer climates, the correlation between phytoplankton biomass and nutrient concentrations was overall weak, especially at low total phosphorus (TP) concentrations where the chl‐a/ TP ratio could be either low or high. 3. Although the enhanced shade tolerance of submerged plants in warmer lakes might promote the stability of their dominance, the potentially high phytoplankton biomass at low nutrient concentrations suggests an overall low predictability of climate effects. 4. We found that near‐bottom oxygen concentrations are lower in warm lakes than in cooler lakes, implying that anoxic P release from eutrophic sediment in warm lakes likely causes higher TP concentrations in the water column. Subsequently, this may lead to a higher phytoplankton biomass in warmer lakes than in cooler lakes with similar external nutrient loadings. 5. Our results indicate that climate effects on the competitive balance between submerged macrophytes and phytoplankton are not straightforward.     

Phytoplankton periodicity of the Waitaki Lakes,New Zealand

The Waitaki River system in the South Island of New Zealand includes three large glacially-formed headwater lakes, Tekapo, Pukaki and Ohau, which drain into the manmade Lake Benmore. Phytoplankton periodicity was followed from December 1975 to January 1980 as part of a study investigating possible changes in these lakes as a consequence of hydroelectric development. The phytoplankton was highly dominated by diatoms, e.g., Diatoma elongatum, Cyclotella stelligera, Asterionella formosa, and Synedra acus, but in lakes Ohau and Benmore populations of green algae occasionally developed. In all four lakes seasonal phytoplankton periodicity was observed with maximum biomass in spring and summer. In Lake Tekapo, the first lake in the chain, maximum biomass did not exceed 300 mg m^–3, but in the very turbid Lake Pukaki the maximum summer biomass ranged between 300 and 800 mg m^–3. In Lake Ohau, the least turbid lake, maximum biomass was around 1 000 mg m^–3. In the newly created Lake Benmore periodicity was less evident and summer maxima reached over 1 500 mg m^–3. The phytoplankton periodicity in these lakes is greatly influenced by seasonal patterns of turbidity from inflowing glacial silt.     

A comparison of zooplankton densities and biomass in Lakes Peipsi and Võrtsjärv (Estonia): rotifers and crustaceans versus ciliates

The abundance and biomass of ciliates, rotifers, cladocerans and copepods were studied in Lake Peipsi and Lake Võrtsjärv, both of which are shallow, turbid and large. Our hypothesis was that in a large shallow eutrophic lake, the ciliates could be the most important zooplankton group. The mean metazooplankton biomass was higher in Peipsi than in Võrtsjärv (mean values and SD, 1.8 ± 0.7 and 1.3 ± 0.6 mg WM l^?1). In Peipsi, the metazooplankton biomass was dominated by filtrators that feed on large-sized phytoplankton and are characteristic of oligo-mesotrophic waters. In Võrtsjärv, the metazooplankton was dominated by species characteristic of eutrophic waters. The planktonic ciliates in both lakes were dominated by oligotrichs. The biomass of ciliates was much greater in Võrtsjärv (mean 2.3 ± 1.4 mg WM l^?1) than in Peipsi (0.1 ± 0.08 mg WM l^?1). Ciliates formed about 60% of the total zooplankton biomass in Võrtsjärv but only 6% in Peipsi. Thus, the food chains in the two lakes differ: a grazing food chain in Peipsi and a detrital food-chain in Võrtsjärv. Consequently, top-down control of phytoplankton can be assumed to be much more important in Peipsi than in Võrtsjärv. When the detrital food chain prevails, the planktonic ciliates become the most important zooplankton group in shallow, eutrophic and large lake. Neglecting protozooplankton can result in serious underestimates of total zooplankton biomass since two-thirds of the zooplankton biomass in Võrtsjärv comprises ciliates.     

Temporal dynamics in epipelic, pelagic and epiphytic algal production in a clear and a turbid shallow lake

SUMMARY 1. Pelagic and epipelic microalgal production were measured over a year in a pre-defined area (depth 0.5 m) in each of two lakes, one turbid and one with clear water. Further estimates of epiphytic production within reed stands were obtained by measuring production of periphyton developed on artificial substrata.
2. Total annual production of phytoplankton and epipelon was 34% greater in the turbid lake (190 g C m⁻² year⁻¹) than in the clearwater lake (141 g C m⁻² year⁻¹). However, the ratio of total production to mean water column TP concentration was two fold greater in the clearwater lake.
3. Phytoplankton accounted for the majority of the annual production (96%) in the turbid lake, while epipelic microalgal production dominated (77%) in the clear lake. The relative contribution of epipelic algae varied over the year, however, and in the turbid lake was higher in winter (11–25%), when the water was relatively clear, than during summer (0.7–1.7%), when the water was more turbid. In the clearwater lake, the relative contribution of epipelon was high both in winter, when the water was most clear, and in mid-summer, when phytoplankton production was constrained either by nutrients or grazing.
4. Compared with pelagic and epipelic primary production, epiphytic production within a reed stand was low and did not vary significantly between the lakes.
5. The study supports the theory of a competitive and compensatory trade-off between primary producers in lakes with contrasting nutrient concentrations, resulting in relatively small differences in overall production between clear and turbid lakes when integrating over the season and over different habitats.     

