Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.     

The role of omnivorous crayfish in littoral communities

Large omnivorous predators may play particularly important roles determining the structure of communities because of their broad diets and simultaneous effects on multiple trophic levels. From June 2001 to June 2002 we quantified community structure and ecosystem attributes of six newly establishing freshwater ponds (660 m² each) after populations of omnivorous crayfish (Orconectes virilis) were introduced to three of the ponds. Crayfish preyed heavily on fish eggs in this experiment, which reduced recruitment of young-of-year fish. This effect indirectly enhanced zooplankton biomass in crayfish ponds. Phytoplankton abundance exhibited a more complex pattern and was probably influenced by non-trophic (e.g., bioturbation) effects of crayfish. Peak dissolved oxygen levels were lower in the crayfish ponds indicating that they had lower primary production: respiration ratios. Metaphytic algae were strongly affected by crayfish presence; filamentous greens quickly disappeared and the blue-green Gleotrichia (a less preferred food item) eventually dominated the composition in crayfish ponds. Chara vulgaris and vascular macrophytes established 34% cover in control ponds by June 2002, but were not able to establish in crayfish ponds. Two important periphyton herbivores (tadpoles and gastropods) were absent or significantly reduced in the crayfish ponds, but periphyton differences were temporally variable and not easily explained by a simple trophic cascade (i.e., crayfish—snails and tadpoles—periphyton). Our results indicate that crayfish can have dramatic direct and indirect impacts on littoral pond communities via feeding links with multiple trophic levels (i.e., fish, invertebrates, and plants) and non-trophic activities (bioturbation). Although the effects of omnivorous crayfish on littoral communities can be large, their complex effects do not fit neatly into current theories about trophic interactions or freshwater community structure.     

Compositional and functional consequences of environmental change in Belgian farmland ponds

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Indirect effects of fish on macrophytes in Bays Mountain Lake: evidence for a littoral trophic cascade

Summary We began this experiment to test specific hypotheses regarding direct and indirect effects of fish predation on the littoral macroinvertebrate community of Bays Mountain Lake, Tennessee. We used 24 m² enclosures in which we manipulated the presence and absence of large redear sunfish (Lepomis microlophus>150 mm SL), and small sunfish (L. macrochirus and L. microlophus <50 mm SL) over a 16-mo period. Here we report on effects of fish predation on gastropod grazers that appear to cascade to periphyton and macrophytes.Both large redear sunfish and small sunfish maintained low snail biomass, but snails in fish-free controls increased significantly during the first 2-mo of the experiment. By late summer of the first year of the experiment, the difference in biomass between enclosures with and without fish had increased dramatically (>10×). Midway through the second summer of the experiment, we noted apparent differences in the abundance of periphyton between enclosures containing fish and those that did not. We also noted differences in the macrophyte distribution among enclosures. To document these responses, we estimated periphyton cover, biovolume and cell size frequencies as well as macrophyte distributions among enclosures at the end of the experiment. When fish were absent, periphyton percent cover was significantly reduced compared to when fish were present. Periphyton cell-size distributions in enclosures without fish were skewed toward small cells (only 12% were greater than 200 m³), which is consistent with intense snail grazing. The macrophyte Najas flexilis had more than 60 x higher biomass in the fish-free enclosures than in enclosures containing fish; Potamogeton diversifolius was found only in fish-free enclosures. These results suggest a chain of strong interactions (i.e. from fish to snails to periphyton to macrophytes) that may be important in lake littoral systems. This contrasts sharply with earlier predictions based on cascading trophic interactions that propose that fish predation on snails would enhance macrophyte biomass.     

Beetle assemblages in ponds: effects of habitat and site age

• 1 Water beetle assemblages were sampled in each of 18 freshwater ponds, including 11 recently constructed sites designed to provide mitigation for wetlands destruction elsewhere, and seven older reference sites. There were three objectives: (a) to relate taxon richness and biomass of the beetles to the same properties of the wider aquatic invertebrate community, (b) to evaluate changes in beetle assemblage structure over time, and (c) to determine habitat effects on taxonomic composition, mean body size and trophic guild structure of the beetle assemblage.
• 2 Forty‐seven beetle genera were identified, representing 77 species. The beetles represented an average of 21.5% of total generic richness, but only 3.7% of total wet biomass of the wider invertebrate community.
• 3 Of all variables evaluated using canonical correspondence analysis (CCA), site age had the greatest influence on the beetle assemblage. Predatory dytiscids were early colonists at younger sites, while herbivorous curculionids and chrysomelids associated with particular types of vegetation typically occurred in older ponds. Mitigation ponds and reference ponds supported similar numbers of species. Reference sites, however, harboured substantially more unique species found at only a single site within the study area.
• 4 The presence of fish was also strongly related to beetle assemblage structure. Ponds with few or no fish contained about 3‐fold higher biomass and 3‐fold greater mean wet weight per individual compared to ponds with substantial fish assemblages.
• 5 Beetle assemblage composition varied among sites and sampling years, but beetle biomass, richness and species composition may be useful tools in evaluating the success of wetland mitigation efforts.

