Farming practices change food web structures in cereal aphid–parasitoid–hyperparasitoid communities

Agricultural intensification has been shown to result in a decline in biodiversity across many taxa, but the changes in community structure and species interactions remain little understood. We have analysed and compared the structure of feeding interactions for cereal aphids and their primary and secondary parasitoids in organically and conventionally managed winter wheat fields using quantitative food web metrics (interaction evenness, generality, vulnerability, link density). Despite little variation in the richness of each trophic group, food web structures between the two farming systems differed remarkably. In contrast to common expectations, aphids and primary parasitoids were characterized by (1) a higher evenness of interaction frequencies (interaction evenness) in conventional fields, which cascaded to interactions at the next trophic level, with (2) a higher interaction evenness, (3) a higher ratio of primary parasitoid taxa per secondary parasitoid (generality) and (4) a higher link density. Aphid communities in the organically managed fields almost exclusively consisted of a single ear-colonizing species, Sitobion avenae, while highly fertilized conventional fields were mainly infested by leaf-colonizing aphids that benefit from the nutritional status of winter wheat. In conclusion, agricultural intensification appears to foster the complexity of aphid–parasitoid food webs, thereby not supporting the general expectation on the importance of organic farming practices for species richness and food web complexity.     

Resource tracking in marine parasites: going with the flow?

Understanding how diversity interacts with energy supply is of broad ecological interest. Most studies to date have investigated patterns within trophic levels, reflecting a lack of food webs which include information on energy flow. We added parasites to a published marine energy‐flow food web, to explore whether parasite diversity is correlated with energy flow to host taxa. Parasite diversity was high with 36 parasite taxa affecting 40 of the 51 animal taxa. Adding parasites increased the number of trophic links per species, trophic link strength, connectance, and food chain lengths. There was evidence of an asymptotic relationship between energy flowing through a food chain and parasite diversity, although there were clear outliers. High parasite diversity was associated with host taxa which were highly connected within the food web. This suggests that energy flow through a taxon may favour parasite diversity, up to a maximal value. The evolutionary and energetic basis for that limitation is of key interest in understanding the basis for parasite diversity in natural food webs and thus their role in food web dynamics.     

Effects of extinction on food web structures on an evolutionary time scale

Extinction affected food web structure in paleoecosystems. Recent theoretical studies that examined the effects of extinction intensity on food web structure on ecological time scales have considered extinction to involve episodic events, with pre-extinction food webs becoming established without dynamics. However, in terms of the paleontological time scale, food web structures are generated from feedback with repeated extinctions, because extinction frequency is affected by food web structure, and food web structure itself is a product of previous extinctions. We constructed a simulation model of changes in tri-trophic-level food webs to examine how continual extinction events affect food webs on an evolutionary time scale. We showed that under high extinction intensity (1) species diversity, especially that of consumer species, decreased; (2) the total population density at each trophic level decreased, while the densities of individual species increased; and (3) the trophic link density of the food web increased. In contrast to previous models, our results were based on an assumption of long-term food web development and are able to explain overall trends posited by empirical investigations based on fossil records.     

Using trophic hierarchy to understand food web structure

Link arrangement in food webs is determined by the species' feeding habits. This work investigates whether food web topology is organized in a gradient of trophic positions from producers to consumers. To this end, we analyzed 26 food webs for which the consumption rate of each species was specified. We computed the trophic positions and the link densities of all species in the food webs. Link density measures how much each species contributes to the distribution of energy in the system. It is expressed as the number of links species establish with other nodes, weighted by their magnitude. We computed these two metrics using various formulations developed in the ecological network analysis framework. Results show a positive correlation between trophic position and link density across all the systems, regardless the specific formulas used to measure the two quantities. We performed the same analysis on the corresponding binary matrices (i.e. removing information about rates). In addition, we investigated the relation between trophic position and link density in: a) simulated binary webs with same connectance as the original ones; b) weighted webs with constant topology but randomized link strengths and c) weighted webs with constant connectance where both topology and link strengths are randomized. The correlation between the two indices attenuates, vanishes or becomes negative in the case of binary food webs and simulated data (weighted and unweighted).
According to our analysis, link density in food webs decreases with trophic position so that it is greatly reduced toward the top of the trophic hierarchy. This outcome, that seems to challenge previous conclusions based on null models, strongly depends on link quantification. Including interaction strengths may improve substantially our understanding of food web organization, and possibly contradict results based on the analysis of binary webs.     

