Shallow Lake Food Webs Research Articles

Small-sized omnivorous fish induce stronger effects on food webs than warming and eutrophication in experimental shallow lakes

Warming, eutrophication, and increased omnivory by small-sized fish are global change processes that induce major effects on the food web structure and primary producers of shallow lakes. Despite the key relevance of phytoplankton and periphyton in freshwaters, the combined and potential synergistic effects of fish omnivory, warming and eutrophication, especially on periphyton, remains little addressed, particularly for subtropical shallow lakes. We experimentally tested the food web effects on phytoplankton and periphyton induced by small visually feeding omnivorous fish (Rhodeus ocellatus), high nutrient enrichment and warming (+4.5 °C) in thirty-two 1000 L-mesocosms simulating littoral conditions of subtropical shallow lakes. We aimed at analysing the mechanisms and responses of periphyton and phytoplankton to these experimental factors. All mesocosms included the submerged macrophytes Vallisneria denseserrulata and Potamogeton lucens and artificial plants at 50% plant volume inhabited, plankton and macroinvertebrates. Small-sized visually feeding omnivorous fish enhanced phytoplankton dominance and periphyton loss. These changes coincided with a decrease in zooplankton biomass and a diversity loss of both zooplankton and macroinvertebrates as well as an increase in snail abundance. Fish presence led to a collapse of cladocerans, thereby releasing the grazing pressure on phytoplankton, and predator and collector macroinvertebrates were replaced with small snails (Radix peregra < 0.5 cm) resulting in enhanced grazing on periphyton. Eutrophication reinforced the fish effects, while warming had weak or no effects. Our results indicate that omnivory by small-sized visually feeding fish may induce stronger effects on the food webs of shallow lakes, towards phytoplankton-dominated states, than the combined effect of nutrient enrichment and warming under the present experimental conditions.

Subfossil cladocerans as quantitative indicators of past ecological conditions in Yangtze River Basin lakes, China

Cladocerans (Crustacea: Branchiopoda) are an essential component of the food webs of shallow lakes. However, their potential significance as sentinels of environmental change in subtropical regions of South-East Asia, is not well elucidated. In this study, we examined the distribution pattern of cladoceran subfossils in the surface sediments of 64 shallow lakes located in the middle and lower reaches of the Yangtze River Basin (MLYB). Using constrained ordination and multivariate regression trees (MRT), we explored the relationships between the distribution of cladocerans and contemporary environmental variables. The results indicated that macrophyte abundance is the most important regulator, followed by chlorophyll a and total phosphorus that all were important drivers of cladoceran community dynamics. Despite intensive aquaculture in the MLYB lakes over the past decades, fish predation did not have a significant influence on the cladoceran community, likely because the fish predation pressure was generally high. The MRT analysis showed distinct variations in the composition of cladoceran communities and was able to identify the key indicator species of various stressors, including Bosmina coregoni, small Alona (A. intermedia, A. rectangula) and Leydigia acanthocercoides, Leydigia leydigi and Chydorus sphaericus. Boosted Regression Tree (BRT), a novel numerical model, was employed to establish the quantitative relationship between cladoceran composition and environmental variables. The model was used to reconstruct historical macrophyte coverage over the past 60 years in a typical Yangtze lake, Liangzi Lake, based on sedimentary cladoceran records. The BRT-inferred macrophyte coverage was validated well by the macrofossil records in the same lake. Using subfossil cladocerans as indicator of the past environmental change in MLYB lakes is potentially useful given their key ecological roles. However, caution should be taken due to species-specific differences in preservation in the sediment and confounding environmental processes that to some extent may bias the conclusions.

Modeling Decreased Resilience of Shallow Lake Ecosystems toward Eutrophication due to Microplastic Ingestion across the Food Web.

The discovery of microplastic (MP) being present in freshwaters has stimulated research on the impacts of MP on freshwater organisms. To date, research has focused on primary effects, leaving questions with respect to secondary effects at the level of freshwater food webs unanswered. Here, we use a theoretical modeling approach to investigate the hypothesis that MP imposes negative impacts on the level of freshwater shallow lake food webs. We find that increasing MP levels have the potential to affect the critical phosphorus loading (CPL), which is defined as the threshold for regime shifts between clear and turbid states of the water column. The possible occurrence of catastrophic cascades due to MP pollution is predominantly driven by the negative effects of MP on zooplankton. We explore the possible states of the food web by scenario analysis and show that the secondary effects of MP at current concentrations are likely to be negligible. However, at the current rate of MP production, a 20-40% reduction in the CPL would occur by the end of this century, suggesting a loss of resilience in shallow lakes that would be subject to abrupt changes in the food web under lower nutrient loading.

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.

