Published Food Webs Research Articles

A food web including parasites for kelp forests of the Santa Barbara Channel, California

We built a high-resolution topological food web for the kelp forests of the Santa Barbara Channel, California, USA that includes parasites and significantly improves resolution compared to previous webs. The 1,098 nodes and 21,956 links in the web describe an economically, socially, and ecologically vital system. Nodes are broken into life-stages, with 549 free-living life-stages (492 species from 21 Phyla) and 549 parasitic life-stages (450 species from 10 Phyla). Links represent three kinds of trophic interactions, with 9,352 predator-prey links, 2,733 parasite-host links and 9,871 predator-parasite links. All decisions for including nodes and links are documented, and extensive metadata in the node list allows users to filter the node list to suit their research questions. The kelp-forest food web is more species-rich than any other published food web with parasites, and it has the largest proportion of parasites. Our food web may be used to predict how kelp forests may respond to change, will advance our understanding of parasites in ecosystems, and fosters development of theory that incorporates large networks.

Quantifying the primary biotic resource use by fisheries: A global assessment

In this paper, the specific primary production required (SPPR expressed as kg-NPP/kg-fish in wet weight) of more than 1700 marine species were calculated directly from 96 published food web models using the newly developed SPPR calculation framework. The relationship between SPPR and other ecological factors were then statistically analyzed. Among- and within-species variability of SPPR were found to be both explained by trophic level (TL), suggesting similar mechanisms underpinning both sources of variability. Among species, we found that harvesting species at higher mean trophic levels (MTL) increases the mean SPPR by a factor of 19 per 1 unit increase in MTL. Based on our empirical relationship, the mean SPPR of more than 9000 marine species were predicted and subsequently used to assess the primary production required (PPR) to support fisheries in five major fishing countries in Europe. The results indicated that conventional approach to estimating PPR, which neglects food web ecology, can underestimate PPR by up to a factor of 5. Within species, we found that harvesting populations occupying a higher TL leads to a higher SPPR. For example, the SPPR of Atlantic cod in the Celtic Sea (TL = 4.75) was 5 times higher than in the Gilbert Bay (TL = 3.3). Our results, which are based on large amounts of field data, highlight the importance of properly accounting for ecological factors during the impact assessment of fisheries.

Water level fluctuations and the ecosystem functioning of lakes

Abstract Hydrological regimes are key drivers of productivity and structure in freshwater ecosystems but are increasingly impacted by human activity. Using 17 published food web models of 13 African lakes as a case study, we explored relationships between seasonal and interannual water level fluctuations and 15 attributes related to ecosystem function. We interpreted our results in the context of Odum's ecosystem maturity hypothesis, as systems with higher magnitude fluctuations may be kept at an earlier maturity stage than those that are relatively stable. The data we compiled indicate that long-term changes in the hydrological regimes of African lakes have already taken place. We used Least Absolute Shrinkage and Selection Operator (LASSO) regression to examine relationships between ecosystem attributes and seven physical characteristics. Of these characteristics, interannual water level fluctuation magnitude was the most frequently retained predictor in the regression models. Our results indicate that interannual water level fluctuations are positively correlated with primary and overall production, but negatively correlated with fish diversity, transfer efficiency, and food chain length. These trends are opposite those expected with increasing ecosystem maturity. Interestingly, we found seasonal water level fluctuations to be positively correlated with biomass. An increase in standing biomass is generally associated with more mature ecosystems. However, we found that less production and biomass occurred at high trophic levels in highly fluctuating compared to relatively stable systems. This synthesis provides evidence that water level fluctuations are a key process influencing ecosystem structure and function in lakes.

What's in a tide pool? Just as much food web network complexity as in large open ecosystems.

Understanding the fundamental laws that govern complex food web networks over large ecosystems presents high costs and oftentimes unsurmountable logistical challenges. This way, it is crucial to find smaller systems that can be used as proxy food webs. Intertidal rock pool environments harbour particularly high biodiversity over small areas. This study aimed to analyse their food web networks to investigate their potential as proxies of larger ecosystems for food web networks research. Highly resolved food webs were compiled for 116 intertidal rock pools from cold, temperate, subtropical and tropical regions, to ensure a wide representation of environmental variability. The network properties of these food webs were compared to that of estuaries, lakes and rivers, as well as marine and terrestrial ecosystems (46 previously published complex food webs). The intertidal rock pool food webs analysed presented properties that were in the same range as the previously published food webs. The niche model predictive success was remarkably high (73–88%) and similar to that previously found for much larger marine and terrestrial food webs. By using a large-scale sampling effort covering 116 intertidal rock pools in several biogeographic regions, this study showed, for the first time, that intertidal rock pools encompass food webs that share fundamental organizational characteristics with food webs from markedly different, larger, open and abiotically stable ecosystems. As small, self-contained habitats, intertidal rock pools are particularly tractable systems and therefore a large number of food webs can be examined with relatively low sampling effort. This study shows, for the first time that they can be useful models for the understanding of universal processes that regulate the complex network organization of food webs, which are harder or impossible to investigate in larger, open ecosystems, due to high costs and logistical difficulties.