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.     

Food web framework for size-structured populations

We synthesise traditional unstructured food webs, allometric body size scaling, trait-based modelling, and physiologically structured modelling to provide a novel and ecologically relevant tool for size-structured food webs. The framework allows food web models to include ontogenetic growth and life-history omnivory at the individual level by resolving the population structure of each species as a size-spectrum. Each species is characterised by the trait ‘size at maturation’, and all model parameters are made species independent through scaling with individual body size and size at maturation. Parameter values are determined from cross-species analysis of fish communities as life-history omnivory is widespread in aquatic systems, but may be reparameterised for other systems. An ensemble of food webs is generated and the resulting communities are analysed at four levels of organisation: community level, species level, trait level, and individual level. The model may be solved analytically by assuming that the community spectrum follows a power law. The analytical solution provides a baseline expectation of the results of complex food web simulations, and agrees well with the predictions of the full model on biomass distribution as a function of individual size, biomass distribution as a function of size at maturation, and relation between predator-prey mass ratio of preferred and eaten food. The full model additionally predicts the diversity distribution as a function of size at maturation.     

Environmental fluctuations can stabilize food web dynamics by increasing synchrony

Natural food webs are species-rich, but classical theory suggests that they should be unstable and extinction-prone. Asynchronous fluctuations in the densities of competing consumers can stabilize food web dynamics in constant environments. However, environmental fluctuations often synchronize dynamics in nature. Using the same 'diamond-shape' food web model first used to demonstrate the stabilizing effects of asynchrony in constant environments, we show that weak-to-moderate environmentally induced fluctuations in consumer mortality rates stabilize food webs while disrupting asynchrony. Synchrony actually promotes stability because: (i) synchronous declines in consumer density reduce the maximum abundance of top predators and (ii) resource competition quickly converts synchronous increases in consumer density into synchronous declines. These results are robust to details of food web topology and the implementation of environmental fluctuations. The fluctuation strengths that enhance stability are within the range experienced naturally by many species, suggesting that stabilization via environmental fluctuations is a realistic possibility.     

Analogous aquatic and terrestrial food webs in the high Arctic: The structuring force of a harsh climate

Understanding food web structure and dynamics remains a central theme in ecology. Whilst differences between aquatic and terrestrial food webs have been the focus of several studies, we aim to reveal similarities where abiotic conditions are particularly extreme such as in the high Arctic. We propose that here, the combination of a short growing season, low temperature and low light, leads to the absence of predator control and the development of typically two-trophic, grazer-dominated food webs with high plant quality in terms of elemental ratios. Moreover, we advocate that this mechanism is evident in both aquatic and terrestrial high-Arctic environments, allowing the build-up of herbivore densities that consume a large fraction of plant primary production and tightly recycle nutrients. Thus, the particular abiotic conditions that characterise the high Arctic give rise to a unique environment that allows biotic factors to orchestrate food web structure and influence ecosystem function. Specially, the short growing season, low temperatures and low light levels collectively constrain the accumulation of structural autotroph tissue that, in temperate regions, effectively keeps herbivores at bay. While fundamental differences between terrestrial and aquatic ecosystems have been frequently advocated, we show here that harsh live-constraining conditions in the high Arctic have led to analogous, grazer-dominated, food web dynamics in both terrestrial and freshwater ecosystems.     

