Temperate Shallow Lakes Research Articles

Seasonal Dynamics of CO2 Flux Across the Surface of Shallow Temperate Lakes

We used data collected from 1989 to 2009 from 151 shallow (mean depth < 3 m) temperate lakes in Denmark to explore the influence of lake trophic status, surface area and catchment size on the seasonal dynamics of the air–water flux of CO2. Monthly CO2 fluxes were derived from measurements of acid neutralizing capacity (ANC), pH, ionic strength, temperature, and wind speed. CO2 fluxes exhibited large seasonal variability, in particular in oligo-mesotrophic lakes. Most of the lakes emitted CO2 during winter (median rates ranging 300–1,900 mg C m−2 day−1), and less CO2 during summer or, in the case of some of the highly eutrophic lakes, retained CO2 during summer. We found that seasonal CO2 fluxes were strongly negatively correlated with pH (r = −0.65, P < 0.01), which in turn was correlated with chlorophyll a concentrations (r = 0.48, P < 0.01). Our analysis suggests that lake trophic status (a proxy for pelagic production) interacts with the lake ANC to drive the seasonal dynamics of CO2 fluxes, largely by changing pH and thereby the equilibrium of the free CO2 and bicarbonate relation. Long-term observations from four lakes, which have all undergone a period of oligotrophication during the past two decades, provide further evidence that CO2 efflux generally increases as trophic status decreases, as a consequence of decreased pH. Across these four lakes, the annual average CO2 emission has increased by 32% during the past two decades, thus, demonstrating the strong link between lake trophic status and CO2 flux.

Bacterial communities in sediments of the shallow Lake Dongping in China

The purpose of this study was to discuss how the environmental inputs and anthropogenic activities impact bacterial communities in the sediments of a shallow, eutrophic and temperate freshwater lake. Sediment cores were collected from Lake Dongping, located in Taian, Shandong, China. All samples were processed within 4 h of collection. Total nitrogen, total phosphorus (TP), total organic carbon, ammonium nitrogen and nitrate nitrogen content of samples were measured by Kjeldahl determination, sulphuric acid-perchloric acid digestion and molybdenum blue colorimetry, potassium dichromate titration, Nessler's reagent colorimetric and the phenol disulphonic acid colorimetric method, respectively. Seasonal and temporal diversity of sediment bacterial communities at six stations in Lake Dongping were investigated using molecular approaches (terminal restriction fragment length polymorphism and 16S rDNA clone libraries). Noticeable seasonal and temporal variations were observed in bacterial diversity and composition at all six stations. Sediment bacterial communities in Lake Dongping belonged to 16 phyla: Proteobacteria (including α-Proteobacteria, β-Proteobacteria, δ-Proteobacteria, ε-Proteobacteria, γ-Proteobacteria), Acidobacteria, Planctomycetes, Bacteroidetes, Firmicutes, Verrucomicrobia, Nitrospira, Chloroflexi, Gemmatimonadetes, Chlorobi, Cyanobacteria, Deferribacteres, Actinobacteria, OP8, Spirochaetes and OP11. Members of β-, δ- and γ-Proteobacterial sequences were predominant in 11 of 12 clone libraries derived from sediment samples. Sediment samples collected at stations 1 and 4 in July had the greatest bacterial diversity while those collected at station 2 in October had the least diversity. TP concentration was significantly correlated with the distribution of bacterial communities. Our results suggested that different environmental nutrient inputs contribute to seasonal and temporal variations of chemical features and bacterial communities in sediments of Lake Dongping. TP concentration was significantly correlated with the distribution of bacterial communities. This study has an important implication for the optimization of integrated ecosystem assessment of shallow temperate freshwater lake and provides interesting information for the subsequent of the ecosystem.

Modeling and Validating Long‐Term Dynamics of Diel Dissolved Oxygen with Particular Reference to pH in a Temperate Shallow Lake (Turkey)

AbstractLong‐term monitoring of changes in dissolved oxygen (DO) and pH is of great importance to quantifying aquatic ecosystem metabolism, particularly for lakes under the changing global environment. During 173 days, diel DO cycles were measured in situ along with the main driving variables of pH, wind speed (WS), and net solar radiation (Rn) in a temperate shallow lake. Best‐fit multiple non‐linear regression (MNLR) models of diel DO time series were built and validated on a monthly basis, with R2 values ranging from 42.4% in September to 95.4% in November for validation. The strong relationship between diel DO and pH (r = 0.6) appeared to be related to the patterns of ecosystem productivity and respiration, and sensitivity of decomposing bacteria to changes in pH. pH‐driven lake metabolism appears to have significant implications for diel and seasonal lake metabolism in a changing global environment.