     

Predator-induced morphological plasticity across local populations of a freshwater snail

The expression of anti-predator adaptations may vary on a spatial scale, favouring traits that are advantageous in a given predation regime. Besides, evolution of different developmental strategies depends to a large extent on the grain of the environment and may result in locally canalized adaptations or, alternatively, the evolution of phenotypic plasticity as different predation regimes may vary across habitats. We investigated the potential for predator-driven variability in shell morphology in a freshwater snail, Radix balthica, and whether found differences were a specialized ecotype adaptation or a result of phenotypic plasticity. Shell shape was quantified in snails from geographically separated pond populations with and without molluscivorous fish. Subsequently, in a common garden experiment we investigated reaction norms of snails from populations' with/without fish when exposed to chemical cues from tench (Tinca tinca), a molluscivorous fish. We found that snails from fish-free ponds had a narrow shell with a well developed spire, whereas snails that coexisted with fish had more rotund shells with a low spire, a shell morphology known to increase survival rate from shell-crushing predators. The common garden experiment mirrored the results from the field survey and showed that snails had similar reaction norms in response to chemical predator cues, i.e. the expression of shell shape was independent of population origin. Finally, we found significant differences for the trait means among populations, within each pond category (fish/fish free), suggesting a genetic component in the determination of shell morphology that has evolved independently across ponds.     

Effects of a top invertebrate predator (<Emphasis Type="Italic">Dytiscus alaskanus</Emphasis>; Coleoptera: Dytiscidae) on fishless pond ecosystems

We investigated the predatory effects of Dytiscus alaskanus, a large predaceous diving beetle, on the biomass, species composition and diversity of fishless pond communities. The effects were tested using presence and absence treatments of D. alaskanus in 24 mesocosms distributed among six ponds. We sampled phytoplankton, zooplankton and macroinvertebrates every two weeks for a six week period. Periphyton was sampled from the mesocosm walls on the final day. Total macroinvertebrate biomass decreased in the presence of dytiscids while species richness was not affected. Macroinvertebrate predators, snails and Gammarus lacustris decreased in the dytiscid treatments. Laboratory feeding experiments confirmed feeding preferences consistent with the mesocosm results. Periphyton biomass was six times greater in the dytiscid enclosures, concomitant with the decreased grazing by gastropods and other invertebrate primary consumers indicating a benthic trophic cascade. Top–down effects of dytiscids on other predatory invertebrates led to increased total zooplankton biomass, largely due to increased abundances of large and small cladocerans. Zooplankton species richness increased in the dytiscid enclosures. Inconsistent with trophic cascade theory, phytoplankton did not respond to top–down effects of D. alaskanus within the study period. Overall, the results show D. alaskanus predation caused trophic effects via two distinct food chains, a dytiscid–snail–periphyton trophic cascade, and a dytiscid–predatory macroinvertebrates–zooplankton partial trophic cascade.     

Food web architecture and basal resources interact to determine biomass and stoichiometric cascades along a benthic food web

Understanding the effects of predators and resources on primary producers has been a major focus of interest in ecology. Within this context, the trophic cascade concept especially concerning the pelagic zone of lakes has been the focus of the majority of these studies. However, littoral food webs could be especially interesting because base trophic levels may be strongly regulated by consumers and prone to be light limited. In this study, the availability of nutrients and light and the presence of an omnivorous fish (Hyphessobrycon bifasciatus) were manipulated in enclosures placed in a humic coastal lagoon (Cabiúnas Lagoon, Macaé - RJ) to evaluate the individual and interactive effects of resource availability (nutrients and light) and food web configuration on the biomass and stoichiometry of periphyton and benthic grazers. Our findings suggest that light and nutrients interact to determine periphyton biomass and stoichiometry, which propagates to the consumer level. We observed a positive effect of the availability of nutrients on periphytic biomass and grazers' biomass, as well as a reduction of periphytic C∶N∶P ratios and an increase of grazers' N and P content. Low light availability constrained the propagation of nutrient effects on periphyton biomass and induced higher periphytic C∶N∶P ratios. The effects of fish presence strongly interacted with resource availability. In general, a positive effect of fish presence was observed for the total biomass of periphyton and grazer's biomass, especially with high resource availability, but the opposite was found for periphytic autotrophic biomass. Fish also had a significant effect on periphyton stoichiometry, but no effect was observed on grazers' stoichiometric ratios. In summary, we observed that the indirect effect of fish predation on periphyton biomass might be dependent on multiple resources and periphyton nutrient stoichiometric variation can affect consumers' stoichiometry.     

