Piscivore enhancement effects on food webs depend on planktivore body size and species composition in replicated whole lake experiments

Recent syntheses of trophic cascade and biomanipulation research have suggested that the effects of piscivores on planktivorous fish populations are reduced, when planktivores are capable of outgrowing predator gape limitation and in systems with complex food web interactions. These hypotheses, however, have not been tested in long-term, whole-lake, experiments where processes such as fish recruitment and compensatory food web responses may be important. We conducted a replicated whole-lake experiment to test for the effects of supplemental piscivore introductions on food webs of eutrophic lakes dominated by deep-bodied planktivores. Responses to piscivore enhancement were compared between lakes differing in food web structure due to the presence of omnivorous gizzard shad (Dorosoma cepedianum). A significant decrease in the relative abundance of juvenile planktivorous fish, and an increase in total benthic macroinvertebrate density was observed in lakes containing mainly bluegills (Lepomus machrochirus). In contrast, lakes containing gizzard shad exhibited no significant responses to piscivore manipulation. Our results support the hypothesis that food webs in lakes dominated by deep-bodied planktivorous fish species respond weakly to piscivore enhancement. In addition, our findings support the hypothesis that cascading trophic interactions are weaker in lake ecosystems with more complex food web interactions such as those containing gizzard shad.     

The influence of productivity and width of littoral zone on the trophic position of a large-bodied omnivore

Omnivory is common in many food webs. Omnivores in different habitats can potentially change their feeding behaviour and alter their trophic position and role according to habitat conditions. Here we examine the trophic level and diet of the omnivorous signal crayfish (Pacifastacus leniusculus) in gradients of trophic status and lake size, both of which have been previously suggested to affect trophic position of predators separately or combined as productive space. We found the trophic position of omnivorous crayfish to be positively correlated with lake trophic status, but found no evidence for any influence of lake size or productive space on crayfish trophic position. The higher trophic position of crayfish in eutrophic lakes was largely caused by a shift in crayfish diet and not by an increase in trophic links in basal parts of the food web. Hence, our results support the “productivity hypothesis,” suggesting that food chains can be longer in more productive systems. Furthermore, stable isotope data indicated that larger crayfish are more predatory than smaller crayfish in lakes with wider littoral zones. Wider littoral zones promoted the development of intrapopulation differences in trophic position whereas narrow littoral zones did not. Hence, differences in habitat quality between and within lakes seem to influence the trophic positions of omnivorous crayfish. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Food Web Topology in High Mountain Lakes

Although diversity and limnology of alpine lake systems are well studied, their food web structure and properties have rarely been addressed. Here, the topological food webs of three high mountain lakes in Central Spain were examined. We first addressed the pelagic networks of the lakes, and then we explored how food web topology changed when benthic biota was included to establish complete trophic networks. We conducted a literature search to compare our alpine lacustrine food webs and their structural metrics with those of 18 published lentic webs using a meta-analytic approach. The comparison revealed that the food webs in alpine lakes are relatively simple, in terms of structural network properties (linkage density and connectance), in comparison with lowland lakes, but no great differences were found among pelagic networks. The studied high mountain food webs were dominated by a high proportion of omnivores and species at intermediate trophic levels. Omnivores can exploit resources at multiple trophic levels, and this characteristic might reduce competition among interacting species. Accordingly, the trophic overlap, measured as trophic similarity, was very low in all three systems. Thus, these alpine networks are characterized by many omnivorous consumers with numerous prey species and few consumers with a single or few prey and with low competitive interactions among species. The present study emphasizes the ecological significance of omnivores in high mountain lakes as promoters of network stability and as central players in energy flow pathways via food partitioning and enabling energy mobility among trophic levels.     