Are zooplankton food resources poor in the vegetated littoral zone of shallow lakes?

Summary 1. The distribution of zooplankton in shallow lakes is negatively related to macrophyte density. However, the abundance of their food along density gradients of macrophytes is unknown. A common but untested assumption is that food quantity and quality for pelagic zooplankton is poor in the littoral zone owing to the deleterious influence of macrophytes on phytoplankton. 2. We tested this assumption with a combination of a field survey and laboratory experiments. We collected seston samples from the littoral and pelagic zones of four shallow temperate lakes and related food quantity (phytoplankton biovolume) and quality to macrophyte abundance (per cent volume infested). Seston food quality was assessed in three ways: N/C and P/C ratios, polyunsaturated fatty acid content and phytoplankton community composition. In the laboratory, we measured the growth and reproduction of Daphnia pulicaria on diets consisting of seston from the littoral and pelagic zones in one lake. 3. In our four study lakes, food quantity was not significantly influenced by macrophyte abundance, and food quality was generally high. Laboratory experiments showed increased juvenile growth, but no significant change in D. pulicaria reproduction, when feeding on littoral resources compared to pelagic resources. 4. Our results suggest that there is no nutritional cost to pelagic zooplankton inhabiting the littoral zone. Therefore, it is likely that other factors (e.g. predation, abiotic factors) are involved in determining zooplankton habitat use.

Reconstructed long-term time series of phytoplankton primary production of a large shallow temperate lake: the basis to assess the carbon balance and its climate sensitivity

Aiming at building the carbon budget for further climate change impact research in the large and shallow northern temperate Lake Vortsjarv, the present paper focuses on reconstructing the full phytoplankton primary production (PP) data series for the lake for the period of 1982–2009 covered by disconnected measurements, and testing the uncertainties involved both in the PP measurements and bio-optical modelling. During this 28-year period, in situ PP was measured in Vortsjarv in 18 years with 14C-assimilation technique. We reconstructed the full time series using a semi-empirical PP simulation model based on continuously measured PAR irradiance and interpolated values of monthly measured chlorophyll a (Cchl). The modelling results, which proved highly reliable during the calibration phase, correlated rather weakly with the annual PP estimates for the 18 years, which were based on 2-h incubations at midday, 1–2 times per month. Being based on continuous irradiance data, the modelled PP can be considered more reliable than the sparse measurements, especially for short to medium term studies. We demonstrate that in the long-term, the bio-optical method can be biased if changes in water colour or water level alter the light climate causing adaptive responses in the cellular chlorophyll content of light-limited phytoplankton.

Does differentversusequal daytime and night-time respiration matter for quantification of lake metabolism using diel dissolved oxygen cycles?

Diel dissolved oxygen (DO) measurements can be used to estimate water metabolism of aquatic systems, in particular, lakes, lagoons and streams in terms of gross primary production (GPP), ecosystem re- spiration (Reco) and net ecosystem production (NEP). One of the main assumptions in the calculation of lake metabolism is that Reco is the same for daytime (Rdaytime) and nighttime (Rdarkhr). This study aimed at testing the equal Rdaytime and Rdarkhr assumption to estimate GPP, Reco and NEP in a littoral zone of a temperate shallow lake (Lake Yenicag˘ a) in northwestern Turkey with and without the assumption. Based on the equal Rdarkhr and Rdaytime assumption, values calculated for GPP and Rdaytime were different than those based on the different Rdarkhr and Rdaytime assumption (P< 0.001). GPP was lower by 7.5% in July, 49.6% in September and 14.9% in October, while Reco was lower by 5.9% in July and 55.8% in September. GPP was higher by 8.9% in August and 55% in November, while Reco was higher by 7.8% in August and 23.9% in November.

