Predicting the effect of climate change on temperate shallow lakes with the ecosystem model PCLake

Global average surface temperatures are expected to rise by about 1.4–5.8°C from the present until the year 2100. This temperature increase will affect all ecosystems on earth. For shallow lakes—which can be either in a clear water or a turbid state—this climate change will expectedly negatively affect water transparency though the prediction is far from conclusive and experimental investigations elucidating the potential climatic effects on shallow lakes are still rare. The aim of this study was to further shape and sharpens hypotheses on the impact of climate change on shallow lakes by applying an existing and well-calibrated ecosystem model, PCLake. We focused on asymptotic model behaviour for a range of temperature and loading scenarios in a factorial design. We conclude that climate change will likely lead to decreased critical nutrient loadings. Combined with an expected increase in the external nutrient loading, this will increase the probability of a shift from a clear to a turbid state. As the model predicts a higher summer chlorophyll-a concentration, a stronger dominance of cyanobacteria during summer and a reduced zooplankton abundance due to climate change, the turbid state itself is likely to become even more severe.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

Mass flux calculations show strong allochthonous support of freshwater zooplankton production is unlikely

Many studies have concluded terrestrial carbon inputs contribute 20-70% of the carbon supporting zooplankton and fish production in lakes. Conversely, it is also known that terrestrial carbon inputs are of very low nutritional quality and phytoplankton are strongly preferentially utilized by zooplankton. Because of its low quality, substantial terrestrial support of zooplankton production in lakes is only conceivable when terrigenous organic matter inputs are much larger than algal production. We conducted a quantitative analysis of terrestrial carbon mass influx and algal primary production estimates for oligo/mesotrophic lakes (i.e., TP ≤ 20 μg L(-1)). In keeping with the principle of mass conservation, only the flux of terrestrial carbon retained within lakes can be utilized by zooplankton. Our field data compilation showed the median (inter-quartile range) terrestrial particulate organic carbon (t-POC), available dissolved organic carbon (t-DOC) inputs, and in-lake bacterial and algal production were 11 (8-17), 34 (11-78), 74 (37-165), and 253 (115-546) mg C m(-2) d(-1), respectively. Despite the widespread view that terrestrial inputs dominate the carbon flux of many lakes, our analysis indicates algal production is a factor 4-7 greater than the available flux of allochthonous basal resources in low productivity lakes. Lakes with high loading of t-DOC also have high hydraulic flushing rates. Because t-DOC is processed, i.e., mineralized or lost to the sediments, in lakes at ≈ 0.1% d(-1), in systems with the highest t-DOC inputs (i.e., 1000 mg m(-2) d(-1)) a median of 98% of the t-DOC flux is advected and therefore is not available to support zooplankton production. Further, advection is the primary fate of t-DOC in lakes with hydraulic retention times <3 years. When taking into account the availability and quality of terrestrial and autochthonous fluxes, this analysis indicates ≈ 95-99% of aquatic herbivore production is supported by in-lake primary production.     

The Importance of Lake Morphometry for the Structureand Function of Lakes

This work demonstrates quantitatively and in a comprehensive way that the size and form of lakes regulate many general transport processes, such as sedimentation, resuspension, diffusion, mixing, burial and outflow, which in turn regulate many abiotic state variables, such as concentrations of phosphorus, suspended particulate matter, many water chemical variables and water clarity, which in turn regulate primary production, which regulate secondary production, for example of zooplankton and fish. Such relationships are discussed not qualitatively but quantitatively using a new generation of validated dynamic ecosystem models (LakeWeb and LakeMab) based on mechanistic principles. It has been shown by critical model tests (including blind tests using data covering wide limnological ranges) that these models give predictions that agree well with empirical data. This should lend credibility to the results presented in this work, which would have been very difficult to obtain using traditional methods with extensive field studies in a few lakes. Simulations have been carried out where the inflow of phosphorus is held constant and the consequences simulated for small, large, shallow and deep lakes. There are striking differences in total phosphorus (TP) concentrations and trophic state (from 10 to 100 µg TP/l) and hence also in changes in many variables characterizing lake structure and function, such as Secchi depth, suspended particulate matter, pH, water temperature, chlorophyll, algal volume, macrophyte cover; as well as production and biomasses of benthic algae, bacterioplankton, macrophytes, herbivorous zooplankton, predatory zooplankton, zoobenthos, prey fish and predatory fish. These changes have been quantified in a comprehensive manner in this work and the approach to calculate such changes are basic for an understanding of how different lakes react to changes in nutrient loading (eutrophication). (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Variability of bio-optical parameters in two North-European large lakes