Complex impacts of an invasive omnivore and native consumers on stream communities in California and Hawaii

The effects of invasive species on native communities often depend on the characteristics of the recipient community and on the food habits of the invasive species, becoming complicated when the invader is omnivorous. In field enclosure experiments, we assessed the direct and interactive effects of an invasive omnivorous crayfish (Procambarus clarkii) and either native herbivorous snails (Physella gyrina) or shrimp (Atyoida bisulcata) on stream communities in California and Hawaii, respectively. Based on literature data and the characteristics of each study site, we predicted that crayfish would affect primarily algal-based trophic linkages in an open California stream but detritus-based trophic linkages in a shaded Hawaiian stream, with trophic cascades mediated through crayfish effects on primary consumers being observed in both systems. As predicted, crayfish in California directly reduced periphyton, filamentous algae, sediment, and snail levels, but generated a cascade by decreasing snail densities and increasing periphyton biomass. Contrary to prediction, crayfish did not reduce total invertebrate biomass. As predicted, crayfish in Hawaii reduced leaf litter, filamentous algae, and benthic invertebrate biomass. Contrary to our predictions, however, a trophic cascade was not observed because shrimp did not affect periphyton levels, crayfish did not reduce shrimp abundance, and crayfish had greater negative impacts on filamentous algae than did shrimp. Our findings highlight that the same invasive species can generate different effects on disparate systems, probably as mediated through the availability of different food types, flexibility in the invasive species’ food habits, and complex pathways of trophic interaction.     

Chemical cues modify species interactions: the ecological consequences of predator avoidance by freshwater snails

Chemical signals released by predators or injured prey often induce shifts in the traits of prey species, which may in turn affect species interactions. Here we investigate the role that chemical cues play in mediating species interactions in the littoral food web of lakes. Previous studies have shown that predators induce shifts in the morphology, life history, and behavior of the freshwater snail Physella, but the ecological consequences of developing these inducible defenses are not well documented. We observed habitat use of the freshwater snail Physella gyrina along a depth gradient in a natural lake, and found they increased their use of covered habitats with increasing depth. We hypothesized that this habitat shift was due to changes in the level and type of predation risk, and that the habitat shift would affect periphyton standing crops. These hypotheses were tested in a mesocosm experiment in which we manipulated the presence of molluscivorous fish and crayfish. Predators were confined to cages and snail density was identical in all treatments, so any effects of predators were mediated through trait shifts induced by chemical cues. In the presence of fish, Physella moved under cover, but in the presence of crayfish, Physella avoided cover and moved to the water surface. These non‐lethal effects of predators on snail habitat use influenced the interaction between snails and their periphyton resources. In the presence of fish, periphyton standing crop in covered habitats was reduced to just 8% of periphyton in the absence of fish. Crayfish had no significant effect on periphyton in covered habitats, but they reduced periphyton in near‐surface habitats to 39% of the standing crop in the absence of crayfish. The combined effects of fish and crayfish were generally intermediate to their individual effects. We conclude that because chemical cues often have strong effects on individual traits and trophic interactions are sensitive to trait values, chemical cues may play an important role in shaping the structure and dynamics of food webs.     

Indirect and direct controls of macroinvertebrates and small fish by abiotic factors and trophic interactions in the Florida Everglades