Long-Term Changes in Nutrients and Phytoplankton Response in Lake Dianshan, a Shallow Temperate Lake in China

ABSTRACT In an effort to understand algal blooms in Dianshan Lake, a shallow temperate lake in China, we analyzed more than 20 years of nutrient data (total nitrogen [TN] and total phosphorus [TP]) and lake responses (chlorophyll a. [chl a], transparency, and algal composition). We found that 1) the trophic state of Dianshan Lake transitioned from eutrophic to hypereutrophic in 1999–2000, and nutrient accumulation has accelerated since then; 2) TP was the primary limiting factor for algal biomass, with the regression coefficients of TP and chl a varying with different trophic states; and 3) TN may limit cyanobacterial growth in autumn.

Filamentous green algae inhibit phytoplankton with enhanced effects when lakes get warmer

1. Filamentous green algae (FGA) may represent an alternative state in high-nutrient shallow temperate lakes. Furthermore, a clear water state is sometimes associated with the dominance of FGA; however, the mechanisms involved remain uncertain. 2. We hypothesised that FGA may promote a clear water state by directly suppressing phytoplankton growth, mostly via the release of allelochemicals, and that this interaction may be affected by temperature. 3. We examined the relationships between FGA, phytoplanktonic chlorophyll a concentrations and zooplankton in a series of mesocosms (2.8 m3) mimicking enriched shallow ponds now and in a future warmer climate (0 and c. 5 °C above ambient temperatures). We then tested the potential allelopathic effects of FGA (Cladophora sp. and Spirogyra sp.) on phytoplankton using several short-term microcosms and laboratory experiments. 4. Mesocosms with FGA evidenced lower phytoplanktonic chlorophyll a concentrations than those without. Zooplankton and zooplankton : phytoplankton biomass ratios did not differ between mesocosms with and without FGA, suggesting that grazing was not responsible for the negative effects on phytoplanktonic biomass (chlorophyll a). 5. Our field microcosm experiments demonstrated that FGA strongly suppressed the growth of natural phytoplankton at non-limiting nutrient conditions and regardless of phytoplankton initial concentrations or micronutrients addition. Furthermore, we found that the negative effect of FGA on phytoplankton growth increased up to 49% under high incubation temperatures. The experiment performed using FGA filtrates confirmed that the inhibitory effect of FGA on phytoplankton may be attributed to allelochemicals. 6. Our results suggest that FGA control of phytoplankton growth may be an important mechanism for stabilising clear water in shallow temperate lakes dominated by FGA and that FGA may play a larger role when lakes get warmer.

The role of macroinvertebrates and fish in regulating the provision by macrophytes of refugia for zooplankton in a warm temperate shallow lake

Summary1. The zooplankton often undergoes diel horizontal migration (DHM) from the open water to the littoral of shallow lakes, thus avoiding predators in the former. This behaviour has functional impacts within the lake, as it enhances zooplankton survival, increases their control of phytoplankton and tends to stabilise the clear water state. However, most of the evidence supporting this migration pattern comes from cold north temperate lakes, and more evidence from tropical and subtropical areas, as well as from southern temperate areas, is needed.2. We conducted a field study of the diel horizontal and vertical migration of zooplankton, and the horizontal distribution of potential predatory macroinvertebrates and fish, over two consecutive days in the summer in a temperate lake in the southern hemisphere. We took zooplankton samples at two depths, at three sampling stations (inside beds of aquatic macrophytes, at their edge and in open water) along three transects running from the centre of a bed of Ceratophyllum demersum to open water. At each sampling station, we also took samples of macroinvertebrates and fish and measured physical and chemical environmental variables.3. Zooplankton (pelagic cladocerans, calanoid copepods and rotifers) avoided the shore, probably because of the greater risk from predators there. Larger and more vulnerable cladocerans, such as Diaphanosoma brachyurum and Moina micrura, were two to four times more abundant in open water than at the edge of or inside beds of macrophytes, respectively, by both day and night. Less vulnerable zooplankton [i.e. of medium body size (Ceriodaphnia dubia) or with the ability to swim fast (calanoid copepods)] were distributed evenly between open water and the edge of the plant beds. Small zooplankton, Bosmina huaronensis and pelagic rotifers, showed an even distribution among the three sampling stations. Accordingly, no DHM of zooplankton occurred, although larger organisms migrated vertically inside C. demersum stands.4. Macrophytes contained high densities of predatory macroinvertebrates and fish. The predator assemblage, composed of large‐bodied macroinvertebrates (including odonates and shrimps) and small littoral fish, was permanently associated with submerged macrophytes. None of these groups moved outside the plant beds or changed their population structure (fish) over the diel cycle.5. Submerged macrophyte beds do not represent a refuge for zooplankton in lakes where predators are numerous among the plants, implying a weaker top‐down control of phytoplankton biomass by zooplankton and, consequently, a more turbid lake. The effectiveness of macrophytes as a refuge for zooplankton depends on the associated assemblage of predatory macroinvertebrates and fish among the plants.

