Defining and measuring trophic role similarity in food webs using regular equivalence

We present a graph theoretic model of analysing food web structure called regular equivalence. Regular equivalence is a method for partitioning the species in a food web into "isotrophic classes" that play the same structural roles, even if they are not directly consuming the same prey or if they do not share the same predators. We contrast regular equivalence models, in which two species are members of the same trophic group if they have trophic links to the same set of other trophic groups, with structural equivalence models, in which species are equivalent if they are connected to the exact same other species. Here, the regular equivalence approach is applied to two published food webs: (1) a topological web (Malaysian pitcher plant insect food web) and (2) a carbon-flow web (St. Marks, Florida seagrass ecosystem food web). Regular equivalence produced a more satisfactory set of classes than did the structural approach, grouping basal taxa with other basal taxa and not with top predators. Regular equivalence models provide a way to mathematically formalize trophic position, trophic group and trophic niche. These models are part of a family of models that includes structural models used extensively by ecologists now. Regular equivalence models uncover similarities in trophic roles at a higher level of organization than do the structural models. The approach outlined is useful for measuring the trophic roles of species in food web models, measuring similarity in trophic relations of two or more species, comparing food webs over time and across geographic regions, and aggregating taxa into trophic groups that reduce the complexity of ecosystem feeding relations without obscuring network relationships. In addition, we hope the approach will prove useful in predicting the outcome of predator-prey interactions in experimental studies.     

Forecasting fine‐scale changes in the food‐web structure of coastal marine communities under climate change

Climate change is inducing deep modifications in local communities worldwide as a consequence of individualistic species range shifts. Understanding how complex interaction networks will be reorganized under climate change represents a major challenge in the fields of ecology and biogeography. However, forecasting the potential effects of climate change on local communities, and more particularly on food‐web structure, requires the consideration of highly structuring processes, such as trophic interactions. A major breakthrough is therefore expected by combining predictive models integrating habitat selection processes, the physiological limits of marine species and their trophic interactions. In this study, we forecasted the potential impacts of climate change on the local food‐web structure of the highly threatened Gulf of Gabes ecosystem located in the south of the Mediterranean Sea. We coupled the climatic envelope and habitat models to an allometric niche food web model, hence taking into account the different processes acting at regional (climate) and local scales (habitat selection and trophic interactions). Our projections under the A2 climate change scenario showed that future food webs would be composed of smaller species with fewer links, resulting in a decrease of connectance, generality, vulnerability and mean trophic level of communities and an increase of the average path length, which may have large consequences on ecosystem functioning. The unified framework presented here, by connecting food‐web ecology, biogeography and seascape ecology, allows the exploration of spatial aspects of interspecific interactions under climate change and improves our current understanding of climate change impacts on local marine food webs.     

Integrating ecosystem engineering and food webs

Ecosystem engineering, the physical modification of the environment by organisms, is a common and often influential process whose significance to food web structure and dynamics is largely unknown. In the light of recent calls to expand food web studies to include non‐trophic interactions, we explore how we might best integrate ecosystem engineering and food webs. We provide rationales justifying their integration and present a provisional framework identifying how ecosystem engineering can affect the nodes and links of food webs and overall organization; how trophic interactions with the engineer can affect the engineering; and how feedbacks between engineering and trophic interactions can affect food web structure and dynamics. We use a simple integrative food chain model to illustrate how feedbacks between the engineer and the food web can alter 1) engineering effects on food web dynamics, and 2) food web responses to extrinsic environmental perturbations. We identify four general challenges to integration that we argue can readily be met, and call for studies that can achieve this integration and help pave the way to a more general understanding of interaction webs in nature. Synthesis All species are affected by their physical environment. Because ecosystem engineering species modify the physical environment and belong to food webs, such species are potentially one of the most important bridges between the trophic and non‐trophic. We examine how to integrate the so far, largely independent research areas of ecosystem engineering and food webs. We present a conceptual framework for understanding how engineering can affect food webs and vice versa, and how feedbacks between the two alter ecosystem dynamics. With appropriate empirical studies and models, integration is achievable, paving the way to a more general understanding of interaction webs in nature.     