The bio-optical properties of some North-European large lakes were examined during 1995–2005 using field data and laboratory measurements. The key variables were optically active substances (OAS: chlorophyll, total suspended matter and dissolved organic matter), Secchi depth, and the “spectrometric” and diffuse light attenuation coefficients. Our main study sites were Lake Peipsi and Lake Võrtsjärv in Estonia, both eutrophic with mean Secchi depth below 3 m. The measured water parameters were compared with those obtained from two clear-water Swedish lakes, Lake Vänern and Lake Vättern. This comparison describes the bio-optical differences of the water in eutrophic and oligotrophic lakes. The variability of water parameters in the turbid Estonian lakes was rather high, e.g. the chlorophyll content varied from 1.8 to 102 mg m^?3 and the diffuse light attenuation coefficient from 0.92 to 6.5 m^?1. The change in water properties depends on the season and the biological activity of phytoplankton. We found no apparent long-time trend in water properties. Regression analysis showed that in the turbid Estonian lakes the optical properties were well correlated with chlorophyll and suspended matter, but not with dissolved organic matter. The highest determination coefficients (between 0.73 and 0.89) were obtained when the optical parameters were correlated with all three OAS together (multiple regressions). Our results concerning the variability and interconnections among bio-optical parameters in two Estonian large lakes illustrate the effect of OAS and light field on the ecological conditions of lakes in general.     

The effect of turbidity state and microhabitat on macroinvertebrate assemblages: a pilot study of six shallow lakes

Shallow lakes can occur in two alternative stable states, a clear-water state and a turbid state. This is associated with separate assemblages of fish, zooplankton and plants. Little is known about whether macroinvertebrate assemblages differ across both stable states. This study investigated this in a connected set of three turbid and three clear-water shallow lakes. To overcome confounding effects of differences in spatial structure of macrophytes in turbid and clear-water lakes, we sampled three microhabitats that occurred in both alternative stable states: open water, sago pondweed (Potamogeton pectinatus) and reed (Phragmites australis). Univariate analyses indicated no differences in the number of organisms, taxon richness or diversity between turbid and clear-water lakes. Multivariate analysis, however, showed significant differences in the macroinvertebrate community structure of both stable states. Nine taxa explained a significant amount of the variation between both lake types, of which seven preferred the clear-water lakes. The number of organisms and the taxon richness were higher in reed than in the other microhabitats, but diversity and evenness did not differ among the microhabitats. Multivariate analyses could separate all three microhabitats. Eight taxa, mainly detritus feeders and collector–gatherers, explained most of the variation in the data and preferred the reed microhabitat. The effects of stable state (6.8% explained variance) and microhabitat (13.1% explained variance) on the macroinvertebrate assemblages were largely independent from each other (1.5% shared variance). Although macroinvertebrates are not implemented in the initial theory of stable states, our results show clearly different assemblages across both stable states.     

Bottom‐up and top‐down effects of browning and warming on shallow lake food webs

Productivity and trophic structure of aquatic ecosystems result from a complex interplay of bottom‐up and top‐down forces that operate across benthic and pelagic food web compartments. Projected global changes urge the question how this interplay will be affected by browning (increasing input of terrestrial dissolved organic matter), nutrient enrichment and warming. We explored this with a process‐based model of a shallow lake food web consisting of benthic and pelagic components (abiotic resources, primary producers, grazers, carnivores), and compared model expectations with the results of a browning and warming experiment in nutrient‐poor ponds harboring a boreal lake community. Under low nutrient conditions, the model makes three major predictions. (a) Browning reduces light and increases nutrient supply; this decreases benthic and increases pelagic production, gradually shifting productivity from the benthic to the pelagic habitat. (b) Because of active habitat choice, fish exert top‐down control on grazers and benefit primary producers primarily in the more productive of the two habitats. (c) Warming relaxes top‐down control of grazers by fish and decreases primary producer biomass, but effects of warming are generally small compared to effects of browning and nutrient supply. Experimental results were consistent with most model predictions for browning: light penetration, benthic algal production, and zoobenthos biomass decreased, and pelagic nutrients and pelagic algal production increased with browning. Also consistent with expectations, warming had negative effects on benthic and pelagic algal biomass and weak effects on algal production and zoobenthos and zooplankton biomass. Inconsistent with expectations, browning had no effect on zooplankton and warming effects on fish depended on browning. The model is applicable also to nutrient‐rich systems, and we propose that it is a useful tool for the exploration of the consequences of different climate change scenarios for productivity and food web dynamics in shallow lakes, the worldwide most common lake type.     