Effects of Multi-chain Omnivory on the Strength of Trophic Control in Lakes

Omnivory has been implicated in both diffusing and intensifying the effects of consumer control in food chains. Some have postulated that the strong, community level, top-down control apparent in lakes is not expressed in terrestrial systems because terrestrial food webs are reticulate, with high degrees of omnivory and diverse plant communities. In contrast, lake food webs are depicted as simple linear chains based on phytoplankton-derived energy. Here, we explore the dynamic implications of recent evidence showing that attached algal (periphyton) carbon contributes substantially to lake primary and secondary productivity, including fish production. Periphyton production represents a cryptic energy source in oligotrophic and mesotrophic lakes that is overlooked by previous theoretical treatment of trophic control in lakes. Literature data demonstrate that many fish are multi-chain omnivores, exploiting food chains based on both littoral and pelagic primary producers. Using consumer-resource models, we examine how multiple food chains affect fourth-level trophic control across nutrient gradients in lakes. The models predict that the stabilizing effects of linked food chains are strongest in lakes where both phytoplankton and periphyton contribute substantially to production of higher trophic levels. This stabilization enables a strong and persistent top down control on the pelagic food chain in mesotrophic lakes. The extension of classical trophic cascade theory to incorporate more complex food web structures driven by multi-chain predators provides a conceptual framework for analysis of reticulate food webs in ecosystems.     

Modification of the diel vertical migration of Bythotrephes longimanus by the cold-water planktivore, Coregonus artedi

1. The weak diel vertical migration observed in the large cladoceran Bythotrephes longimanus seems contradictory to the predator‐avoidance hypothesis that predicts large zooplankton should have long migration amplitudes. However, cold‐water planktivores, especially Coregonus spp., are a main source of mortality for Bythotrephes and hence a deeper migration would result in a greater overlap with these hypolimnetic planktivores. We hypothesized that Coregonus artedi (cisco) modifies the normal vertical migration pattern of Bythotrephes, such that the latter stays higher in the water column during the day and thus migrates less extremely at night. 2. The vertical distribution of Bythotrephes during the day was determined from single visits to six lakes in Ontario, Canada, all of which contain warm‐water, epilimnetic planktivores but differing in whether they contain cisco. One lake of each fish type was sampled day and night every 2–3 weeks over the ice‐free season to examine daytime depths and migration amplitude. 3. The vertical migration of Bythotrephes differed in the presence and absence of cisco. In the lakes with cisco, there were significantly fewer Bythotrephes in the hypolimnion and they were higher in the water column during the day. Migration amplitude was smaller in the cisco than in the non‐cisco lake. These observations were not attributable to differences in physical factors, and, although not conclusively attributable to cisco, are consistent with an effect of cisco. 4. We suggest that diurnal depth selection by Bythotrephes in lakes containing cisco is a trade‐off between the risk of predation by warm‐ versus cold‐water predators, balanced by the benefits of increased temperature and feeding rates near the surface. Even in lakes without cisco, however, the vertical migration of Bythotrephes was less than expected, suggesting that diurnal depth selection is a balance between the risk from warm‐water planktivores and access to sufficient light to feed effectively.     

Cascading effects of generalist fish introduction in oligotrophic lakes

Introduction of strictly planktivorous fish to lakes can alter plankton communities via cascading interactions in food webs. Less is known about the large-scale and long-term effects resulting from the introduction of fish with more generalist feeding habits, and the extent to which these effects depend on lake trophic status. Paleolimnological records of three oligotrophic lakes in Maine, USA were used to analyze the response of plankton communities to the introduction of white perch (Morone americana), a fish that often numerically dominates fish assemblages and switches from strict planktivory to a more generalist diet during ontogeny. After white perch introduction, cladoceran ephippia size increased up to 50 %, suggesting that the most important role of this generalist fish, with respect to water quality, is as a piscivorous trophic link. Algal standing crop declined by a quarter to over a half of pre-introduction levels, suggesting that top-down effects of white perch reduced phytoplankton biomass. In comparing these oligotrophic lakes to prior work in a eutrophic system, white perch introduction had similar effects on zooplankton body size; however, cascading effects to phytoplankton were only observed in low productivity lakes.     

Differences in food webs and trophic states of Brazilian tropical humid and semi-arid shallow lakes: implications of climate change

Global warming may intensify eutrophication of shallow lakes by affecting nutrient loading, evaporation rates, and water level and thus produce major changes in food webs. We investigated to what degree food webs in tropical humid lakes differed from those in more eutrophic semi-arid lakes of the same latitude. Our results indicate that the catchment area-to-lake area ratio, nutrients, chlorophyll a, suspended solids, abundances of phytoplankton, zooplankton, and omnivorous fish as well as total fish catch per unit effort were all higher in the semi-arid lakes, whereas inlet water-to-evaporation ratio (proxy for water balance), water transparency, percentage macrophytes cover, and the piscivores:omnivores ratio were higher in the humid lakes. Our results suggest that reduced inlet water-to-evaporation ratio will increase lake eutrophication, which, in turn, as in temperate regions, will alter trophic structure of the freshwater community.