Food-web structure and ecosystem services: insights from the Serengeti

The central organizing theme of this paper is to discuss the dynamics of the Serengeti grassland ecosystem from the perspective of recent developments in food-web theory. The seasonal rainfall patterns that characterize the East African climate create an annually oscillating, large-scale, spatial mosaic of feeding opportunities for the larger ungulates in the Serengeti; this in turn creates a significant annual variation in the food available for their predators. At a smaller spatial scale, periodic fires during the dry season create patches of highly nutritious grazing that are eaten in preference to the surrounding older patches of less palatable vegetation. The species interactions between herbivores and plants, and carnivores and herbivores, are hierarchically nested in the Serengeti food web, with the largest bodied consumers on each trophic level having the broadest diets that include species from a large variety of different habitats in the ecosystem. The different major habitats of the Serengeti are also used in a nested fashion; the highly nutritious forage of the short grass plains is available only to the larger migratory species for a few months each year. The longer grass areas, the woodlands and kopjes (large partially wooded rocky islands in the surrounding mosaic of grassland) contain species that are resident throughout the year; these species often have smaller body size and more specialized diets than the migratory species. Only the larger herbivores and carnivores obtain their nutrition from all the different major habitat types in the ecosystem. The net effect of this is to create a nested hierarchy of subchains of energy flow within the larger Serengeti food web; these flows are seasonally forced by rainfall and operate at different rates in different major branches of the web. The nested structure that couples sequential trophic levels together interacts with annual seasonal variation in the fast and slow chains of nutrient flow in a way that is likely to be central to the stability of the whole web. If the Serengeti is to be successfully conserved as a fully functioning ecosystem, then it is essential that the full diversity of natural habitats be maintained within the greater Serengeti ecosystem. The best way to do this is by controlling the external forces that threaten the boundaries of the ecosystem and by balancing the economic services the park provides between local, national and international needs. I conclude by discussing how the ecosystem services provided by the Serengeti are driven by species on different trophic levels. Tourism provides the largest financial revenue to the national economy, but it could be better organized to provide more sustained revenue to the park. Ultimately, ecotourism needs to be developed in ways that take lessons from the structure of the Serengeti food webs, and in ways that provide tangible benefits to people living around the park while also improving the experience of all visitors.     

Food web structure in a tropical stream ecosystem

Abstract This study investigated the structure and properties of a tropical stream food web in a small spatial scale, characterizing its planktonic, epiphytic and benthic compartments. The study was carried out in the Potreirinho Creek, a second‐order stream located in the south‐east of Brazil. Some attributes of the three subwebs and of the conglomerate food web, composed by the trophic links of the three compartments plus the fish species, were determined. Among compartments, the food webs showed considerable variation in structure. The epiphytic food web was consistently more complex than the planktonic and benthic webs. The values of number of species, number of links and maximum food chain length were significantly higher in the epiphytic compartment than in the other two. Otherwise, the connectance was significantly lower in epiphyton. The significant differences of most food web parameters were determined by the increase in the number of trophic species, represented mainly by basal and intermediate species. High species richness, detritus‐based system and high degree of omnivory characterized the stream food web studied. The aquatic macrophytes probably provide a substratum more stable and structurally complex than the sediment. We suggest that the greater species richness and trophic complexity in the epiphytic subweb might be due to the higher degree of habitat complexity supported by macrophyte substrate. Despite differences observed in the structure of the three subwebs, they are highly connected by trophic interactions, mainly by fishes. The high degree of fish omnivory associated with their movements at different spatial scales suggests that these animals have a significant role in the food web dynamic of Potreirinho Creek. This interface between macrophytes and the interconnections resultant from fish foraging, diluted the compartmentalization of the Potreirinho food web.     

Food‐web structure and ecosystem function in the Laurentian Great Lakes—Toward a conceptual model

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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.     