Revisiting the Energetic Efficiency Hypothesis: Body Mass, Metabolism, and Food Chain Length

Abstract There is a longstanding interest in describing the factors determining food chain length in ecological communities. Likewise, there has been considerable interest in explaining why food chain lengths differ in terrestrial and aquatic systems. The ‘energetic efficiency hypothesis' predicts endothermy and body size should be negatively correlated with food chain length. Using a large database of published food webs, we assessed the relationships between food chain length, endothermy, body size, and food-web structure. We utilized 77 published food webs spanning marine, freshwater, and terrestrial systems, ranging in species richness from 8–224 taxa. As expected, mean food chain lengths (MFCLs) in aquatic systems were significantly longer than those of terrestrial systems, even when comparing food webs of similar size and species richness. Endotherms were more numerous and occurred lower in terrestrial food chains compared to aquatic ones. Endotherms were also consumed by fewer predators and fed on ...

Pyramids of species richness: the determinants and distribution of species diversity across trophic levels

How species richness is distributed across trophic levels determines several dimensions of ecosystem functioning, including herbivory, predation, and decomposition rates. We perform a meta‐analysis of 72 large published food webs to investigate their trophic diversity structure and possible endogenous, exogenous, and methodological causal variables. Consistent with classic theory, we found that published food webs can generally be described as ‘pyramids of species richness’. The food webs were more predator‐poor, prey‐rich and hierarchical than is expected by chance or by the niche or cascade models. The trophic species richness distribution also depended on centrality, latitude, ecosystem‐type and methodological bias. Although trophic diversity structure is generally pyramidal, under many conditions the structure is consistently uniform or inverse‐pyramidal. Our meta‐analysis adds nuance to classic assumptions about food web structure: diversity decreases with trophic level, but not under all conditions, and the decrease may be scale‐dependent.SynthesisThe distribution of species richness across trophic levels has not been evaluated in recent decades, despite improvement in food web resolution and the relevance of biodiversity distribution to ecosystem function. Our meta‐analysis of 72 large, recent food webs, illustrates that published food webs can generally be described as basal‐rich, top‐poor ‘pyramids of species richness’, consistent with classic theory. Although trophic diversity structure is generally pyramidal, under some environmental and ecological conditions the structure is uniform or inverse‐pyramidal. Our meta‐analysis confirms classic theory about food web structure, while adding nuance by describing conditions under which classic pyramid structure is not observed.

Joining the dots: An automated method for constructing food webs from compendia of published interactions

Food webs are important tools for understanding how complex natural communities are structured and how they respond to environmental change. However their full potential has yet to be realised because of the huge amount of resources required to construct them de novo. Consequently, the current catalogue of networks that are suitable for rigorous and comparative analyses and theoretical development still suffers from a lack of standardisation and replication.Here, we present a novel R function, WebBuilder, which automates the construction of food webs from taxonomic lists, and a dataset of trophic interactions. This function works by matching species against those within a dataset of trophic interactions, and ‘filling in’ missing trophic interactions based on these matches. We also present a dataset of over 20,000 freshwater trophic interactions, and use this and four well-characterised freshwater food webs to test the method.The WebBuilder function facilitates the generation of food webs of comparable quality to the most detailed published food webs, but at a fraction of the research effort or cost. Furthermore, it matched and often outperformed a selection of predictive models, which are currently among the best, in terms of capturing key properties of empirical food webs. The method is simple to use, systematic and, perhaps most importantly, reproducible, which will facilitate (re-) analysis and data sharing. Although developed and tested on a sample of freshwater food webs, this method could easily be extended to cover other types of ecological interactions (such as mutualistic interactions).

Current opinions: Zeros in host–parasite food webs: Are they real?