Parasites in food webs: the ultimate missing links

Parasitism is the most common consumer strategy among organisms, yet only recently has there been a call for the inclusion of infectious disease agents in food webs. The value of this effort hinges on whether parasites affect food‐web properties. Increasing evidence suggests that parasites have the potential to uniquely alter food‐web topology in terms of chain length, connectance and robustness. In addition, parasites might affect food‐web stability, interaction strength and energy flow. Food‐web structure also affects infectious disease dynamics because parasites depend on the ecological networks in which they live. Empirically, incorporating parasites into food webs is straightforward. We may start with existing food webs and add parasites as nodes, or we may try to build food webs around systems for which we already have a good understanding of infectious processes. In the future, perhaps researchers will add parasites while they construct food webs. Less clear is how food‐web theory can accommodate parasites. This is a deep and central problem in theoretical biology and applied mathematics. For instance, is representing parasites with complex life cycles as a single node equivalent to representing other species with ontogenetic niche shifts as a single node? Can parasitism fit into fundamental frameworks such as the niche model? Can we integrate infectious disease models into the emerging field of dynamic food‐web modelling? Future progress will benefit from interdisciplinary collaborations between ecologists and infectious disease biologists.     

Comparing global warming potential,energy use and land use of organic,conventional and integrated winter wheat production

To ensure a sustainable food supply for the growing population, the challenge is to find agricultural systems that can meet production requirements within environmental constraints and demands. This study compares the impacts of winter wheat production on energy use, land use and 100 years Global Warming Potential (GWP₁₀₀) under different arable farming systems and farming practices. Life cycle assessment was used to simulate the impacts of organic, conventional and integrated farming (IF) systems along the production chain from input production up to the farm gate. The IF system models were designed to combine the best practices from organic and conventional systems to reduce negative environmental impacts without significant yield reductions. An integrated system that used food waste digestate as a fertiliser, and utilised pesticides and no‐tillage had the lowest energy use and GWP per functional unit of 1000 kg wheat output. When the impacts of some specific practices for reducing energy use and GWP were compared, the highest energy use reductions were achieved by replacing synthetic nitrogen fertilisers with anaerobically treated food waste or nitrogen fixing crops, increasing yields through crop breeding and using no‐tillage instead of ploughing. The highest GWP reductions were achieved by using nitrification inhibitors, replacing synthetic nitrogen fertilisers and increasing yields. The major contributors to the uncertainty range of energy use were associated with machinery fuel use and the assumed crop yields. For GWP results, the main source of uncertainty related to the N₂O emissions. In conclusion, farming systems that combine the best practices from organic and conventional systems have potential to reduce negative environmental impacts while maintaining yield levels.     

Unravelling the complex structure of forest soil food webs: higher omnivory and more trophic levels

Food web topologies depict the community structure as distributions of feeding interactions across populations. Although the soil ecosystem provides important functions for aboveground ecosystems, data on complex soil food webs is notoriously scarce, most likely due to the difficulty of sampling and characterizing the system. To fill this gap we assembled the complex food webs of 48 forest soil communities. The food webs comprise 89 to 168 taxa and 729 to 3344 feeding interactions. The feeding links were established by combining several molecular methods (stable isotope, fatty acid and molecular gut content analyses) with feeding trials and literature data. First, we addressed whether soil food webs (n = 48) differ significantly from those of other ecosystem types (aquatic and terrestrial aboveground, n = 77) by comparing 22 food web parameters. We found that our soil food webs are characterized by many omnivorous and cannibalistic species, more trophic chains and intraguild‐predation motifs than other food webs and high average and maximum trophic levels. Despite this, we also found that soil food webs have a similar connectance as other ecosystems, but interestingly a higher link density and clustering coefficient. These differences in network structure to other ecosystem types may be a result of ecosystem specific constraints on hunting and feeding characteristics of the species that emerge as network parameters at the food‐web level. In a second analysis of land‐use effects, we found significant but only small differences of soil food web structure between different beech and coniferous forest types, which may be explained by generally strong selection effects of the soil that are independent of human land use. Overall, our study has unravelled some systematic structures of soil food‐webs, which extends our mechanistic understanding how environmental characteristics of the soil ecosystem determine patterns at the community level.     

