Food Web Perspective Research Articles

Habitat heterogeneity stabilizes the spatial and temporal interactions between cereal aphids and parasitic wasps

Although it is widely recognized that landscape heterogeneity (LH) can enhance the activity of natural enemies, little is known about the effect of LH on the spatial and temporal stability of pests and their natural enemies. Here, we designed a gradient of LH in agro-ecosystems and examined experimentally the relationship between LH and the spatial and temporal stability of cereal aphids and their parasitic wasps from 2009 to 2012. Stability was measured by the reciprocal of the spatial and temporal coefficients of variation of the population density. Results showed that LH can significantly homogenize the distribution of cereal aphids and their parasitic wasps, thus enhancing the spatial stability of the system. Increasing LH further dampened the fluctuation of the populations of primary parasitoids and hyperparasitoids, thus enhancing the temporal stability of the system. The stability of parasitism and hyperparasitism was also improved with the increase of LH, with the hyperparasitism being more sensitive to the heterogeneity change. Consequently, integrated pest management in agro-ecosystems could be better studied from a multi-trophic food-web perspective.

Developing a broader scientific foundation for river restoration: Columbia River food webs

Well-functioning food webs are fundamental for sustaining rivers as ecosystems and maintaining associated aquatic and terrestrial communities. The current emphasis on restoring habitat structure--without explicitly considering food webs--has been less successful than hoped in terms of enhancing the status of targeted species and often overlooks important constraints on ecologically effective restoration. We identify three priority food web-related issues that potentially impede successful river restoration: uncertainty about habitat carrying capacity, proliferation of chemicals and contaminants, and emergence of hybrid food webs containing a mixture of native and invasive species. Additionally, there is the need to place these food web considerations in a broad temporal and spatial framework by understanding the consequences of altered nutrient, organic matter (energy), water, and thermal sources and flows, reconnecting critical habitats and their food webs, and restoring for changing environments. As an illustration, we discuss how the Columbia River Basin, site of one of the largest aquatic/riparian restoration programs in the United States, would benefit from implementing a food web perspective. A food web perspective for the Columbia River would complement ongoing approaches and enhance the ability to meet the vision and legal obligations of the US Endangered Species Act, the Northwest Power Act (Fish and Wildlife Program), and federal treaties with Northwest Indian Tribes while meeting fundamental needs for improved river management.

Quantity and quality: unifying food web and ecosystem perspectives on the role of resource subsidies in freshwaters

Although the study of resource subsidies has emerged as a key topic in both ecosystem and food web ecology, the dialogue over their role has been limited by separate approaches that emphasize either subsidy quantity or quality. Considering quantity and quality together may provide a simple, but previously unexplored, framework for identifying the mechanisms that govern the importance of subsidies for recipient food webs and ecosystems. Using a literature review of > 90 studies of open-water metabolism in lakes and streams, we show that high-flux, low-quality subsidies can drive freshwater ecosystem dynamics. Because most of these ecosystems are net heterotrophic, allochthonous inputs must subsidize respiration. Second, using a literature review of subsidy quality and use, we demonstrate that animals select for high-quality food resources in proportions greater than would be predicted based on food quantity, and regardless of allochthonous or autochthonous origin. This finding suggests that low-flux, high-quality subsidies may be selected for by animals, and in turn may disproportionately affect food web and ecosystem processes (e.g., animal production, trophic energy or organic matter flow, trophic cascades). We then synthesize and review approaches that evaluate the role of subsidies and explicitly merge ecosystem and food web perspectives by placing food web measurements in the context of ecosystem budgets, by comparing trophic and ecosystem production and fluxes, and by constructing flow food webs. These tools can and should be used to address future questions about subsidies, such as the relative importance of subsidies to different trophic levels and how subsidies may maintain or disrupt ecosystem stability and food web interactions.

