Rescaling the trophic structure of marine food webs

Measures of trophic position (TP) are critical for understanding food web interactions and human‐mediated ecosystem disturbance. Nitrogen stable isotopes (δ¹⁵N) provide a powerful tool to estimate TP but are limited by a pragmatic assumption that isotope discrimination is constant (change in δ¹⁵N between predator and prey, Δ¹⁵N = 3.4‰), resulting in an additive framework that omits known Δ¹⁵N variation. Through meta‐analysis, we determine narrowing discrimination from an empirical linear relationship between experimental Δ¹⁵N and δ¹⁵N values of prey consumed. The resulting scaled Δ¹⁵N framework estimated reliable TPs of zooplanktivores to tertiary piscivores congruent with known feeding relationships that radically alters the conventional structure of marine food webs. Apex predator TP estimates were markedly higher than currently assumed by whole‐ecosystem models, indicating perceived food webs have been truncated and species‐interactions over simplified. The scaled Δ¹⁵N framework will greatly improve the accuracy of trophic estimates widely used in ecosystem‐based management.     

Top‐down and bottom‐up diversity cascades in detrital vs. living food webs

Apex predators and plant resources are both critical for maintaining diversity in biotic communities, but the indirect (‘cascading’) effects of top‐down and bottom‐up forces on diversity at different trophic levels are not well resolved in terrestrial systems. Manipulations of predators or resources can cause direct changes of diversity at one trophic level, which in turn can affect diversity at other trophic levels. The indirect diversity effects of resource and consumer variation should be strongest in aquatic systems, moderate in terrestrial systems, and weakest in decomposer food webs. We measured effects of top predators and plant resources on the diversity of endophytic animals in an understorey shrub Piper cenocladum (Piperaceae). Predators and resource availability had significant direct and indirect effects on the diversity of the endophytic animal community, but the effects were not interactive, nor were they consistent between living vs. detrital food webs. The addition of fourth trophic level beetle predators increased diversity of consumers supported by living plant tissue, whereas balanced plant resources (light and nutrients) increased the diversity of primary through tertiary consumers in the detrital resources food web. These results support the hypotheses that top‐down and bottom‐up diversity cascades occur in terrestrial systems, and that diversity is affected by different factors in living vs. detrital food webs.     

Consequences of consumer origin and omnivory on stability in experimental food web modules

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A modified niche model for generating food webs with stage‐structured consumers: The stabilizing effects of life‐history stages on complex food webs

1. Almost all organisms grow in size during their lifetime and switch diets, trophic positions, and interacting partners as they grow. Such ontogenetic development introduces life‐history stages and flows of biomass between the stages through growth and reproduction. However, current research on complex food webs rarely considers life‐history stages. The few previously proposed methods do not take full advantage of the existing food web structural models that can produce realistic food web topologies.
2. We extended the niche model developed by Williams and Martinez (Nature, 2000, 404, 180–183) to generate food webs that included trophic species with a life‐history stage structure. Our method aggregated trophic species based on niche overlap to form a life‐history structured population; therefore, it largely preserved the topological structure of food webs generated by the niche model. We applied the theory of allometric predator–prey body mass ratio and parameterized an allometric bioenergetic model augmented with biomass flow between stages via growth and reproduction to study the effects of a stage structure on the stability of food webs.
3. When life‐history stages were linked via growth and reproduction, more food webs persisted, and persisting food webs tended to retain more trophic species. Topological differences between persisting linked and unlinked food webs were small to modest. The slopes of biomass spectra were lower, and weak interaction links were more prevalent in the linked food webs than the unlinked ones, suggesting that a life‐history stage structure promotes characteristics that can enhance stability of complex food webs.
4. Our results suggest a positive relationship between the complexity and stability of complex food webs. A life‐history stage structure in food webs may play important roles in dynamics of and diversity in food webs.