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.     

Understorey environments influence functional diversity in tank‐bromeliad ecosystems

1. A substantial fraction of the freshwater available in neotropical forests is impounded within the rosettes of bromeliads that form aquatic islands in a terrestrial matrix. The ecosystem functioning of bromeliads is known to be influenced by the composition of the contained community but it is not clear whether bromeliad food webs remain functionally similar against a background of variation in the understorey environment. 2. We considered a broad range of environmental conditions, including incident light and incoming litter, and quantified the distribution of a very wide range of freshwater organisms (from viruses to macroinvertebrates) to determine the factors that influence the functional structure of bromeliad food webs in samples taken from 171 tank‐bromeliads. 3. We observed a gradient of detritus‐based to algal‐based food webs from the understorey to the overstorey. Algae, rotifers and collector and predatory invertebrates dominated bromeliad food webs in exposed areas, whereas filter‐feeding insects had their highest densities in shaded forest areas. Viruses, bacteria and fungi showed no clear density patterns. Detritus decomposition is mainly due to microbial activity in understorey bromeliads where filter feeders are the main consumers of microbial and particulate organic matter (POM). Algal biomass may exceed bacterial biomass in sun‐exposed bromeliads where amounts of detritus were lower but functional diversity was highest. 4. Our results provide evidence that tank‐bromeliads, which grow in a broad range of ecological conditions, promote aquatic food web diversity in neotropical forests. Moreover, although bromeliad ecosystems have been categorised as detritus‐based systems in the literature, we show that algal production can support a non‐detrital food web in these systems.     

The relationship between the duration of food web evolution and the vulnerability to biological invasion

I conducted computer simulations of food web evolution and investigated the relationship between the duration of food web evolution and the vulnerability to biological invasion. Model food webs without evolution consisted of animal species with a limited number of prey species and producer species with small intrinsic growth rates. Because these species were not resistant to predation pressure, model food webs without evolution were vulnerable to invasion of powerful omnivores, which had a wide range of feeding preference and a high ecological efficiency. In model food webs without evolution, the number of animal species depending on producer species was small. Therefore, if a producer species invaded and disturbed the base of such food webs, few animal species became extinct. However, model food webs with a long time evolution had a structure that a small number of producer species supported a large number of animal species. When a producer species invaded and disturbed the base of such food webs in this state, many species became extinct by an indirect effect. The mean number of prey species of animal species and the mean intrinsic growth rate of producer species increased rapidly in the early stage of evolution. Therefore, in the early stage of food web evolution, food webs were temporarily resistant to invasion of powerful omnivores. However, this resistibility was not maintained for a long time. The result of this study strongly suggests that food webs change with time, and consequently the vulnerability to invasion changes with time.     

Planktonic Food Web Structure along the Sau Reservoir (Spain) in Summer 1997

We studied the planktonic food web in eutrophic Sau Reservoir (Catalonia, NE Spain). Along the longitudinal axis from the Ter River downstream to the dam, we characterized a microbial succession of food web dominance of bacteria‐HNF‐ciliates. The Ter River transports a large load of organic material into the reservoir, with a bacterial density of ∼9 · 10⁶ large cells per ml. While at the first lacustrine station of the Reservoir HNF were the dominant bacterial consumers, at the others, an oligotrich ciliate, Halteria grandinella, was the main protozoan bacterivore. Most of the bacterial production in the reservoir epilimnion was consumed by grazing. The spatial succession of the reservoir microbial food webs was followed downstream by maximum densities of their potential predators among zoo‐plankters – rotifers, and early developmental stages of copepods.     