A food web perspective on large herbivore community limitation

The exceptional diversity of large mammals in African savannas provides an ideal opportunity to explore the relative importance of top-down and bottom-up controls of large terrestrial herbivore communities. Recent work has emphasized the role of herbivore and carnivore body size in shaping these trophic relationships. However, the lack of across-ecosystem comparisons using a common methodology prohibits general conclusions. Here we used published data on primary production, herbivore and carnivore densities and diets to estimate the consumption fluxes between three trophic levels in four African savanna ecosystems. Our food web approach suggests that the body size distribution within and across trophic levels has a strong influence on the strength of top-down control of herbivores by carnivores and on consumption fluxes within ecosystems, as predicted by theoretical food web models. We generalize findings from the Serengeti ecosystem that suggest herbivore species below 150 kg are more likely to be limited by predation. We also emphasize the key functional role played by the largest species at each trophic level. The abundance of the largest herbivore species largely governs the consumption of primary production in resident communities. Similarly, predator guilds in which the largest carnivore species represent a larger share of carnivore biomass are likely to exert a stronger top-down impact on herbivores. Our study shows how a food web approach allows integrating current knowledge and offers a powerful framework to better understand the functioning of ecosystems.

Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment

Biodiversity is rapidly declining, and this may negatively affect ecosystem processes, including economically important ecosystem services. Previous studies have shown that biodiversity has positive effects on organisms and processes across trophic levels. However, only a few studies have so far incorporated an explicit food-web perspective. In an eight-year biodiversity experiment, we studied an unprecedented range of above- and below-ground organisms and multitrophic interactions. A multitrophic data set originating from a single long-term experiment allows mechanistic insights that would not be gained from meta-analysis of different experiments. Here we show that plant diversity effects dampen with increasing trophic level and degree of omnivory. This was true both for abundance and species richness of organisms. Furthermore, we present comprehensive above-ground/below-ground biodiversity food webs. Both above ground and below ground, herbivores responded more strongly to changes in plant diversity than did carnivores or omnivores. Density and richness of carnivorous taxa was independent of vegetation structure. Below-ground responses to plant diversity were consistently weaker than above-ground responses. Responses to increasing plant diversity were generally positive, but were negative for biological invasion, pathogen infestation and hyperparasitism. Our results suggest that plant diversity has strong bottom-up effects on multitrophic interaction networks, with particularly strong effects on lower trophic levels. Effects on higher trophic levels are indirectly mediated through bottom-up trophic cascades.

Predicting invasion success in complex ecological networks

A central and perhaps insurmountable challenge of invasion ecology is to predict which combinations of species and habitats most effectively promote and prevent biological invasions. Here, we integrate models of network structure and nonlinear population dynamics to search for potential generalities among trophic factors that may drive invasion success and failure. We simulate invasions where 100 different species attempt to invade 150 different food webs with 15–26 species and a wide range (0.06–0.32) of connectance. These simulations yield 11 438 invasion attempts by non-basal species, 47 per cent of which are successful. At the time of introduction, whether or not the invader is a generalist best predicts final invasion success; however, once the invader establishes itself, it is best distinguished from unsuccessful invaders by occupying a lower trophic position and being relatively invulnerable to predation. In general, variables that reflect the interaction between an invading species and its new community, such as generality and trophic position, best predict invasion success; however, for some trophic categories of invaders, fundamental species traits, such as having the centre of the feeding range low on the theoretical niche axis (for non-omnivorous and omnivorous herbivores), or the topology of the food web (for tertiary carnivores), best predict invasion success. Across all invasion scenarios, a discriminant analysis model predicted successful and failed invasions with 76.5 per cent accuracy for properties at the time of introduction or 100 per cent accuracy for properties at the time of establishment. More generally, our results suggest that tackling the challenge of predicting the properties of species and habitats that promote or inhibit invasions from food web perspective may aid ecologists in identifying rules that govern invasions in natural ecosystems.

Causes and consequences of marine mammal population declines in southwest Alaska: a food-web perspective

Populations of sea otters, seals and sea lions have collapsed across much of southwest Alaska over the past several decades. The sea otter decline set off a trophic cascade in which the coastal marine ecosystem underwent a phase shift from kelp forests to deforested sea urchin barrens. This interaction in turn affected the distribution, abundance and productivity of numerous other species. Ecological consequences of the pinniped declines are largely unknown. Increased predation by transient (marine mammal-eating) killer whales probably caused the sea otter declines and may have caused the pinniped declines as well. Springer et al. proposed that killer whales, which purportedly fed extensively on great whales, expanded their diets to include a higher percentage of sea otters and pinnipeds following a sharp reduction in great whale numbers from post World War II industrial whaling. Critics of this hypothesis claim that great whales are not now and probably never were an important nutritional resource for killer whales. We used demographic/energetic analyses to evaluate whether or not a predator-prey system involving killer whales and the smaller marine mammals would be sustainable without some nutritional contribution from the great whales. Our results indicate that while such a system is possible, it could only exist under a narrow range of extreme conditions and is therefore highly unlikely.