Relationships between Fish Species Abundances and Water Transparency in Hypertrophic Turbid Waters of Temperate Shallow Lakes

AbstractWe explored the relationships between Secchi disc depth and the abundance of fish species in very shallow, hypertrophic, turbid waters of Pampa Plain lakes, Argentine.We tested whether the abundance of any of the species present was associated with water transparency for lakes where water transparency, as measured by Secchi disc depth, ranged from 0.1 to 0.4 m. Overall, the abundance of five species (Cnesterodon decemmaculatus, Jenynsia multidentata, Corydoras paleatus, Pimelodella laticeps and Odontesthes bonariensis) seemed to be affected by this narrow gradient in water transparency. These findings represent an interesting result for turbid hypertrophic environments where narrow ranges in water transparency are traditionally neglected as important factors for fishes. We show, however, how water transparency patterns may be still important for some species in highly turbid waters with extremely narrow gradients in Secchi disc depth. (© 2010 WILEY‐VCH Verlag GmbH &amp; Co. KGaA, Weinheim)

High-resolution spectroscopic study of pore-water dissolved organic matter in Holocene sediments of Lake Peipsi (Estonia/Russia)

Ultraviolet–visible absorption and fluorescence spectroscopies allow simultaneous detection of several organic substances. These spectra also contain information on whether the organic matter is produced in the water body or in its catchment. In this work various spectral indices, which are widely used in aquatic studies to determine humic substances, proteinaceous matter, chlorophyllous pigments and the origin (autochthonous versus allochthonous) of organic matter, were applied to track changes in a Holocene record from the large and shallow northern temperate Lake Peipsi (Estonia/Russia) at a high-resolution scale. Absorption and three-dimensional fluorescence spectra of the organic matter fraction dissolved in sediment pore water (pDOM) were analysed. In addition to the spectral measurements of pDOM, the sediment samples were analysed for main constituents (water and organic and mineral matter) and magnetic properties. The core chronology was established by nine radiocarbon datings. Temporal changes in pDOM in the record were quite variable; however, three periods in the development of the lake can be distinguished: a 2500 yr period in the early Holocene and a 2500 yr period in the late Holocene, when drastic changes in the accumulated matter occurred, and a rather stable 5000 yr period in between. We postulate that the changes over the period in the early Holocene reflect a rise of water level in the lake; the oscillations over the relatively steady period were probably caused by climatic factors, and alterations over the period in the late Holocene were due to the expansion of agricultural activity in the lake catchment. Our findings indicate that the use of spectral indices in palaeoinvestigations offers valuable information for reconstructing natural and human-induced developments of lakes.

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.

Ecological influences on larval chironomid communities in shallow lakes: implications for palaeolimnological interpretations