1. The roles of nutrients, disturbance and predation in regulating consumer densities have long been of interest, but their indirect effects have rarely been quantified in wetland ecosystems. The Florida Everglades contains gradients of hydrological disturbance (marsh drying) and nutrient enrichment (phosphorus), often correlated with densities of macroinvertebrate infauna (macroinvertebrates inhabiting periphyton), small fish and larger invertebrates, such as snails, grass shrimp, insects and crayfish. However, most causal relationships have yet to be quantified. 2. We sampled periphyton (content and community structure) and consumer (small omnivores, carnivores and herbivores, and infaunal macroinvertebrates inhabiting periphyton) density at 28 sites spanning a range of hydrological and nutrient conditions and compared our data to seven a priori structural equation models. 3. The best model included bottom‐up and top‐down effects among trophic groups and supported top‐down control of infauna by omnivores and predators that cascaded to periphyton biomass. The next best model included bottom‐up paths only and allowed direct effects of periphyton on omnivore density. Both models suggested a positive relationship between small herbivores and small omnivores, indicating that predation was unable to limit herbivore numbers. Total effects of time following flooding were negative for all three consumer groups even when both preferred models suggested positive direct effects for some groups. Total effects of nutrient levels (phosphorus) were positive for consumers and generally larger than those of hydrological disturbance and were mediated by changes in periphyton content. 4. Our findings provide quantitative support for indirect effects of nutrient enrichment on consumers, and the importance of both algal community structure and periphyton biomass to Everglades food webs. Evidence for top‐down control of infauna by omnivores was noted, though without substantially greater support than a competing bottom‐up‐only model.     

Periphytic food and predatory crayfish: relative roles in determining snail distribution

Summary In the laboratory and field, we examined how periphyton (food of snails) and predatory crayfish influenced snail distribution in Trout Lake, a permanent, northern Wisconsin lake. Laboratory experiments (with no crayfish) tested the importance of periphyton biomass in determining snail preference among rocks, and among rock, sand, and macrophyte substrates. Among rocks with four different amounts of periphyton, periphyton biomass and the number of Lymnaea emarginata, Physa spp., and Amnicola spp. were positively related. A similar, but non-significant, trend occurred for Helisoma anceps. A field experiment at a site in Trout Lake where predation risk was low confirmed the preference by snails for periphyton covered rocks; more snails colonized rocks with periphyton than rocks without. When given a choice of rock, sand, and macrophytes in the laboratory, L. emarginata preferred high periphyton biomass and rock. Laboratory and field results contrasted with the distribution of snails in Trout Lake; no snails occurred in areas with abundant periphyton-covered rocks, but snails were abundant nearby on scattered rocks with little periphyton. However, where snails were absent, crayfish were abundant (14.5 crayfish-trap^–1-day^–1), and where snails were abundant, crayfish were rare (3.2 crayfish-trap^–1-day^–1), suggesting that crayfish predation reduced snails. The hypothesis that the negative association between snail and periphyton biomass resulted from snail grazing was supported by the results of a field snail enclosure-exclosure experiment (1 m² cages; n=3). All experiments and observations therefore suggest that: 1) crayfish predation is more important than a preference for high periphyton biomass in determining snail distribution in Trout Lake; 2) periphyton biomass is negtively related to snail grazing; and 3) crayfish had a positive indirect effect on periphyton by preying on grazing snails.     

Top‐down and bottom‐up control of periphyton by benthivorous fish and light supply in two streams

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Water temperature determines strength of top-down control in a stream food web

1. We examined effects of water temperature on the community structure of a three trophic level food chain (predatory fish, herbivorous caddisfly larvae and periphyton) in boreal streams. We used laboratory experiments to examine (i) the effects of water temperature on feeding activities of fish and caddisfly larvae and on periphyton productivity, to evaluate the thermal effects on each trophic level (species‐level experiment), and (ii) the effects of water temperature on predation pressure of fish on abundance of the lower trophic levels, to evaluate how temperature affects top‐down control by fish (community‐level experiment). 2. In the species‐level experiment, feeding activity of fish was high at 12 °C, which coincides with the mean summer temperature in forested streams of Hokkaido, Japan, but was depressed at 3 °C, which coincides with the mean winter temperature, and also above 18 °C, which coincides with the near maximum summer temperatures. Periphyton productivity increased over the range of water temperatures. 3. In the community‐level experiments, a top‐down effect of fish on the abundance of caddisfly larvae and periphyton was clear at 12 °C. This effect was not observed at 3 and 21 °C because of low predation pressure of fish at these temperatures. 4. These experiments revealed that trophic cascading effects may vary with temperature even in the presence of abundant predators. Physiological depression of predators because of thermal stress can alter top‐down control and lead to changes in community structure. 5. We suggest that thermal habitat alteration can change food web structure via combinations of direct and indirect trophic interactions.     