Trophic cascades promote threshold-like shifts in pelagic marine ecosystems

Fisheries can have a large impact on marine ecosystems, because the effects of removing large predatory fish may cascade down the food web. The implications of these cascading processes on system functioning and resilience remain a source of intense scientific debate. By using field data covering a 30-year period, we show for the Baltic Sea that the underlying mechanisms of trophic cascades produced a shift in ecosystem functioning after the collapse of the top predator cod. We identified an ecological threshold, corresponding to a planktivore abundance of approximately 17 x 10(10) individuals, that separates 2 ecosystem configurations in which zooplankton dynamics are driven by either hydroclimatic forces or predation pressure. Abundances of the planktivore sprat above the threshold decouple zooplankton dynamics from hydrological circumstances. The current strong regulation by sprat of the feeding resources for larval cod may hinder cod recovery and the return of the ecosystem to a prior state. This calls for the inclusion of a food web perspective in management decisions.

QUANTITATIVE APPROACHES TO THE ANALYSIS OF STABLE ISOTOPE FOOD WEB DATA

Ecologists use stable isotopes (delta13C, delta15N) to better understand food webs and explore trophic interactions in ecosystems. Traditionally, delta13C vs. delta15N bi-plots have been used to describe food web structure for a single time period or ecosystem. Comparisons of food webs across time and space are increasing, but development of statistical approaches for testing hypotheses regarding food web change has lagged behind. Here we present statistical methodologies for quantitatively comparing stable isotope food web data. We demonstrate the utility of circular statistics and hypothesis tests for quantifying directional food web differences using two case studies: an arthropod salt marsh community across a habitat gradient and a freshwater fish community from Lake Tahoe, USA, over a 120-year time period. We calculated magnitude and mean angle of change (theta) for each species in food web space using mean delta13C and delta15N of each species as the x, y coordinates. In the coastal salt marsh, arthropod consumers exhibited a significant shift toward dependence on Spartina, progressing from a habitat invaded by Phragmites to a restored Spartina habitat. In Lake Tahoe, we found that all species from the freshwater fish community shifted in the same direction in food web space toward more pelagic-based production with the introduction of nonnative Mysis relicta and onset of cultural eutrophication. Using circular statistics to quantitatively analyze stable isotope food web data, we were able to gain significant insight into patterns and changes in food web structure that were not evident from qualitative comparisons. As more ecologists incorporate a food web perspective into ecosystem analysis, these statistical tools can provide a basis for quantifying directional food web differences from standard isotope data.

A Bioenergetics Based Comparison of Growth Conversion Efficiency of Atlantic Cod on Georges Bank and in the Gulf of Maine

Several authors state that foraging conditions and food web dynamics may be contributing to declines in Atlantic cod stocks. Therefore, it is essential to take a food web perspective to understand the complicated array of potential interactions affecting marine communities. The widely used Wisconsin Bioenergetics Model uses an energy-balance approach calculated on a daily time step and can be used to evaluate the effect of changes in prey resources on growth or food consumption. Bioenergetics models have been made for many important fishes within the Great Lakes. However, few have been developed for US Northeastern Continental Shelf fish species. Growth performance of Georges Bank and Gulf of Maine Atlantic cod during 2004 was examined using a bioenergetics model and available data on gut contents and growth of different ages. Growth efficiency, calculated from daily growth and consumption rates, was used as a measure of growth performance. Overall growth performance for Atlantic cod was significantly lower at Georges Bank than in Gulf of Maine. Monthly individual consumption demand and specific growth rates for Atlantic cod calculated from the bioenergetic model were significantly higher on Georges Bank than in the Gulf of Maine. Increasing water temperatures approached the upper limits of thermal tolerances for cod in Georges Bank, possibly leading to decreasing growth efficiencies. Growth efficiency of cod in the Gulf of Maine was less variable than on Georges Bank due to the more homogenous temperatures and energetic content of diets found in the former location.