Predator and detritivore niche width helps to explain biocomplexity of experimental detritus-based food webs in four aquatic and terrestrial ecosystems

In the study of food webs, the existence and explanation of recurring patterns, such as the scale invariance of linkage density, predator–prey ratios and mean chain length, constitute long-standing issues. Our study focused on litter-associated food webs and explored the influence of detritivore and predator niche width (as δ¹³C range) on web topological structure. To compare patterns within and between aquatic and terrestrial ecosystems and take account of intra-habitat variability, we constructed 42 macroinvertebrate patch-scale webs in four different habitats (lake, lagoon, beech forest and cornfield), using an experimental approach with litterbags. The results suggest that although web differences exist between ecosystems, patterns are more similar within than between aquatic and terrestrial web types. In accordance with optimal foraging theory, we found that the niche width of predators and prey increased with the number of predators and prey taxa as a proportion of total taxa in the community. The tendency was more marked in terrestrial ecosystems and can be explained by a lower per capita food level than in aquatic ecosystems, particularly evident for predators. In accordance with these results, the number of links increased with the number of species but with a significantly sharper regression slope for terrestrial ecosystems. As a consequence, linkage density, which was found to be directly correlated to niche width, increased with the total number of species in terrestrial webs, whereas it did not change significantly in aquatic ones, where connectance scaled negatively with the total number of species. In both types of ecosystem, web robustness to rare species removal increased with connectance and the niche width of predators. In conclusion, although limited to litter-associated macroinvertebrate assemblages, this study highlights structural differences and similarities between aquatic and terrestrial detrital webs, providing field evidence of the central role of niche width in determining the structure of detritus-based food webs and posing foraging optimisation constraints as a general mechanistic explanation of food web complexity differences within and between ecosystem types.     

Shrinking by numbers: landscape context affects the species composition but not the quantitative structure of local food webs

1. With habitat fragmentation spreading around the world, there is a pressing need to understand its impacts on local food webs. To date, few studies have examined the effects of landscape context on multiple local communities in a quantitative, spatially realistic setting. 2. To examine how the isolation of a food web affects its structure, we construct local food webs of specialist herbivores and their natural enemies on 82 individual oaks (Quercus robur) growing in different landscape contexts. 3. Across this set of webs, we find that communities in isolated habitat patches not only contained fewer species than did well-connected ones, but also differed in species composition. 4. Surprisingly, the effects observed in terms of species composition were not reflected in the quantitative interaction structure of local food webs: landscape context had no detectable effect on either the interaction evenness, linkage density, connectance, generality or vulnerability of local webs. 5. We conclude that the quantitative structure of food webs may be stable in the face of habitat fragmentation, despite clear-cut impacts on individual species. This finding offers hope-inspiring news for conservation, but should clearly be verified by empirical studies across both naturally and more recently fragmented systems.     

Food web complexity and stability across habitat connectivity gradients

The effects of habitat connectivity on food webs have been studied both empirically and theoretically, yet the question of whether empirical results support theoretical predictions for any food web metric other than species richness has received little attention. Our synthesis brings together theory and empirical evidence for how habitat connectivity affects both food web stability and complexity. Food web stability is often predicted to be greatest at intermediate levels of connectivity, representing a compromise between the stabilizing effects of dispersal via rescue effects and prey switching, and the destabilizing effects of dispersal via regional synchronization of population dynamics. Empirical studies of food web stability generally support both this pattern and underlying mechanisms. Food chain length has been predicted to have both increasing and unimodal relationships with connectivity as a result of predators being constrained by the patch occupancy of their prey. Although both patterns have been documented empirically, the underlying mechanisms may differ from those predicted by models. In terms of other measures of food web complexity, habitat connectivity has been empirically found to generally increase link density but either reduce or have no effect on connectance, whereas a unimodal relationship is expected. In general, there is growing concordance between empirical patterns and theoretical predictions for some effects of habitat connectivity on food webs, but many predictions remain to be tested over a full connectivity gradient, and empirical metrics of complexity are rarely modeled. Closing these gaps will allow a deeper understanding of how natural and anthropogenic changes in connectivity can affect real food webs.     

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.