Characteristics of consumer trophic resources for Waikato shallow lake food webs

ABSTRACTWe characterised the quantity (mass per unit area or volume), quality (cabon:nitrogen ratio) and enrichment (δ15N) of basal food resources and benthic macroinvertebrates in six shallow lakes in the Waikato region, New Zealand, to elucidate patterns related to lake type (peat, riverine), habitat (littoral, mid-lake) and enrichment status (Trophic Level Index >6 or ≤5). Total seston dry mass was dominated by fine material that was correlated with chlorophyll a concentration. Seston was a higher quality food resource than benthic particulate organic matter which had higher biomass overall in littoral than in mid-lake habitats. Biomass of Chironomidae was highest in the less-enriched peat lakes, possibly due to clearer water supporting benthic algae in shallow littoral areas. Our results highlight the potential role of multiple bottom-up influences on shallow lake food webs. The more abundant and higher quality pelagic food resources in highly enriched lakes may have implications for resilience to non-indigenous fish invasion.

Stable isotope analysis confirms substantial differences between subtropical and temperate shallow lake food webs

Differences in trophic web structure in otherwise similar ecosystems as a consequence of direct or indirect effects of ambient temperature differences can lead to changes in ecosystem functioning. Based on nitrogen and carbon stable isotope analysis, we compared the food-web structure in a series of subtropical (Uruguay, 30–35°S) and temperate (Denmark, 55–57°N) shallow lakes. The food-web length was on average one trophic position shorter in the subtropical shallow lakes compared with their temperate counterparts. This may reflect the fact that the large majority of subtropical fish species are omnivores (i.e., feed on more than one trophic level) and have a strong degree of feeding niche overlap. The shapes of the food webs of the subtropical lakes (truncated and trapezoidal) suggest that they are fuelled by a combination of different energy pathways. In contrast, temperate lake food webs tended to be more triangular, likely as a result of more simple pathways with a top predator integrating different carbon sources. The effects of such differences on ecosystem functioning and stability, and the connection with ambient temperature as a major underlying factor, are, however, still incipiently known.

Whole‐lake experiments reveal the fate of terrestrial particulate organic carbon in benthic food webs of shallow lakes

Lake ecosystems are strongly linked to their terrestrial surroundings by material and energy fluxes across ecosystem boundaries. However, the contribution of terrestrial particulate organic carbon (tPOC) from annual leaf fall to lake food webs has not yet been adequately traced and quantified. In this study, we conducted whole-lake experiments to trace artificially added tPOC through the food webs of two shallow lakes of similar eutrophic status, but featuring alternative stable regimes (macrophyte rich vs. phytoplankton dominated). Lakes were divided with a curtain, and maize (Zea mays) leaves were added, as an isotopically distinct tPOC source, into one half of each lake. To estimate the balance between autochthonous carbon fixation and allochthonous carbon input, primary production and tPOC and tDOC (terrestrial dissolved organic carbon) influx were calculated for the treatment sides. We measured the stable isotope ratios of carbon (delta13C) of about 800 samples from all trophic consumer levels and compared them between lake sides, lakes, and three seasons. Leaf litter bag experiments showed that added maize leaves were processed at rates similar to those observed for leaves from shoreline plants, supporting the suitability of maize leaves as a tracer. The lake-wide carbon influx estimates confirmed that autochthonous carbon fixation by primary production was the dominant carbon source for consumers in the lakes. Nevertheless, carbon isotope values of benthic macroinvertebrates were significantly higher with maize additions compared to the reference side of each lake. Carbon isotope values of omnivorous and piscivorous fish were significantly affected by maize additions only in the macrophyte-dominated lake and delta13C of zooplankton and planktivorous fish remained unaffected in both lakes. In summary, our results experimentally demonstrate that tPOC in form of autumnal litterfall is rapidly processed during the subsequent months in the food web of shallow lakes and is channeled to secondary and tertiary consumers predominantly via the benthic pathways. A more intense processing of tPOC seems to be connected to a higher structural complexity in littoral zones, and hence may differ between shallow lakes of alternative stable states.

Bimodality in stable isotope composition facilitates the tracing of carbon transfer from macrophytes to higher trophic levels

Even though the suitability of macrophytes to act as a carbon source to food webs has been questioned by some studies, some others indicate that macrophyte-derived carbon may play an important role in the trophic transfer of organic matter in the food web of shallow lakes. To evaluate the importance of macrophytes to food webs, we collected primary producers—macrophytes and periphyton—and consumers from 19 South American shallow lakes and analyzed their carbon stable isotopes composition (δ13C). Despite the diversity of inorganic carbon sources available in our study lakes, the macrophytes’ δ13C signatures showed a clear bimodal distribution: 13C-depleted and 13C-enriched, averaging at −27.2 and −13.5‰, respectively. We argue that the use of either CO2 or HCO3− by the macrophytes largely caused the bimodal pattern in δ13C signals. The contribution of carbon from macrophytes to the lake’s food webs was not straightforward in most of the lakes because the macrophytes’ isotopic composition was quite similar to the isotopic composition of periphyton, phytoplankton, and terrestrial carbon. However, in some lakes where the macrophytes had a distinct isotopic signature, our data suggest that macrophytes can represent an important carbon source to shallow lake food webs.