Linking Landscapes and Food Webs: Effects of Omnivorous Fish and Watersheds on Reservoir Ecosystems

Ecologists increasingly recognize the need to understand how landscapes and food webs interact. Reservoir ecosystems are heavily subsidized by nutrients and detritus from surrounding watersheds, and often contain abundant populations of gizzard shad, an omnivorous fish that consumes plankton and detritus. Gizzard shad link terrestrial landscapes and pelagic reservoir food webs by consuming detritus, translocating nutrients from sediment detritus to the water column, and consuming zooplankton. The abundance of gizzard shad increases with watershed agriculturalization, most likely through a variety of mechanisms operating on larval and adult life stages. Gizzard shad have myriad effects on reservoirs, including impacts on nutrients, phytoplankton, zooplankton, and fish, and many of their effects vary with ecosystem productivity (i.e., watershed land use). Interactive feedbacks among watersheds, gizzard shad populations, and reservoir food webs operate to maintain dominance of gizzard shad in highly productive systems. Thus, effective stewardship of reservoir ecosystems must incorporate both watershed and food-web perspectives.

Water quality management in Lake Kinneret (Israel): hydrological and food web perspectives

Long term (1969-2001) data record of nutrient and plankton temporal distribution, and hydrological parameters in Lake Kinneret, combined with metabolic parameters of zooplankton, which were experimentally measured, were statistically (ANOVA) analyzed. Trophic relations between food web compartments were quantitatively considered to evaluate directional combination of ecological forces. Monthly data of inflow discharges, and lake volume were used to calculate residence time values and the data were incorporated into the ecological analysis. The seasonal fluctuations of the hydrological parameters, nutrients, and plankton inventories represent typical subtropical climate conditions: high level in winter and low in summer months. It was found that nitrogen inventories in the lake declined and the biomass of grazable phytoplankton was enhanced since early 1980’s. Dissolved phosphorus was decreased mostly in summer months when the lake is nutrient limited, as a result of phytoplankton uptake. Zooplankton was declined until 1993 and increased later. Zooplankton preferably feed on chlorophytes and diatoms with supplemental resources of detritus, bacteria and protozoa. The most abundant zooplanktivorous fish, Lavnun (Bleak, Acanthobrama spp.) populated the lake very densely during 1993-95 and biomanipulation management of subsidized fishery caused lowering of predation pressure resulted in zooplankton enhancement and suppression of additional primary produced matter. It is concluded that zooplankton in Lake Kinneret is not food limited and fishery management (Lavnun removal) might be efficient to enhance zooplankton grazing capacity and algal suppression if phosphorus flux is reduced. Long term changes of nano-phytoplankton are affected by both phosphorus availability and zooplankton grazing and fish predation has a significant impact on zooplankton density. Fishery management aimed at algal suppression might be efficient if phosphorus supply is reduced. The similarity between seasonal fluctuations of food web components and residence time is a result of the natural subtropical conditions. The major effect on nutrient loads is due to inputs from the catchment and to a lesser extend to internal processes. Consequently, management implications aimed at lake water quality protection should be mostly directed towards nutrient removal in the drainage basin and loads reduction by pumping water for supply. When such a management is successfully implemented biomanipulation (fish removal) might be efficient.

Insect-resistant transgenic plants in a multi-trophic context.

So far, genetic engineering of plants in the context of insect pest control has involved insertion of genes that code for toxins, and may be characterized as the incorporation of biopesticides into classical plant breeding. In the context of pesticide usage in pest control, natural enemies of herbivores have received increasing attention, because carnivorous arthropods are an important component of insect pest control. However, in plant breeding programmes, natural enemies of herbivores have largely been ignored, although there are many examples that show that plant breeding affects the effectiveness of biological control. Negative influences of modified plant characteristics on carnivorous arthropods may induce population growth of new, even more harmful pest species that had no pest status prior to the pesticide treatment. Sustainable pest management will only be possible when negative effects on non-target, beneficial arthropods are minimized. In this review, we summarize the effects of insect-resistant crops and insect-resistant transgenic crops, especially Bt crops, from a food web perspective. As food web components, we distinguish target herbivores, non-target herbivores, pollinators, parasitoids and predators. Below-ground organisms such as Collembola, nematodes and earthworms should also be included in risk assessment studies, but have received little attention. The toxins produced in Bt plants retain their toxicity when bound to the soil, so accumulation of these toxins is likely to occur. Earthworms ingest the bound toxins but are not affected by them. However, earthworms may function as intermediaries through which the toxins are passed on to other trophic levels. In studies where effects of insect-resistant (Bt) plants on natural enemies were considered, positive, negative and no effects have been found. So far, most studies have concentrated on natural enemies of target herbivores. However, Bt toxins are structurally rearranged when they bind to midgut receptors, so that they are likely to lose their toxicity inside target herbivores. What happens to the toxins in non-target herbivores, and whether these herbivores may act as intermediaries through which the toxins may be passed on to the natural enemies, remains to be studied.