Clay-Turbid Interactions May Not Cascade—A Reminder for Lake Managers

Food web management is a frequently used lake restoration method, which aims to reduce phytoplankton biomass by strengthening herbivorous zooplankton through reduction of planktivorous fish. However, in clay‐turbid lakes several factors may reduce the effectivity of food web management. Increasing turbidity reduces the effectivity of fish predation and weakens the link between zooplankton and phytoplankton. Therefore, the effects of fish stock manipulations may not cascade to lower trophic levels as expected. Additionally, in clay‐turbid conditions invertebrate predators may coexist in high densities with planktivorous fish and negate the effects of fish reductions. For instance, in the stratifying regions of the clay‐turbid Lake Hiidenvesi, Chaoborus flavicans is the main regulator of cladocerans and occupies the water column throughout the day, although planktivorous Osmerus eperlanus is very abundant. The coexistence of chaoborids and fish is facilitated by a metalimnetic turbidity peak, which prevents efficient predation by fish. In the shallow parts of the lake, chaoborids are absent despite high water turbidity. We suggest that, generally, the importance of invertebrate predators in relation to vertebrate predators may change along turbidity and depth gradients. The importance of fish predation is highest in shallow waters with low turbidity. When water depth increases, the importance of fish in the top‐down regulation of zooplankton declines, whereas that of chaoborids increases, the change along the depth gradient being moderate in clear‐water lakes and steep in highly turbid lakes. Thus, especially deep clay‐turbid lakes may be problematic for implementing food web management as a restoration tool.     

The relationship between nutrients and trophic‐level biomass in turbid tropical ponds

1. The total phosphorus–algal biomass relationship from a set of turbid tropical ponds in Kenya was compared with predictions derived from surveys of temperate and subtropical lakes. Despite high concentrations of total phosphorus (TP) (up to 797 μg L ^–1) and inorganic turbidity (up to 800 mg L^–1), the log–log relationship between algal biomass and TP was steeper than expected.
2. No evidence of nitrogen limitation was found at high TP, and total nitrogen (TN):TP ratios were higher than in lakes with similar TP levels studied previously. High TN:TP ratios may be a consequence of excretion by cattle into the ponds, a nutrient source characterized by a high ratio of available N to available P.
3. Despite extremely high turbidity, the ratio of mixed layer depth to euphotic depth was generally low because these ponds are shallow (≤ 2 m), and was not related to algal yield. A positive relationship was also found between TP and zooplankton biomass, and between TP and the density of the zooplanktivorous bug, Anisops . In contrast, no relationship was found between fish biomass and TP, algal biomass or zooplankton biomass.     

Colourful coexistence of red and green picocyanobacteria in lakes and seas

Hutchinson's paradox of the plankton inspired many studies on the mechanisms of species coexistence. Recent laboratory experiments showed that partitioning of white light allows stable coexistence of red and green picocyanobacteria. Here, we investigate to what extent these laboratory findings can be extrapolated to natural waters. We predict from a parameterized competition model that the underwater light colour of lakes and seas provides ample opportunities for coexistence of red and green phytoplankton species. To test this prediction, we sampled picocyanobacteria of 70 aquatic ecosystems, ranging from clear blue oceans to turbid brown peat lakes. As predicted, red picocyanobacteria dominated in clear waters, whereas green picocyanobacteria dominated in turbid waters. We found widespread coexistence of red and green picocyanobacteria in waters of intermediate turbidity. These field data support the hypothesis that niche differentiation along the light spectrum promotes phytoplankton biodiversity, thus providing a colourful solution to the paradox of the plankton.     

Impact of climatic fluctuations on Characeae biomass in a shallow,restored lake in The Netherlands

External phosphorus load to a wetland with two shallow lakes in the Botshol Nature Reserve, The Netherlands, was reduced, resulting in a rapid reduction of phytoplankton biomass and turbidity, and after 4 years, explosive growth of Characeae. The clear water state was unstable, however, and the ecosystem then alternated between clear, high-vegetation and turbid, low-vegetation states. A model of water quality processes was used in conjunction with a 14-year nutrient budget for Botshol to determine if fluctuations in precipitation and nutrient load caused the ecosystem instability. The results indicate that, during wet winters when groundwater level rose above surface water level, phosphorus from runoff was stored in the lake bottom and banks. Stored phosphorus was released the following spring and summer under anaerobic sediment conditions, resulting in increased phytoplankton density and light attenuation in the water column. During years with high net precipitation, flow from land to surface water also transported humic acids, further increasing light attenuation. In years with dry winters, the phosphorus and humic acid loads to surface water were reduced, and growth of submerged macrophytes was enhanced by clear water. Thus, the temporal pattern of precipitation and flow from land to water gave a coherent, quantitative explanation of the observed dynamics in phosphorus, phytoplankton, turbidity, and Characeae. Global warming has caused winters in The Netherlands to become warmer and wetter during the last 50 years, increasing flow from land to water of humic acids and phosphorus and, ultimately, enhancing instability of Characeae populations. In the first half of the 20th century interannual variation in precipitation was not sufficient to cause large changes in internal P flux in Botshol, and submerged macrophyte populations were stable.     