     

Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food‐web stability. In lakes, littoral and pelagic food‐web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high‐latitude lakes. We analyzed food‐web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km²) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food‐chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate‐dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high‐latitude lakes.     

A stable carbon and nitrogen isotope study of lake food webs in Canada's Boreal Plain

1.  Stable carbon and nitrogen isotope and fish stomach content analyses were used to investigate food webs in five relatively undisturbed lakes on the Boreal Plain of Canada. Stable isotope analysis was also used to determine the importance of external and internal carbon sources.
2.  Overlap in the carbon and nitrogen signatures of primary producers made it difficult to determine unambiguously the feeding habits of many invertebrates. However, isotope analysis suggested that external carbon inputs were detectable in the aquatic food chains of the one lake with a short water residence time («1 year). In the other four lakes, with water residence times ≥1 year, autochthonous carbon was the only detectable carbon source in the food webs.
3.  Food webs in these lakes spanned a range of four to five trophic levels. Both invertebrates and fish appeared to eat a variety of food, often feeding at more than one trophic level.
4.  With the exception of one lake (SPH20), top predators in these lakes, northern pike ( Esox lucius ) and fathead minnows ( Pimephales promelas ), occupied similar trophic positions despite large differences in body size and trophic morphology. In SPH20, where there were two additional fish species, pike occupied a higher trophic position. However, all the top predators in each lake appeared to be omnivores and generalists.
5.  The prevalence of omnivory and the apparent generalist feeding habits of fish in these lakes suggest that organisms are flexible in their feeding habits and that these food webs will be resilient to disturbance.     

The mixotroph Ochromonas tuberculata may invade and suppress specialist phago- and phototroph plankton communities depending on nutrient conditions

Mixotrophic organisms combine light, mineral nutrients, and prey as supplementary resources. Based on theoretical assumptions and field observations, we tested experimentally the hypothesis that mixotrophs may invade established plankton communities depending on the trophic status of the system, and investigated possible effects on food web structure, species diversity, and nutrient dynamics. To test our hypothesis, we inoculated the mixotrophic nanoflagellate Ochromonas tuberculata into established planktonic food webs, consisting of specialist phototrophs, specialist phagotrophs, and bacteria at different supplies of soluble inorganic nutrients and dissolved organic carbon. Oligotrophic systems facilitated the invasion of O. tuberculata in two different ways. First, the combination of photosynthesis and phagotrophy gave mixotrophs a competitive advantage over specialist phototrophs and specialist phagotrophs. Second, low nutrient supplies supported the growth of small plankton organisms that fell into the food size spectrum of mixotrophs. Conversely, high nutrient supplies prevented O. tuberculata from successfully invading the food webs. Two important conclusions were derived from our experiments. First, in contrast to a paradigm of ecology, specialization may not necessarily be the most successful strategy for survival under stable conditions. Indeed, the use of several resources with lower efficiency can be an equally, or even more, successful strategy in nature. Second, when limiting nutrients promote the growth of bacterio- and picophytoplankton, invading mixotrophs may have a habitat-ameliorating effect for higher trophic levels, gauged in terms of food quantity and quality. Using given resources more efficiently, O. tuberculata generated higher biomasses and expressed an increased nutritional value for potential planktivores, due to decreased cellular carbon to phosphorus (C:P) ratios compared to specialized plankton taxa. Our findings may help to explain why energy transfer efficiency between phytoplankton and higher trophic levels is generally higher in oligotrophic systems than in nutrient rich environments.     