As the data have poured in, and the number of published food webs containing parasites has increased, questions have been raised as to why free-living species consistently outnumber parasites, even though most general reviews on the subject of host:parasite species richness suggest the contrary. Here, I describe this pattern as it exists in the literature, posit both real and artifactual sources of these findings, and suggest ways that we might interpret existing parasite-inclusive food webs. In large part, the reporting of free-living species devoid of any associated parasites (termed here in the coding of food web matrices as "zeros") is a consequence of either sampling issues or the intent of the study. However, there are also several powerful explanatory features that validate real cases of this phenomenon. Some hosts appear to authentically lack parasitism in portions of their geographic ranges, and parasites are often lost from systems that are either in early phases of community re-colonization or are compromised by environmental perturbation. Additionally, multi-stage parasite life cycles and broad host spectra allow some parasite species to partially saturate systems without providing a corresponding increase in parasite species richness, leading to low parasite species richness values relative to the free-living community. On the whole, the existing published food webs are sufficient to, at least in principle, determine basic patterns and pathways associated with parasite establishment and persistence in free-living communities because (1) for the purpose of those features, species rarity is roughly analogous to absence and (2) the existing data seem to suggest that the addition of more parasite taxa would reinforce the patterns already observed. This is particularly true for helminth parasites, in which our understanding and the resolution of our work is most robust.

The reduction of food web robustness by parasitism: Fact and artefact

A robust food web is one which suffers few secondary extinctions after primary species losses. While recent research has shown that a food web with parasitism is less robust than one without, it still remains unclear whether the reduction in robustness is due to changes in network complexity or unique characteristics associated with parasitism. Here, using several published food webs, simulation experiments with different food web models and extinction scenarios were conducted to elucidate how such reduction can be achieved. Our results show that, regardless of changes in network complexity and preferential parasitism, the reduction in food web robustness is mainly due to the life cycle constraint of parasites. Our findings further demonstrate that parasites are prone to secondary extinctions and that their extinctions occur earlier than those involving free-living species. These findings suggest that the vulnerable nature of parasites to species loss makes them highly sensitive indicators of food web integrity.

Food webs are built up with nested subwebs

Nested structure, in which specialists interact with subsets of species with which generalists interact, has been repeatedly found in networks of mutualistic interactions and thus is considered a general feature of mutualistic communities. However, it is uncertain how exclusive nested structure is for mutualistic communities since few studies have evaluated nestedness in other types of networks. Here, we show that 31 published food webs consist of bipartite subwebs that are as highly nested as mutualistic networks, contradicting the hypothesis that antagonistic interactions disfavor nested structure. Our findings suggest that nested networks may be a common pattern of communities that include resource-consumer interactions. In contrast to the hypothesis that nested structure enhances biodiversity in mutualistic communities, we also suggest that nested food webs increase niche overlap among consumers and thus prevent their coexistence. We discuss potential mechanisms for the emergence of nested structure in food webs and other types of ecological networks.

Exploitation of asymmetric predator–prey interactions by trophically transmitted parasites

The directionality of asymmetric interactions between predators (definitive hosts) and prey (intermediate hosts) should impact trophic transmission in parasites. This study tests the prediction that trophically transmitted parasites are funneled towards asymmetric predator–prey interactions where intermediate hosts have few predators and definitive hosts feed upon many prey (‘downward asymmetry’). The distribution of trophically transmitted parasites was examined in four published food webs in relation to mismatch asymmetry of predator–prey interactions. We found that trophically transmitted parasites exploit downwardly asymmetric interactions in a nonrandom manner, and particular predator–prey pairs contain more trophically transmitted parasites than would be expected by random chance alone. These findings suggest that food web topology has great bearing on the ecology of trophically transmitted parasites, and that consideration of parasite life cycles in the context of food web organization can provide insights into the forces affecting the evolution of trophic transmission.

Major dimensions in food‐web structure properties

The covariance among a range of 20 network structural properties of food webs plus net primary productivity was assessed for 14 published food webs using principal components analysis. Three primary components explained 84% of the variability in the data sets, suggesting substantial covariance among the properties employed in the literature. The first dimension explained 48% of the variance and could be ascribed to connectance, covarying significantly with the proportion of intermediate species and characteristic path length. The second dimension explained 19% and was related to trophic species richness. The third axis explained 17% and was related to ecosystem net primary productivity. A distinct opposite clustering of connectance, the proportion of intermediate species, and mean trophic level vs. the proportion of top and basal species and path length suggests a dichotomy in food-web structure. Food webs appear either clustered and highly interconnected or elongated with fewer links.