Summary1. To correctly interpret chironomid faunas for palaeoenvironmental reconstruction, it is essential that we improve our understanding of the relative influence of ecosystem variables, biotic as well as physicochemical, on chironomid larvae. To address this, we analysed the surface sediments from 39 shallow lakes (29 Norfolk, U.K., 10 Denmark) for chironomid head capsules, and 70 chironomid taxa (including Chaoborus) were identified.2. The shallow lakes were selected over large environmental gradients of aquatic macrophytes, total phosphorus (TP) and fish communities. Redundancy analysis (RDA) identified two significant variables that explained chironomid distribution: macrophyte species richness (P &lt; 0.001) and TP (P &lt; 0.005). Generalised linear models (GLM) identified specific taxa that had significant relationships with both these variables. Macrophyte percentage volume infested (PVI) and species richness were significant in classifying the lake types based on chironomid communities under twinspan analysis, although other factors, notably nutrient concentrations and fish communities, were also important, illustrating the complexities of classifying shallow lake ecosystems. Lakes with plant species richness &gt;10 all had relatively diverse (Hill’s N2) chironomid assemblages, and lakes with Hill’s N2 &gt;10 all had TP &lt;250 μg L−1 and total fish densities &lt;2 fish per m2.3. Plant density (PVI), and perhaps more importantly species richness, were primary controls on the distribution of chironomid communities within these lakes. This clearly has implications for palaeoenvironmental reconstructions using zoobenthos remains (i.e. chironomids) and suggests that they could be used to track changes in benthic/pelagic production and could be used as indicators of changing macrophyte habitat.4. Measuring key biological gradients, in addition to physicochemical gradients, allowed the major controls on chironomid distribution to be assessed more directly, in terms of plant substrate, food availability, competition and predation pressure, rather than implying indirect mechanisms through relationships with nutrients. Many of these variables, notably macrophyte abundance and species richness, are not routinely measured in such studies, despite their importance in determining zoobenthos in temperate shallow lakes.5. When physical, chemical and ecological gradients are considered, as is often the case with palaeo‐reconstructions rather than training sets chosen to maximise one gradient, complex relationships exist, and attempting to reconstruct a single trophic variable quantitatively may not be appropriate or reliable.

The response of periphyton and submerged macrophytes to nitrogen and phosphorus loading in shallow warm lakes: a mesocosm experiment

Summary1. Recent experimental and field studies on temperate shallow lakes indicate that nitrogen may play a greater role in their functioning than previously thought. Several studies document that abundance and richness of submerged macrophytes, both central in shallow lake ecology, may decrease with increasing nitrogen loading, especially at high phosphorus levels. However, the role of nitrogen in warm lakes with fluctuating water regimes remains to be described in detail.2. The effect of increasing nitrate and phosphate concentrations on submerged macrophyte growth was examined in a 3‐month mesocosm experiment conducted in summer in a shallow freshwater lake on the north western coast of Turkey with a Mediterranean climate. Twenty four field mesocosms, open to the sediment and atmosphere, were stocked with Myriophyllum spicatum shoots and small cyprinid fish. Three nitrate loadings in combination with two phosphate loadings were applied in a fourfold replicated design.3. Mean ± SD nutrient concentrations maintained throughout the experiment were 0.55 ± 0.17, 2.2 ± 0.97, 9.2 ± 5.45 mg L−1 total nitrogen and 55 ± 19.2, 73 ± 22.9 μg L−1 total phosphorus. Mean periphyton biomass increased with increasing nutrient concentrations and peaked at the highest nitrogen and phosphorus loadings, while the mean phytoplankton biomass remained relatively low in all treatments.4. Percent volume inhabited (% PVI) by macrophytes throughout the experiment and total macrophyte biomass at the end of the experiment did not differ among treatments. In addition to stocked M. spicatum, Ceratophyllum demersum and Potamogeton crispus appeared in the majority of the mesocosms. The plants grew continuously up to 50% PVI throughout the experiment and remained resilient to shading provided by periphyton and phytoplankton.5. The mean summer air temperature in 2007 was 2.2 °C higher than the average of the last 32 years, which resulted in a water level decrease of 0.3 m in the mesocosms over three months. This might have counteracted the shading of submerged macrophytes provided by phytoplankton and periphyton. The results of the experiment are consistent with observations of higher macrophyte resilience to nutrient loading in Mediterranean lakes compared with northern temperate lakes.