Complexity and structural properties of food webs in the Barents Sea

A food web topology describes the diversity of species and their trophic interactions, i.e. who eats whom, and structural analysis of food web topologies can provide insight into ecosystem structure and function. It appears simple, at first sight, to list all species and their trophic interactions. However, the very large number of species at low trophic levels and the impossibility to monitor all trophic interactions in the ocean makes it impossible to construct complete food web topologies. In practice, food web topologies are simplified by aggregating species into groups termed trophospecies. It is not clear though, how much simplified versions of food webs retain the structural properties of more detailed networks. Using the most comprehensive Barents Sea food web to date, we investigate the performance of methods to construct simplified food webs using three approaches: taxonomic, structural and regular clustering. We then evaluate how topological properties vary with the level of network simplification. Results show that alteration of food web structural properties due to aggregation are highly sensitive to the methodology used for grouping species and trophic links. In the specific case of the Barents Sea, we show that it is possible to preserve key structural properties of the original complex food web in simplified versions when using taxonomic or structural clustering combined with intermediate 25% linkage for trophic aggregation.     

Trophic disruption: a meta‐analysis of how habitat fragmentation affects resource consumption in terrestrial arthropod systems

Habitat fragmentation is a complex process that affects ecological systems in diverse ways, altering everything from population persistence to ecosystem function. Despite widespread recognition that habitat fragmentation can influence food web interactions, consensus on the factors underlying variation in the impacts of fragmentation across systems remains elusive. In this study, we conduct a systematic review and meta‐analysis to quantify the effects of habitat fragmentation and spatial habitat structure on resource consumption in terrestrial arthropod food webs. Across 419 studies, we found a negative overall effect of fragmentation on resource consumption. Variation in effect size was extensive but predictable. Specifically, resource consumption was reduced on small, isolated habitat fragments, higher at patch edges, and neutral with respect to landscape‐scale spatial variables. In general, resource consumption increased in fragmented settings for habitat generalist consumers but decreased for specialist consumers. Our study demonstrates widespread disruption of trophic interactions in fragmented habitats and describes variation among studies that is largely predictable based on the ecological traits of the interacting species. We highlight future prospects for understanding how changes in spatial habitat structure may influence trophic modules and food webs.     

The Serengeti food web: empirical quantification and analysis of topological changes under increasing human impact

1.?To address effects of land use and human overexploitation on wildlife populations, it is essential to better understand how human activities have changed species composition, diversity and functioning. Theoretical studies modelled how network properties change under human-induced, non-random species loss. However, we lack data on realistic species-loss sequences in threatened, real-world food webs to parameterize these models. 2.?Here, we present a first size-structured topological food web of one of the most pristine terrestrial ecosystems in the world, the Serengeti ecosystem (Tanzania). The food web consists of 95 grouped nodes and includes both invertebrates and vertebrates ranging from body masses between 10(-7) and 10(4) kg. 3.?We study the topological changes in this food web that result from the simulated IUCN-based species-loss sequence representing current species vulnerability to human disturbances in and around this savanna ecosystem. We then compare this realistic extinction scenario with other extinction sequences based on body size and connectance and perform an analysis of robustness of this savanna food web. 4.?We demonstrate that real-world species loss in this case starts with the biggest (mega) herbivores and top predators, causing higher predator-prey mass ratios. However, unlike theoretically modelled linear species deletion sequences, this causes poor-connected species to be lost first, while more highly connected species become lost as human impact progresses. This food web shows high robustness to decreasing body size and increasing connectance deletion sequences compared with a high sensitivity to the decreasing connectance deletion scenario. 5.?Furthermore, based on the current knowledge of the Serengeti ecosystem, we discuss how the focus on food web topology alone, disregarding nontrophic interactions, may lead to an underestimation of human impacts on wildlife communities, with the number of trophic links affected by a factor of two. 6.?This study underlines the importance of integrative efforts between the development of food web theory and basic field work approaches in the quantification of the structure of interaction networks to sustain natural ecosystems in a changing world.     

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.     