Are network indices robust indicators of food web functioning? A Monte Carlo approach

Indices based on network theory are often used to describe food web functioning. These indices take as input food web flows that are estimated based on merging of (scarce) data with linear inverse methods (LIMs). Due to under sampling, most food webs are highly uncertain and can only be quantified within a specific uncertainty range. The linear inverse method (LIM) can estimate food web flows using a variety of techniques, e.g. the parsimonious or minimum norm (MN) solution, which selects one food web, based on a quadratic minimization technique or the Monte Carlo solution where a finitely many random solutions are generated which are then averaged. We use the Monte Carlo approach (MCA) to estimate the values of several indices from four published food webs, the Gulf of Riga for the autumn, summer and spring seasons, and the Takapoto atoll system. We first show that network indices are much better constrained than the uncertain food webs from which they are calculated. Therefore, even in the face of food web uncertainty, they are robust estimators of food web functioning. We then use the MCA-derived network indices to generate cumulative density functions for each index. These serve to compute the probabilities of the MN indices estimates being an extreme solution as compared to the median values. Our findings show that 82% of the MN solutions are smaller than the MCA solutions, and 63% of the network indices are significantly under-estimated.

Parasites alter the topology of a stream food web across seasons

Relatively few published food webs have included parasites, and in this study we examined the animal community in a stream across eight contiguous seasons to test how inclusion of helminth parasites alters the topology or structure of the food web. Food webs constructed for each season and analyzed using common binary matrix measures show that species richness, linkage density, and the number of observed and possible links increased when parasites were included as individual species nodes. With parasite-parasite and predator-parasite links omitted, measures of community complexity, such as connectance (C), generally increased over multiple seasons. However, relative nestedness (n*) decreased when parasites were included, which may be a result of low resolution of basal resources inflating specialist-to-specialist links. Overall, adding parasites resulted in moderate changes in food web measures when compared to those of four other published food webs representing different ecosystems. In addition, including parasites in the food web revealed consistent pathways of energy flow, and the association of parasite life histories along these pathways suggest stable evolutionary groups of interacting species within the community.

Gaining insight into food webs reconstructed by the inverse method

The use of the inverse method to analyze flow patterns of organic components in ecological systems has had wide application in ecological modeling. Through this approach, an infinite number of food web flows describing the food web and satisfying biological constraints are generated, from which one (parsimonious) solution is drawn. Here we address two questions: (1) is there justification for the use of the parsimonious solution or is there a better alternative and (2) can we use the infinitely many solutions that describe the same food web to give more insight into the system? We reassess two published food webs, from the Gulf of Riga in the Baltic Sea and the Takapoto Atoll lagoon in the South Pacific. A finite number of random food web solutions is first generated using the Monte Carlo simulation technique. Using the Wilcoxon signed ranks test, we cannot find significant differences in the parsimonious solution and the average values of the finite random solutions generated. However, as the food web composed of the average flows has more attractive properties, the choice of the parsimonious solution to describe underdetermined food webs is challenged. We further demonstrate the use of the factor analysis technique to characterize flows that are closely related in the food web. Through this process sub-food webs are extracted within the plausible set of food webs, a property that can be utilized to gain insight into the sampling strategy for further constraining of the model.

Modeling the impact of oyster culture on a mudflat food web in Marennes-Oléron Bay (France)

We used a carbon-based food web model to investigate the effects of oyster cultivation on the ecosystem of an intertidal mudflat. A previously published food web model of a mudflat in Marennes-Oleron Bay, France, was updated with revised parameters, and a realistic surface area and density of existing oyster cultures on the mudflat. We developed 2 hypothetical scenarios to estimate the impact of oyster cultivation on the food web structure of the ecosystem: one with no oysters, the other with a doubled area devoted to cultivated oysters in the bay. Oysters are direct trophic competitors of other filter feeders, and their presence modifies benthic-pelagic coupling by forcing a shift from pelagic consumers to benthic consumers. Increasing the surface area of cultivated oysters caused secondary production to increase, providing food for top predators (in particular juvenile nekton), reinforcing the nursery role of the mudflat in the ecosystem, and altering the species composition available to the top predators.

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.

Connectance in stream food webs

Summary The properties of food webs are important both in theoretical ecology and environmental management, yet remain elusive. Here, we examined 12 new stream food webs of higher taxonomic resolution and completeness than any previously published data set and combined them with other 10 published stream webs. Compared with most previously published food webs, these stream communities (containing between 22 and 212 species) had more feeding links per species, a higher fraction of intermediate species (0·78–0·88) and lower fractions of top (0·04–0·09) and ‘basal’ species (0·07–0·15). The exponent of the log link‐log species relationship (S1·3) differed significantly (ancova: P < 0·001) from the link‐species ‘law’ (S1) and the constant connectance hypothesis (S2). Based on these results, the link scaling law and the constant connectance hypothesis must be rejected for food webs in stream systems. Connectance was markedly lower than previously reported and decreased with web size. We attribute this to the body‐size disparity of organisms at the top and the bottom of the web, flow disturbance and to the high physical heterogeneity and complexity of the sedimentary habitat. All these may reduce the fraction of possible feeding links that are realized.