Biogenic methane in freshwater food webs

1. It has long been known that substantial amounts of methane are produced in anoxic lake sediments, and the components of the methane cycle in lakes have been well described. At oxic–anoxic interfaces, methane‐oxidising bacteria (MOB) convert methane to microbial biomass and can be highly productive. However, only recently has methane been recognised as a potentially important carbon and energy source for lake food webs, and some instances have also been reported of methane contribution to river food webs. Stable isotope analysis (SIA) has provided compelling evidence in this respect and has been supplemented by other lines of evidence. 2. In the benthic food webs of lakes, profundal chironomid larvae appear to be the main conduits for trophic transfer of biogenic methane via grazing on MOB. The mode of feeding of these larvae and the microhabitats they generate both promote larval ability to exploit MOB production. Support to chironomid larvae from methane is rather widespread, but its degree is highly variable; estimates suggest that in some lakes methane‐carbon might contribute more than 60% of chironomid carbon biomass. 3. Evidence of crustacean zooplankton in lakes deriving part of their carbon from methane is currently more limited. Reports from some lakes have indicated Daphnia with a substantial (>50%) contribution of methane‐carbon in their biomass. However, for this to happen, an oxic–anoxic interface where sufficient MOB production can occur needs to be within the range of vertical migrations by zooplankton, which may only rarely be the case. Hence, a significant methane subsidy of pelagic food webs in lakes is probably much less widespread than for benthic food webs. 4. There is also recent and currently very limited evidence that some stream benthos derives biomass carbon (reported values up to 30%) from methane. This can occur in stagnant backwater pools where conditions can be analogous to those in lake sediments. However, groundwater aquifers can also supply water supersaturated with methane to some rivers, providing a basis for a microbially‐mediated transfer of methane‐carbon to river benthos. 5. Evidence for significant transfer of methane‐derived carbon to higher trophic levels is still very limited. Within some lakes, those fish species that feed extensively on chironomid larvae can derive a substantial part (perhaps up to 20%) of their carbon biomass from methane. It is also likely that methane‐carbon produced in lakes or rivers is exported to riparian ecosystems when emerging chironomids or other insects are eaten by invertebrate or avian predators. 6. We argue that conceptual models of freshwater food webs, and especially those for lakes, need to be modified to enable incorporation of biogenic methane as a carbon and energy source. For some types of lakes, carbon and energy budgets certainly need to take account of the production and utilisation of biogenic methane, and the accumulating evidence indicates that this is a more widespread phenomenon that has generally been acknowledged hitherto.     

Impact of nitrogen deposition on forest and lake food webs in nitrogen‐limited environments

Increased reactive nitrogen (N_r) deposition has raised the amount of N available to organisms and has greatly altered the transfer of energy through food webs, with major consequences for trophic dynamics. The aim of this review was to: (i) clarify the direct and indirect effects of N_r deposition on forest and lake food webs in N‐limited biomes, (ii) compare and contrast how aquatic and terrestrial systems respond to increased N_r deposition, and (iii) identify how the nutrient pathways within and between ecosystems change in response to N_r deposition. We present that N_r deposition releases primary producers from N limitation in both forest and lake ecosystems and raises plants' N content which in turn benefits herbivores with high N requirements. Such trophic effects are coupled with a general decrease in biodiversity caused by different N‐use efficiencies; slow‐growing species with low rates of N turnover are replaced by fast‐growing species with high rates of N turnover. In contrast, N_r deposition diminishes below‐ground production in forests, due to a range of mechanisms that reduce microbial biomass, and decreases lake benthic productivity by switching herbivore growth from N to phosphorus (P) limitation, and by intensifying P limitation of benthic fish. The flow of nutrients between ecosystems is expected to change with increasing N_r deposition. Due to higher litter production and more intense precipitation, more terrestrial matter will enter lakes. This will benefit bacteria and will in turn boost the microbial food web. Additionally, N_r deposition promotes emergent insects, which subsidize the terrestrial food web as prey for insectivores or by dying and decomposing on land. So far, most studies have examined N_r‐deposition effects on the food web base, whereas our review highlights that changes at the base of food webs substantially impact higher trophic levels and therefore food web structure and functioning.     

Influences of fish on food web structure and function in mountain lakes

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Global change‐driven effects on dissolved organic matter composition: Implications for food webs of northern lakes