Invasions cause biodiversity loss and community simplification in vertebrate food webs

Global change is increasing the occurrence of perturbation events on natural communities, with biological invasions posing a major threat to ecosystem integrity and functioning worldwide. Most studies addressing biological invasions have focused on individual species or taxonomic groups to understand both, the factors determining invasion success and their effects on native species. A more holistic approach that considers multispecies communities and species’ interactions can contribute to a better understanding of invasion effects on complex communities. Here we address biological invasions on species‐rich food webs. We performed in silico experiments on empirical vertebrate food webs by introducing virtual species characterised by different ecological roles and belonging to different trophic groups. We varied a number of invasive species traits, including their diet breadth, the number of predators attacking them, and the bioenergetic thresholds below which invader and native species become extinct. We found that simpler food webs were more vulnerable to invasions, and that relatively less connected mammals were the most successful invaders. Invasions altered food web structure by decreasing species richness and the number of links per species, with most extinctions affecting poorly connected birds. Our food web approach allows identifying the combinations of trophic factors that facilitate or prevent biological invasions, and it provides testable predictions on the effects of invasions on the structure and dynamics of multitrophic communities.     

Interplay between the paradox of enrichment and nutrient cycling in food webs

Nutrient cycling is fundamental to ecosystem functioning. Despite recent major advances in the understanding of complex food web dynamics, food web models have so far generally ignored nutrient cycling. However, nutrient cycling is expected to strongly impact food web stability and functioning. To make up for this gap, we built an allometric and size structured food web model including nutrient cycling. By releasing mineral nutrients, recycling increases the availability of limiting resources for primary producers and links each trophic level to the bottom of food webs. We found that nutrient cycling can provide a significant part of the total nutrient supply of the food web, leading to a strong enrichment effect that promotes species persistence in nutrient poor ecosystems but leads to a paradox of enrichment at high nutrient inputs. The presence of recycling loops linking each trophic level to the basal resources weakly affects species biomass temporal variability in the food web. Recycling loops tend to slightly dampen the destabilising effect of nutrient enrichment on consumer temporal variability while they have opposite effects for primary producers. By considering nutrient cycling, this new model improves our understanding of the response of food webs to nutrient availability and opens perspectives to better link studies on food web dynamics and ecosystem functioning.     

Food web modularity and biodiversity promote species persistence in polluted environments

Pollution represents a major threat to biodiversity. A wide class of pollutants tends to accumulate within organisms and propagate within communities via trophic interactions. Thus the final effects of accumulable pollutants may be determined by the structure of food webs and not only by the susceptibility of their constituent species. Species within real food webs are typically arranged into modules, which have been proposed to be determinants of network stability. In this study we evaluate the effect of network modularity and species richness on long‐term species persistence in communities perturbed by pollutant stress. We built model food webs with different levels of modularity and used a bioenergetic model to project the dynamics of species. Further, we modeled the dynamics of bioaccumulated and environmental pollutants. We found that modularity promoted the stability of food webs subjected to pollutant stress. We also found that richer food webs were more robust at all modularity levels. Nevertheless, modularity did not promote stability of communities facing a perturbation that shared most features with the pollutant perturbation, but does not spread through trophic interactions. The positive effect of both modularity and species richness on species persistence was cancelled and even reversed when the structure of food web departed from a realistic body size distribution or a hierarchical feeding structure. Our results support the idea that modularity implies important dynamic consequences for communities facing pollution, highlighting a main role of network structure on ecosystem stability.     

The effects of energy input,immigration and habitat size on food web structure: a microcosm experiment

It has been hypothesised that larger habitats should support more complex food webs. We consider three mechanisms which could lead to this pattern. These are increased immigration rates, increased total productivity and spatial effects on the persistence of unstable interactions. Experiments designed to discriminate between these mechanisms were carried out in laboratory aquatic microcosm communities of protista and bacteria, by independently manipulating habitat size, total productivity and immigration rate. Larger habitats supported more complex food webs, with more species, more links per species and longer maximum and mean food chains, even in the absence of differences in total energy input. Increased immigration rate resulted in more complex food webs, but habitats with higher energy input per unit area supported less complex food webs. We conclude that spatial effects on the persistence of unstable interactions, and variation in immigration rates, are plausible mechanisms by which habitat size could affect food web structure. Variation in total productivity with habitat area seems a less likely explanation for variation in food web structure.     