Predators, parasitoids and pathogens: species richness, trophic generality and body sizes in a natural food web

Summary 1. A food web is presented which describes trophic interactions among the herbivores, parasitoids, predators and pathogens associated with broom, Cytisus scoparius (L.) Link. The data come from published work on the community at a single site. The web comprises a total of 154 taxa: one plant, 19 herbivores, 66 parasitoids, 60 predators, five omnivores and three pathogens. There are 370 trophic links between these taxa in the web. The taxa form 82 functionally distinct groups, called trophic species. 2. Predators consumed significantly more species than did parasitoids: a median of two prey species per species of predator (range = 1–9), compared to a median of one host species per species of parasitoid (range = 1–4). Significant differences in the number of species consumed were also found among the five predator groups: birds (median = 4), spiders (median = 5), Coleoptera (median = 1), Diptera (median = 2) and Hemiptera (median = 7). 3. Vulnerability, measured by numbers of consumer species, was significantly affected by the herbivores’ feeding styles: externally feeding herbivores were most vulnerable and the concealed herbivores were least vulnerable. Miners were vulnerable to the most parasitoid species and externally feeding herbivores were the most vulnerable to predators. 4. Resource species had a median vulnerability of 13 consumer species, a figure far higher than that in most published food webs. No significant relationship was found between species’ vulnerability to predators and vulnerability to parasitoids. However, there was a strong negative relationship between the percentage mortality due to predation and percentage mortality due to parasitism. 5. The broom food web contains nine orders of insects, a figure higher than previously recorded. The web also contains vertebrates, arachnids, bacteria and fungi. Most of the interactions between the orders were weak. Connectance was calculated for the complete web, the parasitoid sub‐web and the predator sub‐web. The connectance of the predator sub‐web, a value of 0·0364, was more than an order of magnitude larger than the connectance of the entire web (0·0156) or the parasitoid sub‐web (0·018). 6. The body lengths of 52 species in the food web were estimated from field guides or museum specimens. Larger predators consumed smaller prey in 93% of predator–prey interactions. Smaller parasitoids consumed larger hosts in 79% of parasitoid–host interactions. Parasitoids were significantly smaller than predators. 7. The 52 species were arranged in order of increasing body length along the columns and down the rows of a food web matrix. The predator sub‐web was predominantly upper triangular with 8% of non‐zero elements falling below the leading diagonal. The parasitoid sub‐web was predominantly lower triangular with 21% non‐zero elements falling above the leading diagonal. The entire web contains entries both above and below the main diagonal and thus violates a central assumption of the cascade model.

Effects of taxonomic and trophic aggregation on food web properties.

Historically, ecologists have been more interested in organisms feeding at the tops of food chains than in organisms feeding at or near the bottom. The problem of taxonomic and trophic inconsistency within and among described food webs is central to criticisms of contemporary food web research. To study the effects of taxonomic and trophic aggregation on food web properties, 38 published food webs, each containing a large fraction of investigator-defined biological species, were systematically aggregated by taxonomy and trophic (functional) group similarity. During each step of taxonomic and trophic aggregation, eight food web properties (MIN, MAX, mean chain lengths; the fractions of basal, intermediate and top species; the ratio of all links by the total number of species, L/S; and rigid circuits) were calculated and their departures from the original, unaggregated version were recorded. We found only two␣properties showing wide systematic departure from initial values after both taxonomic and trophic group aggregation: the fraction of basal species and L/S. One reason for the relative `constancy' of the six other properties was due in part to large numbers of trophically equivalent species (species with identical sets of prey and predators) found in these and other published webs. In the 38 webs, the average number of trophically equivalent species was 45% and ranged from a low of 13% in aquatic webs to a high of 71% in certain terrestrial systems (i.e., carrion webs). Six of the eight properties (MIN, MAX and mean chain lengths, the fractions of top and basal species, and the L/S ratio) were found to be more sensitive to taxonomic than to trophic aggregation. The relatively smaller variations observed in trophically lumped versions suggest that food web properties more aptly reflect functional, rather than taxonomic, attributes of real food webs. These findings parallel earlier trophic-based results, and bolster the conclusion that uneven lumping of taxonomic and trophic groups in published food web reports do not modify markedly the scaling behaviour of most of their descriptive properties.