Planktonic ciliate community structure in shallow lakes of lowland Western Europe

Temperate shallow meso- to eutrophic lakes can exist in one of two alternative states with contrasting foodwebs, referred to as the clear-water and the turbid state. We describe the planktonic ciliate communities of such lakes based on a survey of 66 northwestern European lakes. Ciliates were enumerated and identified to species level according to the quantitative protargol staining technique. Ciliate biomass was on average twice as high in the turbid than in the clear-water lakes. The ciliate communities were dominated by oligotrichs and protostomatids, and no differences in functional composition or α-diversity could be detected between turbid and clear-water lakes, although β-diversity tended to be higher in the latter. At the species level, however, community structure strongly differed between turbid and clear-water lakes, and several indicator species could be identified for the different lake categories. Variation partitioning showed that nutrient status did not explain ciliate community structure independent of the alternative states, while lake area was identified as an additional structuring factor for the ciliate communities. These results stress the importance of the ecosystem structure in shaping ciliate communities in temperate shallow lakes and suggest that nutrient status has little direct effect on ciliate community structure in such lakes.     

Habitat structure determines resource use by zooplankton in temperate lakes

While the importance of terrestrial linkages to aquatic ecosystems is well appreciated, the degree of terrestrial support of aquatic consumers remains debated. Estimates of terrestrial contributions to lake zooplankton have omitted a key food source, phytoplankton produced below the mixed layer. We used carbon and nitrogen stable isotope data from 25 Pacific Northwest lakes to assess the relative importance of particulate organic matter (POM) from the mixed layer, below the mixed layer and terrestrial detritus to zooplankton. Zooplankton and deep POM were depleted in 13C relative to mixed layer POM in lakes that can support deep primary production. A Bayesian stable isotope mixing model estimated that terrestrial detritus contributed <5% to zooplankton production, and confirms the role of lake optical and thermal properties; deep POM accounted for up to 80% of zooplankton production in the clearest lakes. These results suggest terrestrial support of lake zooplankton production is trivial.     

Impacts of elevated terrestrial nutrient loads and temperature on pelagic food‐web efficiency and fish production

Both temperature and terrestrial organic matter have strong impacts on aquatic food‐web dynamics and production. Temperature affects vital rates of all organisms, and terrestrial organic matter can act both as an energy source for lower trophic levels, while simultaneously reducing light availability for autotrophic production. As climate change predictions for the Baltic Sea and elsewhere suggest increases in both terrestrial matter runoff and increases in temperature, we studied the effects on pelagic food‐web dynamics and food‐web efficiency in a plausible future scenario with respect to these abiotic variables in a large‐scale mesocosm experiment. Total basal (phytoplankton plus bacterial) production was slightly reduced when only increasing temperatures, but was otherwise similar across all other treatments. Separate increases in nutrient loads and temperature decreased the ratio of autotrophic:heterotrophic production, but the combined treatment of elevated temperature and terrestrial nutrient loads increased both fish production and food‐web efficiency. CDOM: Chl a ratios strongly indicated that terrestrial and not autotrophic carbon was the main energy source in these food webs and our results also showed that zooplankton biomass was positively correlated with increased bacterial production. Concomitantly, biomass of the dominant calanoid copepod Acartia sp. increased as an effect of increased temperature. As the combined effects of increased temperature and terrestrial organic nutrient loads were required to increase zooplankton abundance and fish production, conclusions about effects of climate change on food‐web dynamics and fish production must be based on realistic combinations of several abiotic factors. Moreover, our results question established notions on the net inefficiency of heterotrophic carbon transfer to the top of the food web.     

Terrestrial subsidies to lake food webs: an experimental approach

Cross-ecosystem movements of material and energy are ubiquitous. Aquatic ecosystems typically receive material that also includes organic matter from the surrounding catchment. Terrestrial-derived (allochthonous) organic matter can enter aquatic ecosystems in dissolved or particulate form. Several studies have highlighted the importance of dissolved organic carbon to aquatic consumers, but less is known about allochthonous particulate organic carbon (POC). Similarly, most studies showing the effects of allochthonous organic carbon (OC) on aquatic consumers have investigated pelagic habitats; the effects of allochthonous OC on benthic communities are less well studied. Allochthonous inputs might further decrease primary production through light reduction, thereby potentially affecting autotrophic resource availability to consumers. Here, an enclosure experiment was carried out to test the importance of POC input and light availability on the resource use in a benthic food web of a clear-water lake. Corn starch (a C(4) plant) was used as a POC source due to its insoluble nature and its distinct carbon stable isotope value (δ(13)C). The starch carbon was closely dispersed over the bottom of the enclosures to study the fate of a POC source exclusively available to sediment biota. The addition of starch carbon resulted in a clear shift in the isotopic signature of surface-dwelling herbivorous and predatory invertebrates. Although the starch carbon was added solely to the sediment surface, the carbon originating from the starch reached zooplankton. We suggest that allochthonous POC can subsidize benthic food webs directly and can be further transferred to pelagic systems, thereby highlighting the importance of benthic pathways for pelagic habitats.     