Dragonfly predators influence biomass and density of pond snails

Studies in lakes show that fish and crayfish predators play an important role in determining the abundance of freshwater snails. In contrast, there are few studies of snails and their predators in shallow ponds and marshes. Ponds often lack fish and crayfish but have abundant insect populations. Here we present the results of field surveys, laboratory foraging trials, and an outdoor mesocosm experiment, testing the hypothesis that insects are important predators of pulmonate snails. In laboratory foraging trials, conducted with ten species of insects, most insect taxa consumed snails, and larval dragonflies were especially effective predators. The field surveys showed that dragonflies constitute the majority of the insect biomass in fishless ponds. More focused foraging trials evaluated the ability of the dragonflies Anax junius and Pantala hymenaea to prey upon different sizes and species of pulmonate snails (Helisoma trivolvis, Physa acuta, and Stagnicola elodes). Anax junius consumed all three species up to the maximum size tested. Pantala hymenaea consumed snails with a shell height of 3 mm and smaller, but did not kill larger snails. P. acuta were more vulnerable to predators than were H. trivolvis or S. elodes. In the mesocosm experiment, conducted with predator treatments of A. junius, P. hymenaea, and the hemipteran Belostoma flumineum, insect predators had a pronounced negative effect on snail biomass and density. A. junius and B. flumineum reduced biomass and density to a similar degree, and both reduced biomass more than did P. hymenaea. Predators did not have a strong effect on species composition. A model suggested that A. junius and P. hymenaea have the largest effects on snail biomass in the field. Given that both pulmonate snails and dragonfly nymphs are widespread and abundant in marshes and ponds, snail assemblages in these water bodies are likely regulated in large part by odonate predation.     

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

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Habitat structural complexity mediates food web dynamics in a freshwater macrophyte community

A considerable amount of research has investigated the influence of habitat structure on predator success, yet few studies have explored the implications for community structure and food-web dynamics. The relative importance of macrophyte structure and fish predation on the composition of the macroinvertebrate and periphyton communities in a lowland river was investigated using a multifactorial caging experiment. We hypothesised that: (1) fish predators are less effective in a more structurally complex macrophyte analogue; (2) strong direct and indirect effects of fish predators (e.g. trophic cascades) are less likely to occur in a structurally complex habitat; and (3) the strength of these patterns is influenced by the composition of the prevailing community assemblage. We measured the abundance and composition of the macroinvertebrate and periphyton communities associated with three different-shaped macrophyte analogues, under different fish predator treatments and at different times. Macrophyte analogue architecture had strong, consistent effects on both the macroinvertebrate and periphyton communities; both were most abundant and diverse on the most structurally complex plant analogue. In contrast, the fish predators affected only a subset of the macroinvertebrate community and there was a suggestion of minor indirect effects on periphyton community composition. Contrary to expectations, the fish predators had their strongest effects in the most structurally complex macrophyte analogue. We conclude that in this system, macrophyte shape strongly regulates the associated freshwater assemblage, resulting in a diverse community structure less likely to exhibit strong effects of fish predation.     

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.     

Relationship between Snail Population Density and Infection Status of Snails and Fish with Zoonotic Trematodes in Vietnamese Carp Nurseries

Background

Fish-borne zoonotic trematodes (FZT) are a food safety and health concern in Vietnam. Humans and other final hosts acquire these parasites from eating raw or under-cooked fish with FZT metacercariae. Fish raised in ponds are exposed to cercariae shed by snail hosts that are common in fish farm ponds. Previous risk assessment on FZT transmission in the Red River Delta of Vietnam identified carp nursery ponds as major sites of transmission. In this study, we analyzed the association between snail population density and heterophyid trematode infection in snails with the rate of FZT transmission to juvenile fish raised in carp nurseries.

Methodology/Principal Findings

Snail population density and prevalence of trematode (Heterophyidae) infections were determined in 48 carp nurseries producing Rohu juveniles, (Labeo rohita) in the Red River Delta area. Fish samples were examined at 3, 6 and 9 weeks after the juvenile fish were introduced into the ponds. There was a significant positive correlation between prevalence of FZT metacercariae in juvenile fish and density of infected snails. Thus, the odds of infection in juvenile fish were 4.36 and 11.32 times higher for ponds with medium and high density of snails, respectively, compared to ponds where no infected snails were found. Further, the intensity of fish FZT infections increased with the density of infected snails. Interestingly, however, some ponds with no or few infected snails were collected also had high prevalence and intensity of FZT in juvenile fish. This may be due to immigration of cercariae into the pond from external water sources.

Conclusions/Significance

The total number and density of potential host snails and density of host snails infected with heterophyid trematodes in the aquaculture pond is a useful predictor for infections in juvenile fish, although infection levels in juvenile fish can occur despite low density or absence infected snails. This suggests that intervention programs to control FZT infection of fish should include not only intra-pond snail control, but also include water sources of allochthonous cercariae, i.e. canals supplying water to ponds as well as snail habitats outside the pond such as rice fields and surrounding ponds.