Back to the future: using palaeolimnology to infer long‐term changes in shallow lake food webs

Summary1. Shallow lakes are often cited as classic examples of systems that exhibit trophic cascades but, whilst they provide good model systems with which to test general ecological theory and to assess long‐term community change, their food web linkages have rarely been resolved, so changes associated with the structure and dynamics of the ecological network as a whole are still poorly understood.2. We sought to redress this, and to demonstrate the potential benefits of integrating palaeolimnological and contemporary data, by constructing highly resolved food webs and stable isotope derived measures of trophic interactions and niche space, for the extant communities of two shallow U.K. lakes from different positions along a gradient of eutrophication. The contemporary surface sediment cladoceran and submerged macrophyte assemblages in the less enriched site, Selbrigg Pond, matched the palaeolimnological assemblages of the more enriched site, Felbrigg Hall Lake, in its more pristine state during the 1920s. Thus, Selbrigg was a temporal analogue for Felbrigg, from which the consequences of long‐term eutrophication on food web structure could be inferred. These data represent the first steps towards reconstructing not only past assemblages (i.e. nodes within a food web), but also past interactions (i.e. links within a food web): a significant departure from much of the previous research in palaeolimnology.3. The more eutrophic food web had far fewer nodes and links, and thus a less reticulate network, than was the case for the more pristine system. In isotopic terms, there was vertical compression in δ15N range (NR) and subsequent increased overlap in isotopic niche space, indicating increased trophic redundancy within Felbrigg. This structural change, which was associated with a greater channelling of energy through a smaller number of nodes as alternative feeding pathways disappear, could lead to reduced dynamic stability, pushing the network towards further simplification. These changes reflected a general shift from a benthic‐dominated towards a more pelagic system, as the plant‐associated subweb eroded.4. Although these data are among the first of their kind, the palaeo‐analogue approach used here demonstrates the huge potential for applying food web theory to understand how and why these ecological networks change during eutrophication. Furthermore, because of the rich biological record preserved in their sediments, shallow lakes represent potentially important models for examining long‐term intergenerational dynamics, thereby providing a means by which models and data can be integrated on meaningful timescales – a goal that has long proved elusive in food web ecology.

Mesocosm experiments on nutrient and fish effects on shallow lake food webs in a Mediterranean climate

Summary1. Nutrient and fish manipulations in mesocosms were carried out on food‐web interactions in a Mediterranean shallow lake in south‐east Spain. Nutrients controlled biomass of phytoplankton and periphyton, while zooplankton, regulated by planktivorous fish, influenced the relative percentages of the dominant phytoplankton species.2. Phytoplankton species diversity decreased with increasing nutrient concentration and planktivorous fish density. Cyanobacteria grew well in both turbid and clear‐water states.3. Planktivorous fish increased concentrations of soluble reactive phosphorus (SRP). Larger zooplankters (mostly Ceriodaphnia and copepods) were significantly reduced when fish were present, whereas rotifers increased, after fish removal of cyclopoid predators and other filter feeders (cladocerans, nauplii). The greatest biomass and diversity of zooplankton was found at intermediate nutrient levels, in mesocosms without fish and in the presence of macrophytes.4. Water level decrease improved underwater light conditions and favoured macrophyte persistence. Submerged macrophytes (Chara spp.) outcompeted algae up to an experimental nutrient loading equivalent to added concentrations of 0.06 mg L−1 PO4‐P and 0.6 mg L−1 NO3‐N, above which an exponential increase in periphyton biomass and algal turbidity caused characean biomass to decline.5. Declining water levels during summer favoured plant‐associated rotifer species and chroococcal cyanobacteria. High densities of chroococcal cyanobacteria were related to intermediate nutrient enrichment and the presence of small zooplankton taxa, while filamentous cyanobacteria were relatively more abundant in fishless mesocosms, in which Crustacea were more abundant, and favoured by dim underwater light.6. Benthic macroinvertebrates increased significantly at intermediate nutrient levels but there was no relationship with planktivorous fish density.7. The thresholds of nutrient loading and in‐lake P required to avoid a turbid state and maintain submerged macrophytes were lower than those reported from temperate shallow lakes. Mediterranean shallow lakes may remain turbid with little control of zooplankton on algal biomass, as observed in tropical and subtropical lakes. Nutrient loading control and macrophyte conservation appear to be especially important in these systems to maintain high water quality.

Environmental quality criteria for organic chemicals predicted from internal effect concentrations and a food web model

Environmental quality criteria (EQC) for hydrophobic organic chemicals were calculated with a model for bioaccumulation in food webs. The model was calibrated and verified using polychlorinated biphenyl concentrations in food webs of shallow lakes. The EQCs in water and sediment were derived based on internal effect concentrations (IECs) for several modes of toxic action. By reverse calculation with the food web model for each organism in the web, a different water or sediment concentration is calculated corresponding to the IEC in each organism. A statistical procedure with an acute-to-chronic value is used to derive chronic EQCs based on bioaccumulation. The model-based chronic EQCs were compared with previously established EQCs. The EQCs calculated with the food web model generally are within an order of magnitude of the previously derived EQCs based on toxicity data on individual chemicals. Some previously derived EQCs are much lower than model predictions and usually based on small samples of toxicity data such as no-observed-effect concentrations (NOECs) with large assessment factors. When faced with data gaps, it is proposed to use model-based chronic EQCs for (polar) narcotic chemicals. Other modes of action require a different model concept to account for receptor-based toxicity.