     

Phylogeny versus body size as determinants of food web structure

Food webs are the complex networks of trophic interactions that stoke the metabolic fires of life. To understand what structures these interactions in natural communities, ecologists have developed simple models to capture their main architectural features. However, apparently realistic food webs can be generated by models invoking either predator-prey body-size hierarchies or evolutionary constraints as structuring mechanisms. As a result, this approach has not conclusively revealed which factors are the most important. Here we cut to the heart of this debate by directly comparing the influence of phylogeny and body size on food web architecture. Using data from 13 food webs compiled by direct observation, we confirm the importance of both factors. Nevertheless, phylogeny dominates in most networks. Moreover, path analysis reveals that the size-independent direct effect of phylogeny on trophic structure typically outweighs the indirect effect that could be captured by considering body size alone. Furthermore, the phylogenetic signal is asymmetric: closely related species overlap in their set of consumers far more than in their set of resources. This is at odds with several food web models, which take only the view-point of consumers when assigning interactions. The echo of evolutionary history clearly resonates through current food webs, with implications for our theoretical models and conservation priorities.     

Uncovering trophic positions and food resources of soil animals using bulk natural stable isotope composition

Despite the major importance of soil biota in nutrient and energy fluxes, interactions in soil food webs are poorly understood. Here we provide an overview of recent advances in uncovering the trophic structure of soil food webs using natural variations in stable isotope ratios. We discuss approaches of application, normalization and interpretation of stable isotope ratios along with methodological pitfalls. Analysis of published data from temperate forest ecosystems is used to outline emerging concepts and perspectives in soil food web research. In contrast to aboveground and aquatic food webs, trophic fractionation at the basal level of detrital food webs is large for carbon and small for nitrogen stable isotopes. Virtually all soil animals are enriched in ¹³C as compared to plant litter. This ‘detrital shift’ likely reflects preferential uptake of ¹³C‐enriched microbial biomass and underlines the importance of microorganisms, in contrast to dead plant material, as a major food resource for the soil animal community. Soil organic matter is enriched in ¹⁵N and ¹³C relative to leaf litter. Decomposers inhabiting mineral soil layers therefore might be enriched in ¹⁵N resulting in overlap in isotope ratios between soil‐dwelling detritivores and litter‐dwelling predators. By contrast, ¹³C content varies little between detritivores in upper litter and in mineral soil, suggesting that they rely on similar basal resources, i.e. little decomposed organic matter. Comparing vertical isotope gradients in animals and in basal resources can be a valuable tool to assess trophic interactions and dynamics of organic matter in soil. As indicated by stable isotope composition, direct feeding on living plant material as well as on mycorrhizal fungi is likely rare among soil invertebrates. Plant carbon is taken up predominantly by saprotrophic microorganisms and channelled to higher trophic levels of the soil food web. However, feeding on photoautotrophic microorganisms and non‐vascular plants may play an important role in fuelling soil food webs. The trophic niche of most high‐rank animal taxa spans at least two trophic levels, implying the use of a wide range of resources. Therefore, to identify trophic species and links in food webs, low‐rank taxonomic identification is required. Despite overlap in feeding strategies, stable isotope composition of the high‐rank taxonomic groups reflects differences in trophic level and in the use of basal resources. Different taxonomic groups of predators and decomposers are likely linked to different pools of organic matter in soil, suggesting different functional roles and indicating that trophic niches in soil animal communities are phylogenetically structured. During last two decades studies using stable isotope analysis have elucidated the trophic structure of soil communities, clarified basal food resources of the soil food web and revealed links between above‐ and belowground ecosystem compartments. Extending the use of stable isotope analysis to a wider range of soil‐dwelling organisms, including microfauna, and a larger array of ecosystems provides the perspective of a comprehensive understanding of the structure and functioning of soil food webs.     