Wetland macrophyte diversity, trophic status and phytoplankton： a case study of the cascading effects of an invasive herbivore

Shallow water bodies can exist in alternative stable states, a clear water state with high coverage of macrophytes or a turbid state with high phytoplankton biomass. The alternative equilibria hypothesis has been proposed to explain the occurrence of the alternative stable states (Scheffer et al., 1993)[1], which assumes that: 1),     

Alternation between clear,high-vegetation and turbid,low-vegetation states in a shallow lake: the role of birds

External phosphorus load to a wetland with two shallow lakes in the Botshol Nature Reserve, the Netherlands, was reduced since 1989, resulting in a rapid reduction of phosphorus levels, phytoplankton biomass and turbidity, and after 4 years, explosive growth of Characeae. The clear water state was unstable, however, and the ecosystem subsequently alternated between clear, high-vegetation and turbid, low-vegetation states. The switch from clear to turbid states occurred at a higher phosphorus level than the switch from turbid to clear states and area covered by macrophytes was significantly correlated to Secchi disc depth (r = 0.86, p < 0.001). Beginning in 1997, phosphorus input from droppings of greylag geese and black-headed gulls increased. The hypothesis that grazing prevented the return of Characeae populations was tested during 1996–1997 using large-scale exclosure experiments. The biomass and cover of Characeae were monitored in 17 exclosures for 2 years. The experiments showed no significant increase in Characeae in the absence of grazers compared with the controls. During the course of the experiments, the entire lake switched from turbid water to clear water and high Chara biomass. These ecosystem developments suggest that light limitation was the main factor controlling the collapse and return of Characeae in Botshol.     

Changes in physicochemical and biological factors during regime shifts in a restoration demonstration of macrophytes in a small hypereutrophic Chinese lake

Shallow, eutrophic lakes are usually characterized by a turbid state devoid of submerged vegetation subject to human-induced eutrophication. In most cases, it is rather hard to restore a vegetated clear state due to reduced resilience caused by a blend of complicated factors. In this study, we successfully reestablished a plant community in a small hypereutrophic lake over a certain period. In winter and spring with transparency of >55 cm and temperature of <20 °C, a submerged stands bed formed gradually under strong human interventions. The reestablished plant bed displayed obvious seasonal succession and prolonged the clear-water stage until July 2005, when it collapsed. The regime shift to a turbid state was mainly attributed to the decreasing biomass of stands bed and mechanical damage brought about by the elimination of Spirodela polyrhiza, increasing water temperature, P concentration as well as periphyton biomass, etc. The reestablishment also changed the aquatic ecosystem greatly. A ‘clear-water’ stage was characterized by higher NO₃^?–N, NH₄⁺–N, electrical conductivity, transparency and TN/TP level and more cladocerans (mainly Daphnia pulex), while the turbid state was characterized by higher temperature, chlorophyll a and TP level and more abundant rotifers. It is thus viable to restore submerged macrophytes in such lakes in winter and spring, when transparency is relatively high while temperature and water level are low. Nevertheless, to obtain a long-term, vegetated clear state, control of internal nutrient loading by means of obstruction, purification, dredging or solidification, is extremely necessary since nutrients play an important role in regime shifts as evidenced by the present case, too.     

Zooplankton, phytoplankton and the microbial food web in two turbid and two clearwater shallow lakes in Belgium

Components of the pelagic food web in four eutrophic shallow lakes in two wetland reserves in Belgium (Blankaart and De Maten) were monitored during the course of 1998–1999. In each wetland reserve, a clearwater and a turbid lake were sampled. The two lakes in each wetland reserve had similar nutrient loadings and occurred in close proximity of each other. In accordance with the alternative stable states theory, food web structure differed strongly between the clearwater and turbid lakes. Phytoplankton biomass was higher in the turbid than the clearwater lakes. Whereas chlorophytes dominated the phytoplankton in the turbid lakes, cryptophytes were the most important phytoplankton group in the clearwater lakes. The biomass of microheterotrophs (bacteria, heterotrophic nanoflagellates and ciliates) was higher in the turbid than the clearwater lakes. Biomass and community composition of micro- and macrozooplankton was not clearly related to water clarity. The ratio of macrozooplankton to phytoplankton biomass – an indicator of zooplankton grazing pressure on phytoplankton – was higher in the clearwater when compared to the turbid lakes. The factors potentially regulating water clarity, phytoplankton, microheterotrophs and macrozooplankton are discussed. Implications for the management of these lakes are discussed.