Effects of hydrological forcing on the structure of a tropical estuarine food web

River flow can impact which sources of particulate organic matter (POM) fuel estuarine food webs. Here, we used stable carbon (C) and nitrogen (N) isotope analyses to compare how contributions of different POM sources (terrestrial, estuarine, and marine) to the diets of zooplankton and juvenile fishes differed between low and high river flow conditions, as well as spatially across a tropical estuary, Hilo Bay, Hawaii, USA. Diets of zooplankton and juvenile fishes were affected by river flow conditions, but the magnitude and the change in the basal resources depended on the location of the station in the estuary relative to the ocean and the river mouths. Consumers from the station most isolated from the ocean and with groundwater and overland flow inputs, utilized a combination of estuarine and terrestrial POM during both low and high river flow conditions and exhibited less variability in their basal resources than stations with direct ocean exchange. Consumers from stations in the Bay most affected by ocean exchange and river inputs utilized a combination of estuarine, terrestrial, and marine POM during low flow conditions, but shifted to marine and terrestrial POM during high river flow conditions. This shift to using terrestrial POM during high river flow conditions was substantial and up to 40% higher than values measured in other estuaries. Factors suspected to be affecting which POM source(s) consumers use in Hilo Bay are gross primary production, biological availability of exported terrestrial OM, and estuarine bacteria biomass, all of which are affected by river flow. Overall, our results suggest that Hilo Bay's food web and possibly those from other tropical estuaries are vulnerable to changes in hydrology, which may be further enhanced by global climate change.     

Stable isotope analysis of the origins of zooplankton carbon in lakes of differing trophic state

Carbon stable isotope analysis was carried out on zooplankton from 24 United Kingdom lakes to examine the hypothesis that zooplankton dependence on allochthonous sources of organic carbon declines with increasing lake trophy. Stable isotope analysis was also carried out on particulate and dissolved organic matter (POM and DOM) and, in 11 of the lakes, of phytoplankton isolates. In 21 of the 24 lakes, the zooplankton were depleted in ¹³C relative to bulk POM, consistent with previous reports. δ¹³C for POM showed relatively little variation between lakes compared to high variation in values for DOM and phytoplankton. δ¹³C values for phytoplankton and POM converged with increasing lake trophy, consistent with the expected greater contribution of autochthonous production to the total organic matter pool in eutrophic lakes. The difference between δ¹³C for zooplankton and that for POM was also greatest in oligotrophic lakes and reduced in mesotrophic lakes, in accordance with the hypothesis that increasing lake trophic state leads to greater dependence of zooplankton on phytoplankton production. However, the difference increased again in hypertrophic lakes, where higher δ¹³C values for POM may have been due to greater inputs of ¹³C-enriched organic matter from the littoral zone. The very wide variation in phytoplankton δ¹³C between lakes of all trophic categories made it difficult to detect robust patterns in the variation in δ¹³C for zooplankton. Received: 2 November 1998 / Accepted: 3 December 1999     

Climate drivers of diatom distribution in shallow subarctic lakes

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Identification of total phosphate,submerged vegetation cover and zooplankton size thresholds for success of biomanipulation in peri-urban eutrophic ponds

In small shallow lakes and ponds, the clear-water state can generally be maintained at higher nutrient concentrations compared to larger shallow lakes. The main objective of this study was to identify thresholds for total phosphorus (TP), submerged vegetation cover and zooplankton size that determine biomanipulation success in peri-urban eutrophic ponds. Additionally, the relationship between transparency and TP is discussed with regard to similar relationships and thresholds reported for shallow lakes. Using classification trees, a threshold TP concentration of 0.300 mg P L^?1 was determined below which a clear-water state was generally maintained after biomanipulation. When the average TP concentration was >0.300 mg P L^?1, the stability of the clear-water state largely depended on the presence of sufficiently large zooplankton (>0.87 mm) or a submerged vegetation cover of >82% at some point during the year. This threshold TP concentration is considerably higher than the threshold of 0.1 mg L^?1 which is generally suggested for longer-term success of biomanipulation in shallow lakes. Such threshold nutrient concentration is important when restoring ecological quality in eutrophic small lakes and ponds. Extended follow-up of biomanipulation success in eutrophic ponds could provide more insight into the feasibility of these thresholds on the longer term.     