Trophic coupling of the microbial and the classical food web in Lake Baikal,Siberia

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Sensory cues of a top‐predator indirectly control a reef fish mesopredator

Behavioural trophic cascades highlight the importance of indirect/risk effects in the maintenance of healthy trophic‐level links in complex ecosystems. However, there is limited understanding on how the loss of indirect top–down control can cascade through the food‐web to modify lower level predator–prey interactions. Using a reef fish food‐web, our study examines behavioural interactions among predators to assess how fear elicited by top‐predator cues (visual and chemical stimuli) can alter mesopredator behaviour and modify their interaction with resource prey. Under experimental conditions, the presence of any cue (visual, chemical, or both) from the top‐predator (coral trout Plectropomus leopardus) strongly restricted the distance swum, area explored and foraging activity of the mesopredator (dottyback Pseudochromis fuscus), while indirectly triggering a behavioural release of the resource prey (recruits of the damselfish Pomacentrus chrysurus). Interestingly, the presence of a large non‐predator species (thicklip wrasse Hemigymnus melapterus) also mediated the impact of the mesopredator on prey, as it provoked mesopredators to engage in an ‘inspection’ behaviour, while significantly reducing their feeding activity. Our study describes for the first time a three‐level behavioural cascade of coral reef fish and stresses the importance of indirect interactions in marine food‐webs.     

Boosted food web productivity through ocean acidification collapses under warming

Future climate is forecast to drive bottom‐up (resource driven) and top‐down (consumer driven) change to food web dynamics and community structure. Yet, our predictive understanding of these changes is hampered by an over‐reliance on simplified laboratory systems centred on single trophic levels. Using a large mesocosm experiment, we reveal how future ocean acidification and warming modify trophic linkages across a three‐level food web: that is, primary (algae), secondary (herbivorous invertebrates) and tertiary (predatory fish) producers. Both elevated CO₂ and elevated temperature boosted primary production. Under elevated CO₂, the enhanced bottom‐up forcing propagated through all trophic levels. Elevated temperature, however, negated the benefits of elevated CO₂ by stalling secondary production. This imbalance caused secondary producer populations to decline as elevated temperature drove predators to consume their prey more rapidly in the face of higher metabolic demand. Our findings demonstrate how anthropogenic CO₂ can function as a resource that boosts productivity throughout food webs, and how warming can reverse this effect by acting as a stressor to trophic interactions. Understanding the shifting balance between the propagation of resource enrichment and its consumption across trophic levels provides a predictive understanding of future dynamics of stability and collapse in food webs and fisheries production.     

Predicting food‐web structure with metacommunity models

Synthesis Metacommunity theory aims to elucidate the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity research has focused largely on assemblages of competing organisms within a single trophic level. Here, we test the ability of metacommunity models to predict the network structure of the aquatic food web found in the leaves of the northern pitcher plant Sarracenia purpurea. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions play an important role in structuring Sarracenia food webs. Our approach can be applied to any well‐resolved food web for which data are available from multiple locations. The metacommunity framework explores the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity models and empirical studies have focused mostly on assemblages of competing organisms within a single trophic level. Studies of multi‐trophic metacommunities are predominantly restricted to simplified trophic motifs and rarely consider entire food webs. We tested the ability of the patch‐dynamics, species‐sorting, mass‐effects, and neutral metacommunity models, as well as three hybrid models, to reproduce empirical patterns of food web structure and composition in the complex aquatic food web found in the northern pitcher plant Sarracenia purpurea. We used empirical data to determine regional species pools and estimate dispersal probabilities, simulated local food‐web dynamics, dispersed species from regional pools into local food webs at rates based on the assumptions of each metacommunity model, and tested their relative fits to empirical data on food‐web structure. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions were important in structuring Sarracenia food webs. However, differences in dispersal abilities were also important in models that accurately reproduced empirical food web properties. Although the models were tested using pitcher‐plant food webs, the approach we have developed can be applied to any well‐resolved food web for which data are available from multiple locations.     