Determinants of biodiversity in subtropical shallow lakes (Atlantic coast, Uruguay)

Summary1. Shallow lakes and ponds contribute disproportionally to species richness relative to other aquatic ecosystems. In‐lake conditions (e.g. presence of submerged plants) seem to play a key role in determining diversity, as has been demonstrated for temperate lakes. When water quality deteriorates and turbidity increases, conditions in such lakes are affected drastically resulting in a loss of diversity. However, it is not clear whether subtropical lakes show the same pattern and whether the richness of all groups reacts similarly to environmental changes.2. Our aim was to analyse the main factors explaining patterns of species richness in plankton, fish and submerged macrophyte assemblages in both turbid and clear subtropical shallow lakes. We analysed abiotic and biotic features of 18 subtropical, small‐ to medium‐sized, shallow lakes along the Uruguayan coast. We compared both turbid and clear ecosystem states and evaluated the relative variance explained by the factors measured.3. Variables describing lake and catchment morphology, as well as the percentage of the water column occupied by submerged macrophytes (%PVI) and water turbidity, had strong effects on taxon richness. Interestingly, individual biotic groups had dissimilar richness patterns. Macrophyte %PVI decreased with increasing lake area, while fish species richness showed the opposite pattern. Phytoplankton species richness increased with macrophyte %PVI, while the zooplankton richness pattern varied depending on the taxonomic group considered.4. Overall, our results indicate that, as found for temperate lakes, a greater submerged plant cover promotes higher species richness in several groups, and that this may overwhelm the otherwise expected positive effect of lake size on species richness. On the other hand, small‐bodied zooplankton predominated in lakes with high plant abundance. Our findings concur with recent studies, indicating that refuge capacity of aquatic plants might be weaker in (sub)tropical than in temperate shallow lakes.5. The extremely high plant cover, frequently observed in warm lakes, could potentially lead to different richness patterns in some groups. This conclusion has important consequences for local managers and conservationists.

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.

Substantial differences in littoral fish community structure and dynamics in subtropical and temperate shallow lakes

Summary1. Fish play a key role in the functioning of temperate shallow lakes by affecting nutrient exchange among habitats as well as lake trophic structure and dynamics. These processes are, in turn, strongly influenced by the abundance of submerged macrophytes, because piscivorous fish are often abundant at high macrophyte density. Whether this applies to warmer climates as well is virtually unknown.2. To compare fish community structure and dynamics in plant beds between subtropical and temperate shallow lakes we conducted experiments with artificial submerged and free‐floating plant beds in a set of 10 shallow lakes in Uruguay (30°–35°S) and Denmark (55°–57°N), paired along a gradient of limnological characteristics.3. The differences between regions were more pronounced than differences attributable to trophic state. The subtropical littoral fish communities were characterised by higher species richness, higher densities, higher biomass, higher trophic diversity (with predominance of omnivores and lack of true piscivores) and smaller body size than in the comparable temperate lakes. On average, fish densities were 93 ind. m−2 (±10 SE) in the subtropical and 10 ind. m−2 (±2 SE) in the temperate lakes. We found a twofold higher total fish biomass per unit of total phosphorus in the subtropical than in the temperate lakes, and as fish size is smaller in the former, the implication is that more energy reaches the littoral zone fish community of the warmer lakes.4. Plant architecture affected the spatial distribution of fish within each climate zone. Thus, in the temperate zone fish exhibited higher densities among the artificial free‐floating plants while subtropical fish were denser in the artificial submerged plant beds. These patterns appeared in most lakes, regardless of water turbidity or trophic state.5. The subtropical littoral fish communities resembled the fish communities typically occurring in temperate eutrophic and hypertrophic lakes. Our results add to the growing evidence that climate warming may lead to more complex and omnivory‐dominated food webs and higher density and dominance of smaller‐sized fish. This type of community structure may lead to a weakening of the trophic cascading effects commonly observed in temperate shallow lakes and a higher risk of eutrophication.

Experimental evidence of the dynamic effect of free‐floating plants on phytoplankton ecology

Summary1. There is an increasing concern to understand the role of free‐floating plants (FFP) on the functioning and structure of shallow lakes, particularly the mechanisms by which their dominance is self‐stabilizing and how they may outcompete phytoplankton.2. In a field experiment with mesocosms, we simulated three commonly encountered scenarios in warm temperate shallow lakes: FFP dominance, FFP fluctuation and FFP absence. We explored the effects of several key processes, triggered by FFP dynamics, on the composition, diversity and production of phytoplankton, and on the physicochemical conditions.3. The effects of persistent floating mats on light interception triggered a complex response by the phytoplankton: species were lost and biomass was low, yet high diversity and productivity potential were maintained. A contrasting phytoplankton response characterized mesocosms lacking FFP, where light was sufficient but nitrogen was limiting. Fluctuating FFP cover brought periodic shifts between these two limiting resources for the phytoplankton, which most probably explain the maintenance of richness, diversity and production in these shallow lakes.4. These results support the recently proposed framework of alternative stable states, providing experimental field evidence of the mechanisms resulting from the shifts between floating plants and phytoplankton dominance.