稻田节肢动物群落的营养联系

根据田间调查和室内饲养观察的资料,研究了稻田节肢动物群落的营养结构及类型。在稻田生态系统中,物种之间由于取食与被取食、寄生与被寄生、捕食与被捕食的营养联系,形成了复杂的食物链和食物网。依据物种在食物网中的位置和功能,可将福州市郊区稻田节肢动物群落的营养结构分为3种类型:1)食物网中尚未发现有重寄生环节;2)食物网中有重寄生环节;3)食物网中有兼寄生环节。为了探讨定量研究生物群落营养联系的可能性,本文运用图论的知识把食物网的结构描述为标向图、集合或邻接矩阵,同时用图论的运算法则解决了各种类型的食物网的合并问题,为研究复杂群落的营养关系提供了一种新方法。     

How habitat-modifying organisms structure the food web of two coastal ecosystems

The diversity and structure of ecosystems has been found to depend both on trophic interactions in food webs and on other species interactions such as habitat modification and mutualism that form non-trophic interaction networks. However, quantification of the dependencies between these two main interaction networks has remained elusive. In this study, we assessed how habitat-modifying organisms affect basic food web properties by conducting in-depth empirical investigations of two ecosystems: North American temperate fringing marshes and West African tropical seagrass meadows. Results reveal that habitat-modifying species, through non-trophic facilitation rather than their trophic role, enhance species richness across multiple trophic levels, increase the number of interactions per species (link density), but decrease the realized fraction of all possible links within the food web (connectance). Compared to the trophic role of the most highly connected species, we found this non-trophic effects to be more important for species richness and of more or similar importance for link density and connectance. Our findings demonstrate that food webs can be fundamentally shaped by interactions outside the trophic network, yet intrinsic to the species participating in it. Better integration of non-trophic interactions in food web analyses may therefore strongly contribute to their explanatory and predictive capacity.     

What Can Interaction Webs Tell Us About Species Roles?

The group model is a useful tool to understand broad-scale patterns of interaction in a network, but it has previously been limited in use to food webs, which contain only predator-prey interactions. Natural populations interact with each other in a variety of ways and, although most published ecological networks only include information about a single interaction type (e.g., feeding, pollination), ecologists are beginning to consider networks which combine multiple interaction types. Here we extend the group model to signed directed networks such as ecological interaction webs. As a specific application of this method, we examine the effects of including or excluding specific interaction types on our understanding of species roles in ecological networks. We consider all three currently available interaction webs, two of which are extended plant-mutualist networks with herbivores and parasitoids added, and one of which is an extended intertidal food web with interactions of all possible sign structures (+/+, -/0, etc.). Species in the extended food web grouped similarly with all interactions, only trophic links, and only nontrophic links. However, removing mutualism or herbivory had a much larger effect in the extended plant-pollinator webs. Species removal even affected groups that were not directly connected to those that were removed, as we found by excluding a small number of parasitoids. These results suggest that including additional species in the network provides far more information than additional interactions for this aspect of network structure. Our methods provide a useful framework for simplifying networks to their essential structure, allowing us to identify generalities in network structure and better understand the roles species play in their communities.     

Major food web properties of two sandy beaches with contrasting morphodynamics, and effects on the stability

We determined major structural properties influencing the food webs of two sandy beaches with contrasting morphodynamics in the Atlantic coast of Uruguay: reflective (narrow and steep) and dissipative beaches (wide and flat). Furthermore, we evaluated how these characteristics could influence the stability of the local food webs. To this end, we examined the correlation of several food web properties with different ecosystem types (including freshwater habitats, estuary, marine, and terrestrial environments) using a principal components analysis. Sandy beach food web components included detritus, phytoplankton, zooplankton, benthic invertebrates, fishes, and seabirds. Our results revealed that the dissipative beach presented higher trophic levels, a higher number of trophic species, more links per species, as well as a higher proportion of intermediate trophic species, but lower connectance and proportion of omnivorous species than the reflective beach. The variation in the food web properties was explained by two principal components. Sandy beach food webs contribute mainly to one dimension of the principal components analysis that was determined by the number of trophic species, links per species, the trophic similarity, and the characteristic path length. We suggest that species and link characteristics, such as predominance of scavengers and detritivorous, the relatively high connectance and the short path length are drivers in the food web structure and may play a role in the community dynamic.