Physical and chemical limnology of 204 lakes from the Canadian Arctic Archipelago

The physical and chemical limnology of 204 lakes from across the Canadian Arctic Archipelago was examined. Mean summer air temperature did not correlate well with lake chlorophyll levels due to the predominance of ultra-oligotrophic hard-water lakes located in a polar climate. Local geology influences ion budgets and is an important factor in determining pelagic phosphorus availability, carbon cycling and metal concentrations. Ratios of particulate carbon, particulate nitrogen and chlorophyll a indicate that planktonic microorganisms are not always the major producers of organic carbon in arctic lakes. Allochthonous particulate matter contributes significantly to the carbon and phosphorus budgets of small and mid-sized lakes across the Arctic, although the availability of these elements is controlled by many interacting geochemical and biological factors. Phosphorus is generally limiting, however, increases in available phosphorus, nitrogen and carbon are all required to make significant long-term differences in lake productivity. Particulate phosphorus levels can be high in lakes where phosphorus-rich shales or carbonatite bedrock are present. These phosphorus-enriched lakes are found in several areas across the mid-arctic islands, however, only small amounts of this nutrient are available as soluble reactive phosphorus. Although lakes throughout the Arctic are typically ultra-oligotrophic, they still represent an important sink for allochthonous nutrient deposition.     

Essential fatty acids and phosphorus in seston from lakes with contrasting terrestrial dissolved organic carbon content

1. It is often assumed that lakes highly influenced by terrestrial organic matter (TOM) have low zooplankton food quality because of elemental and/or biochemical deficiencies of the major particulate organic carbon pools. We used the biochemical [polyunsaturated fatty acids, especially eicosapentaenoic acid (EPA) – 20:5ω3] and elemental (C : P ratio) composition of particulate matter (PM) as qualitative measures of potential zooplankton food in two categories of lakes of similar primary productivity, but with contrasting TOM influence (clear water versus humic lakes). 2. C : P ratios (atomic ratio) in PM were similar between lake categories and were above 400. The concentration (μg L⁻¹) and relative content (μg mg C⁻¹) of EPA, as well as the particulate organic carbon concentration, were higher in the humic lakes than in the clear‐water lakes. 3. Our results show high fatty acid quality of PM in the humic lakes. The differences in the biochemical quality of the potential zooplankton food between lake categories can be attributed to the differences in their phytoplankton communities. 4. High biochemical quality of the food can result in high efficiency of energy transfer in the food chain and stimulate production at higher trophic levels, assuming that zooplankton are able to ingest and digest the resource available.     

Tiny Is Mighty: Seagrass Beds Have a Large Role in the Export of Organic Material in the Tropical Coastal Zone

Ecosystems in the tropical coastal zone exchange particulate organic matter (POM) with adjacent systems, but differences in this function among ecosystems remain poorly quantified. Seagrass beds are often a relatively small section of this coastal zone, but have a potentially much larger ecological influence than suggested by their surface area. Using stable isotopes as tracers of oceanic, terrestrial, mangrove and seagrass sources, we investigated the origin of particulate organic matter in nine mangrove bays around the island of Phuket (Thailand). We used a linear mixing model based on bulk organic carbon, total nitrogen and δ¹³C and δ¹⁵N and found that oceanic sources dominated suspended particulate organic matter samples along the mangrove-seagrass-ocean gradient. Sediment trap samples showed contributions from four sources oceanic, mangrove forest/terrestrial and seagrass beds where oceanic had the strongest contribution and seagrass beds the smallest. Based on ecosystem area, however, the contribution of suspended particulate organic matter derived from seagrass beds was disproportionally high, relative to the entire area occupied by mangrove forests, the catchment area (terrestrial) and seagrass beds. The contribution from mangrove forests was approximately equal to their surface area, whereas terrestrial contributions to suspended organic matter under contributed compared to their relative catchment area. Interestingly, mangrove forest contribution at 0 m on the transects showed a positive relationship with the exposed frontal width of the mangrove, indicating that mangrove forest exposure to hydrodynamic energy may be a controlling factor in mangrove outwelling. However we found no relationship between seagrass bed contribution and any physical factors, which we measured. Our results indicate that although seagrass beds occupy a relatively small area of the coastal zone, their role in the export of organic matter is disproportional and should be considered in coastal management especially with respect to their importance as a nutrient source for other ecosystems and organisms.     

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.