Northern ecosystems are experiencing some of the most dramatic impacts of global change on Earth. Rising temperatures, hydrological intensification, changes in atmospheric acid deposition and associated acidification recovery, and changes in vegetative cover are resulting in fundamental changes in terrestrial–aquatic biogeochemical linkages. The effects of global change are readily observed in alterations in the supply of dissolved organic matter (DOM)—the messenger between terrestrial and lake ecosystems—with potentially profound effects on the structure and function of lakes. Northern terrestrial ecosystems contain substantial stores of organic matter and filter or funnel DOM, affecting the timing and magnitude of DOM delivery to surface waters. This terrestrial DOM is processed in streams, rivers, and lakes, ultimately shifting its composition, stoichiometry, and bioavailability. Here, we explore the potential consequences of these global change‐driven effects for lake food webs at northern latitudes. Notably, we provide evidence that increased allochthonous DOM supply to lakes is overwhelming increased autochthonous DOM supply that potentially results from earlier ice‐out and a longer growing season. Furthermore, we assess the potential implications of this shift for the nutritional quality of autotrophs in terms of their stoichiometry, fatty acid composition, toxin production, and methylmercury concentration, and therefore, contaminant transfer through the food web. We conclude that global change in northern regions leads not only to reduced primary productivity but also to nutritionally poorer lake food webs, with discernible consequences for the trophic web to fish and humans.     

Benthos as the basis for arctic lake food webs

Plankton have traditionally been viewed as the basis for limnetic food webs, with zooplankton acting as a gateway for energy passing between phytoplanktonic primary producers and fish. Often, benthic production has been considered to be important primarily in shallow systems or as a subsidy to planktonic food web pathways. Stable isotope food web analyses of two arctic lakes (NE14 and I minus) in the Toolik Lake region of Alaska indicate that benthos are the primary source of carbon for adults of all species of benthic and pelagic fish present. We found no effect of turbidity, which may suppress benthic algae by shading, on food web structure. Even though Secchi transparency varied from 10.2 m in NE14 to 0.55–2.6 m in I minus, food webs in both lakes were based upon benthos, had four trophic levels, and culminated with omnivorous lake trout. We suggest that the importance of benthos in the food webs of these lakes is due to their extreme oligotrophy, resulting in planktonic resources that are insufficient for the support of planktivorous consumers.     

From clear lakes to murky waters – tracing the functional response of high‐latitude lake communities to concurrent ‘greening’ and ‘browning’

Climate change and the intensification of land use practices are causing widespread eutrophication of subarctic lakes. The implications of this rapid change for lake ecosystem function remain poorly understood. To assess how freshwater communities respond to such profound changes in their habitat and resource availability, we conducted a space‐for‐time analysis of food‐web structure in 30 lakes situated across a temperature‐productivity gradient equivalent to the predicted future climate of subarctic Europe (temperature +3°C, precipitation +30% and nutrient +45 μg L^?1 total phosphorus). Along this gradient, we observed an increase in the assimilation of pelagic‐derived carbon from 25 to 75% throughout primary, secondary and tertiary consumers. This shift was overwhelmingly driven by the consumption of pelagic detritus by benthic primary consumers and was not accompanied by increased pelagic foraging by higher trophic level consumers. Our data also revealed a convergence of the carbon isotope ratios of pelagic and benthic food web endmembers in the warmest, most productive lakes indicating that the incorporation of terrestrial derived carbon into aquatic food webs increases as land use intensifies. These results, reflecting changes along a gradient characteristic of the predicted future environment throughout the subarctic, indicate that climate and land use driven eutrophication and browning are radically altering the function and fuelling of aquatic food webs in this biome.     

Between‐lake variation in the trophic ecology of an invasive crayfish

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Analysis of the community structure of planktonic ciliated protozoa relative to trophic state in Florida lakes

The planktonic ciliate populations of 30 Florida lakes constituting a broad trophic gradient were examined to determine the response of protozoan community structure to increasing eutrophication. Both ciliate abundance and biomass were strongly related to lake trophic state. Comparison of the Florida data base with a comparable north temperate lake group indicated that subtropical lakes generally possess higher ciliate abundance and biomass at a given trophic state than temperate lakes. However the equations derived for each data base were not significantly different. Community diversity and species richness increased with increasing lake productivity. Highly acidic lakes displayed significantly reduced diversity and numbers of species when contrasted with nonacidic oligotrophic lakes. Small-bodied (< 30 um) ciliates dominated all lakes but were proportionally less important in oligotrophic lakes. Presence-absence data produced three assemblages: an ubiquitous association of primarily small ciliate taxa, a group of large ciliates mainly restricted to eutrophic-hypereutrophic lakes, and a very large ciliate,Stentor niger, which dominated the protozoan communities of acidic oligotrophic lakes.