Seasonal dynamics of phytoplankton in two Amazon flood plain lakes of varying hydrologic connectivity to the main river channel

Seasonal dynamics of phytoplankton were evaluated in Lakes Amapa and Pirapora, small oxbows in the floodplain of Rio Acre, a tributary of the Amazon River in western Brazil. The two lakes differ primarily in their degree of connectivity to the main river channel, with the younger Lake Pirapora maintaining a connection year-round and displaying physical and chemical characteristics responding to rising river water levels, versus the older Lake Amapa connecting to the river only during a 2-3 month flooding phase and having a distinct period of isolation at the end of the dry season. The phytoplankton of Lake Pirapora has a low numeric density year-round and is dominated by small C-strategists (cryptomonads). The phytoplankton of Lake Amapa has a similar low density when the lake is connected to the river. However, once that lake is isolated and turbidity declines, density increases an order of magnitude and the assemblage becomes dominated by filamentous N-fixing Aphanizomenon tropicalis. Coincidentally concentrations of dissolved oxygen increase and levels of TN decline, similar to changes during autogenic succession in temperate shallow lakes. The seasonal changes in phytoplankton of Lake Amapa, from the time it is connected to the time when it is isolated from the river, generally conform to the C-S-R conceptual model of succession. This is the first study to directly compare lakes of different developmental stages, in terms of connectivity with the river channel. There remains a need for more comprehensive long-term comparisons of these unique lake ecosystems.

Effects of increased temperature and nutrient enrichment on the stoichiometry of primary producers and consumers in temperate shallow lakes

Summary1. We studied the effects of increased water temperatures (0–4.5 °C) and nutrient enrichment on the stoichiometric composition of different primary producers (macrophytes, epiphytes, seston and sediment biofilm) and invertebrate consumers in 24 mesocosm ecosystems created to mimic shallow pond environments. The nutrient ratios of primary producers were used as indicative of relative nitrogen (N) or phosphorus (P) limitation. We further used carbon stable isotopic composition (δ13C) of the different primary producers to elucidate differences in the degree of CO2 limitation.2. Epiphytes were the only primary producer with significantly higher δ13C in the enriched mesocosms. No temperature effects were observed in δ13C composition of any primary producer. Independently of the treatment effects, the four primary producers had different δ13C signatures indicative of differences in CO2 limitation. Seston had signatures indicating negligible or low CO2 limitation, followed by epiphytes and sediment biofilm, with moderate CO2 limitation, while macrophytes showed the strongest CO2 limitation. CO2 together with biomass of epiphytes were the key variables explaining between 50 and 70% of the variability in δ13C of the different primary producers, suggesting that epiphytes play an important role in carbon flow of temperate shallow lakes.3. The ratio of carbon to chlorophyll a decreased with increasing temperature and enrichment in both epiphytes and seston. The effects of temperature were mainly attributed to changes in algal Chl a content, while the decrease with enrichment was probably a result of a higher proportion of algae in the seston and epiphytes.4. Macrophytes, epiphytes and seston decreased their C : N with enrichment, probably as an adaptation to the different N availability levels. The C : N of epiphytes and Elodea canadensis decreased with increasing temperature in the control mesocosms. Sediment biofilm was the only primary producer with lower C : P and N : P with enrichment, probably as a result of higher P accumulation in the sediment.5. Independently of nutrient level and increased temperature effects the four primary producers had significantly different stoichiometric compositions. Macrophytes had higher C : N and C : P and, together with epiphytes, also the highest N : P. Seston had no N or P limitation, while macrophytes and epiphytes may have been P limited in a few mesocosms. Sediment biofilm indicated strong N deficiency.6. Consumers had strongly homeostatic stoichiometric compositions in comparison to primary producers, with weak or no significant treatment effects in any of the groups (insects, leeches, molluscs and crustaceans). Among consumers, predators had significantly higher N content and lower C : N than grazers.