The influence of a parasitoid's response to temperature on the performance of a tri‐trophic food web

1. Food web interactions are complex and can respond to environmental changes in unpredictable ways that do not necessarily equate to the individual responses of each of the components of the food web. 2. Biomass can be used to evaluate the productivity of the three individual trophic levels, in the form of the Net Generational Productivity (NGP) and the performance of the entire food web with the newly developed tri‐trophic food web performance ratio (?_3t). 3. These parameters were used to evaluate the performance of nine plant‐based tri‐trophic food webs composed of: potato, Solanum tuberosum L. and two cultivars of bell pepper Capsicum annuum L; three biotypes of the aphid Macrosiphum euphorbiae Thomas (Hemiptera: Aphididae) and the parasitoid wasp Aphidius ervi (Haliday) (Hymenoptera: Braconidae). 4. The NGP showed that the thermal window for biomass productivity for each trophic level is different and is reduced by approximately 4 °C with respect to the inferior level. Aphidius ervi had the smallest thermal window for biomass productivity and development. 5. The present results showed that the performance (?_3t) of the tested food webs is influenced in a top‐down fashion, where the intra‐specific variation of the food web, namely the host plant, played a major role in the productivity of each of the subsequent trophic levels. 6. The ?_3t suggested that exposure to high and low temperatures might severely affect the effectiveness of A. ervi as a biocontrol agent of the aphid M. euphorbiae in bell pepper and potato crops.     

Organic matter sources and size structuring in stream invertebrate food webs across a tropical to temperate gradient

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Food web complexity and higher-level ecosystem services

Studies mostly focused on communities of primary producers have shown that species richness provides and promotes fundamental ecosystem services. However, we know very little about the factors influencing ecosystem services provided by higher trophic levels in natural food webs. Here we present evidence that differences in food web structure and the richness of herbivores in 19 plant‐herbivore‐parasitoid food webs influence the service supplied by natural enemies, namely, the parasitism rates on hosts. Specifically, we find that parasitoids function better in simple food webs than in complex ones, a result relevant to biological control practice. More generally, we show that species richness per se only contributes partially to the understanding of higher‐level ecosystem services in multitrophic communities, and that changes in food web complexity should also be taken into account when predicting the effects of human‐driven disturbances in natural communities.     

Rewiring and indirect effects underpin modularity reshuffling in a marine food web under environmental shifts

Species are characterized by physiological and behavioral plasticity, which is part of their response to environmental shifts. Nonetheless, the collective response of ecological communities to environmental shifts cannot be predicted from the simple sum of individual species responses, since co‐existing species are deeply entangled in interaction networks, such as food webs. For these reasons, the relation between environmental forcing and the structure of food webs is an open problem in ecology. To this respect, one of the main problems in community ecology is defining the role each species plays in shaping community structure, such as by promoting the subdivision of food webs in modules—that is, aggregates composed of species that more frequently interact—which are reported as community stabilizers. In this study, we investigated the relationship between species roles and network modularity under environmental shifts in a highly resolved food web, that is, a “weighted” ecological network reproducing carbon flows among marine planktonic species. Measuring network properties and estimating weighted modularity, we show that species have distinct roles, which differentially affect modularity and mediate structural modifications, such as modules reconfiguration, induced by environmental shifts. Specifically, short‐term environmental changes impact the abundance of planktonic primary producers; this affects their consumers’ behavior and cascades into the overall rearrangement of trophic links. Food web re‐adjustments are both direct, through the rewiring of trophic‐interaction networks, and indirect, with the reconfiguration of trophic cascades. Through such “systemic behavior,” that is, the way the food web acts as a whole, defined by the interactions among its parts, the planktonic food web undergoes a substantial rewiring while keeping almost the same global flow to upper trophic levels, and energetic hierarchy is maintained despite environmental shifts. This behavior suggests the potentially high resilience of plankton networks, such as food webs, to dramatic environmental changes, such as those provoked by global change.     