The Responses of a Natural Bacterioplankton Community to Different Levels of Ultraviolet-B Radiation: A Food Web Perspective.

With the continuing increase of ultraviolet-B radiation (UVBR: 280-320 nm) fluxes toward the Earth's surface, there is concern regarding a possible negative impact on heterotrophic bacterioplankton. The effects of enhanced UVBR on a natural bacterioplankton community were studied during a 7-day experiment conducted in mesocosms (1500 L). Four light regimes were tested: natural light, 280 to 313 nm excluded UVBR, and two levels of UVBR enhancement. During the first 3 days of the experiment characterized by high inorganic nutrient concentrations (nitrates > 1 µmol L-1 and ammonium > 0.1 µmol L-l), UVBR had no effect on both bacterial abundances and activities. From day 4 to the end of the experiment, nitrate concentrations remained low (

Insect herbivory accelerates nutrient cycling and increases plant production.

Ecologists hold two views about the role of herbivory in ecosystem dynamics. First, from a food web perspective in population/community ecology, consumption by herbivores reduces plant abundance. Second, from a nutrient cycling perspective in ecosystem ecology, herbivory sometimes slows down cycling, which decreases plant abundance, but at other times speeds up cycling, which possibly increases plant abundance. The nutrient cycling perspective on herbivory has been experimentally addressed more thoroughly in aquatic systems than in terrestrial systems. We experimentally examined how grasshoppers influence nutrient cycling and, thereby, plant abundance and plant species composition over a period of 5 years. We examined how grasshoppers influence nutrient (nitrogen) cycling (i) by their excrement, (ii) by changing the abundance of and the decomposition rate of plant litter, and (iii) by both. Grasshoppers may speed up nitrogen cycling by changing the abundance and decomposition rate of plant litter, which increases total plant abundance (up to 32.9 g/m(2) or 18%), especially, the abundance of plants that are better competitors when nitrogen is more available. However, whether grasshoppers enhance plant abundance depends on how much they consume. Consequently, ecosystems and food web perspectives are not mutually exclusive. Finally, under some conditions, grasshoppers may decrease nutrient cycling and plant abundance.

Dams and Downstream Aquatic Biodiversity: Potential Food Web Consequences of Hydrologic and Geomorphic Change

/ Responses of rivers and river ecosystems to dams are complex and varied, as they depend on local sediment supplies, geomorphic constraints, climate, dam structure and operation, and key attributes of the biota. Therefore, "one-size-fits-all" prescriptions cannot substitute for local knowledge in developing prescriptions for dam structure and operation to protect local biodiversity. One general principle is self-evident: that biodiversity is best protected in rivers where physical regimes are the most natural. A sufficiently natural regime of flow variation is particularly crucial for river biota and food webs. We review our research and that of others to illustrate the ecological importance of alternating periods of low and high flow, of periodic bed scour, and of floodplain inundation and dewatering. These fluctuations regulate both the life cycles of river biota and species interactions in the food webs that sustain them. Even if the focus of biodiversity conservation efforts is on a target species rather than whole ecosystems, a food web perspective is necessary, because populations of any species depend critically on how their resources, prey, and potential predators also respond to environmental change. In regulated rivers, managers must determine how the frequency, magnitude, and timing of hydrologic events interact to constrain or support species and food webs. Simple ecological modeling, tailored to local systems, may provide a framework and some insight into explaining ecosystem response to dams and should give direction to mitigation efforts.KEY WORDS: Dams; Food webs; Hydrologic disturbance; Predator-prey dynamics; Succession

Effects of Disturbance on River Food Webs

A multitrophic model integrating the effects of flooding disturbance and food web interactions in rivers predicted that removing floods would cause increases of predator-resistant grazing insects, which would divert energy away from the food chain leading to predatory fish. Experimental manipulations of predator-resistant grazers and top predators, and large-scale comparisons of regulated and unregulated rivers, verified the model predictions. Thus, multitrophic models can successfully synthesize a variety of ecological processes, and conservation programs may benefit by taking a food web perspective instead of concentrating on a single species.