Ecological emergence of thermal clines in body size

The unprecedented rate of global warming requires a better understanding of how ecosystems will respond. Organisms often have smaller body sizes under warmer climates (Bergmann's rule and the temperature‐size rule), and body size is a major determinant of life histories, demography, population size, nutrient turnover rate, and food‐web structure. Therefore, by altering body sizes in whole communities, current warming can potentially disrupt ecosystem function and services. However, the underlying drivers of warming‐induced body downsizing remain far from clear. Here, we show that thermal clines in body size are predicted from universal laws of ecology and metabolism, so that size‐dependent selection from competition (both intra and interspecific) and predation favors smaller individuals under warmer conditions. We validate this prediction using 4.1 × 10⁶ individual body size measurements from French river fish spanning 29 years and 52 species. Our results suggest that warming‐induced body downsizing is an emergent property of size‐structured food webs, and highlight the need to consider trophic interactions when predicting biosphere reorganizations under global warming.     

Detecting trophic‐level variation in consumer assemblages

1. Stable isotopes of nitrogen are useful for quantifying the trophic structure of food webs, but only if the variation in trophic enrichment (Δ_N), which is the difference in δ¹⁵N between a consumer and its food, is small relative to the value of Δ_N itself. 2. We examined the sources of variation in zooplankton Δ_N by measuring the trophic enrichment (Δ_N) of seven species of freshwater cladocerans, and by testing for an effect of age and temperature on the Δ_N of Daphnia pulicaria. 3. We found that Δ_N was similar among Cladocera and was not correlated with body size. Overall, the Δ_N for D. pulicaria was 1.4‰ (SE = 0.69, n = 57), as was expected for the detritus diet that we used in our experiments. We found no effect of temperature (15–25 °C) on Δ_N, but found that Δ_N of D. pulicaria increased with increasing age (10–30 days). 4. We developed a new method to test for trophic‐level variation in a group of consumers that explicitly accounts for the uncertainty in Δ_N. Using this approach, we confirmed that natural assemblages of zooplankton feed at several trophic levels in lake food webs.     

Aquatic food‐web structure along a salinized dryland river

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Energetically relevant predator–prey body mass ratios and their relationship with predator body size

Food web structure and dynamics depend on relationships between body sizes of predators and their prey. Species‐based and community‐wide estimates of preferred and realized predator–prey mass ratios (PPMR) are required inputs to size‐based size spectrum models of marine communities, food webs, and ecosystems. Here, we clarify differences between PPMR definitions in different size spectrum models, in particular differences between PPMR measurements weighting prey abundance in individual predators by biomass (r^bio) and numbers (r^num). We argue that the former weighting generates PPMR as usually conceptualized in equilibrium (static) size spectrum models while the latter usually applies to dynamic models. We use diet information from 170,689 individuals of 34 species of fish in Alaskan marine ecosystems to calculate both PPMR metrics. Using hierarchical models, we examine how explained variance in these metrics changed with predator body size, predator taxonomic resolution, and spatial resolution. In the hierarchical analysis, variance in both metrics emerged primarily at the species level and substantially less variance was associated with other (higher) taxonomic levels or with spatial resolution. This suggests that changes in species composition are the main drivers of community‐wide mean PPMR. At all levels of analysis, relationships between weighted mean r^bio or weighted mean r^num and predator mass tended to be dome‐shaped. Weighted mean r^num values, for species and community‐wide, were approximately an order of magnitude higher than weighted mean r^bio, reflecting the consistent numeric dominance of small prey in predator diets. As well as increasing understanding of the drivers of variation in PPMR and providing estimates of PPMR in the north Pacific Ocean, our results demonstrate that that r^bio or r^num, as well as their corresponding weighted means for any defined group of predators, are not directly substitutable. When developing equilibrium size‐based models based on bulk energy flux or comparing PPMR estimates derived from the relationship between body mass and trophic level with those based on diet analysis, weighted mean r^bio is a more appropriate measure of PPMR. When calibrating preference PPMR in dynamic size spectrum models then weighted mean r^num will be a more appropriate measure of PPMR.