Threshold extinction in food webs

Food web response to species loss has been investigated in several ways in the previous years. In binary food webs, species go secondarily extinct if no resource item remains to be exploited. In this work, we considered that species can go extinct before the complete loss of their resources and we introduced thresholds of minimum energy requirement for species survival. According to this approach, extinction of a node occurs whenever an initial extinction event eliminates its incoming links so it is left with an overall energy intake lower than the threshold value. We tested the robustness of 18 real food webs by removing species from most to least connected and considering different scenarios defined by increasing the extinction threshold. Increasing energy requirement threshold negatively affects food web robustness. We found that a very small increase of the energy requirement substantially increases system fragility. In addition, above a certain value of energy requirement threshold we found no relationship between the robustness and the connectance of the web. Further, food webs with more species showed higher fragility with increasing energy threshold. This suggests that the shape of the robustness–complexity relationship of a food web depends on the sensitivity of consumers to loss of prey.     

Current food web models cannot explain the overall topological structure of observed food webs

Topological food webs illustrating “who eats whom” in different systems exhibit similar, non‐random, structures suggesting that general rules govern food web structure. Current food web models correctly predict many measures of food web topology from knowledge of species richness and connectance (fraction of possible predator–prey links that actually occur), together with assumptions about the ecological rules governing “who eats whom”. However, current measures are relatively insensitive to small changes in topology. Improvement of, and discrimination among, current models requires development of new measures of food web structure. Here I examine whether current food web models (cascade, niche, and nested hierarchy models, plus a random null model) can predict a new measure of food web structure, structural stability. Structural stability complements other measures of food web topology because it is sensitive to changes in topology that other measures often miss. The cascade and null models respectively over‐ and underpredict structural stability for a set of 17 high‐quality food webs. While the niche and nested hierarchy models provide unbiased predictions on average, their 95% confidence intervals frequently fail to include the observed data. Observed structural stabilities for all models are overdispersed compared to model predictions, and predicted and observed structural stabilities are uncorrelated, indicating that important sources of variation in structural stability are not captured by the models. Crucially, poor model performance arises because observed variation in structural stability is unrelated to variation in species richness and connectance. In contrast, almost all other measures of food web topology vary with species richness and connectance in natural webs. No model that takes species richness and connectance as the only input parameters can reproduce observed variation in structural stability. Further progress in predicting and explaining food web topology will require fundamentally new models based on different input parameters.     

Global synthesis suggests that food web connectance correlates to invasion resistance

Biological invasions are a key component of global change, and understanding the drivers of global invasion patterns will aid in assessing and mitigating the impact of invasive species. While invasive species are most often studied in the context of one or two trophic levels, in reality species invade communities comprised of complex food webs. The complexity and integrity of the native food web may be a more important determinant of invasion success than the strength of interactions between a small subset of species within a larger food web. Previous efforts to understand the relationship between food web properties and species invasions have been primarily theoretical and have yielded mixed results. Here, we present a synthesis of empirical information on food web connectance and species invasion success gathered from different sources (estimates of food web connectance from the primary literature and estimates of invasion success from the Global Invasive Species Database as well as the primary literature). Our results suggest that higher‐connectance food webs tend to host fewer invaders and exert stronger biotic resistance compared to low‐connectance webs. We argue that while these correlations cannot be used to infer a causal link between food web connectance and habitat invasibility, the promising findings beg for further empirical research that deliberately tests for relationships between food web connectance and invasion.     

Food web stability and weighted connectance: the complexity-stability debate revisited

How the complexity of food webs relates to stability has been a subject of many studies. Often, unweighted connectance is used to express complexity. Unweighted connectance is measured as the proportion of realized links in the network. Weighted connectance, on the other hand, takes link weights (fluxes or feeding rates) into account and captures the shape of the flux distribution. Here, we used weighted connectance to revisit the relation between complexity and stability. We used 15 real soil food webs and determined the feeding rates and the interaction strength matrices. We calculated both versions of connectance, and related these structural properties to food web stability. We also determined the skewness of both flux and interaction strength distributions with the Gini coefficient. We found no relation between unweighted connectance and food web stability, but weighted connectance was positively correlated with stability. This finding challenges the notion that complexity may constrain stability, and supports the ‘complexity begets stability’ notion. The positive correlation between weighted connectance and stability implies that the more evenly flux rates were distributed over links, the more stable the webs were. This was confirmed by the Gini coefficients of both fluxes and interaction strengths. However, the most even distributions of this dataset still were strongly skewed towards small fluxes or weak interaction strengths. Thus, incorporating these distribution with many weak links via weighted instead of unweighted food web measures can shed new light on classical theories.     

Persistence increases with diversity and connectance in trophic metacommunities

Background

We are interested in understanding if metacommunity dynamics contribute to the persistence of complex spatial food webs subject to colonization-extinction dynamics. We study persistence as a measure of stability of communities within discrete patches, and ask how do species diversity, connectance, and topology influence it in spatially structured food webs.

Methodology/Principal Findings

We answer this question first by identifying two general mechanisms linking topology of simple food web modules and persistence at the regional scale. We then assess the robustness of these mechanisms to more complex food webs with simulations based on randomly created and empirical webs found in the literature. We find that linkage proximity to primary producers and food web diversity generate a positive relationship between complexity and persistence in spatial food webs. The comparison between empirical and randomly created food webs reveal that the most important element for food web persistence under spatial colonization-extinction dynamics is the degree distribution: the number of prey species per consumer is more important than their identity.

Conclusions/Significance

With a simple set of rules governing patch colonization and extinction, we have predicted that diversity and connectance promote persistence at the regional scale. The strength of our approach is that it reconciles the effect of complexity on stability at the local and the regional scale. Even if complex food webs are locally prone to extinction, we have shown their complexity could also promote their persistence through regional dynamics. The framework we presented here offers a novel and simple approach to understand the complexity of spatial food webs.     

Connectance determines invasion success via trophic interactions in model food webs

Human induced global change has greatly altered the structure and composition of food webs through the invasion of non‐native species and the extinction of native species. Much attention has been paid to the effects of species deletions on food web structure and stability. However, recent empirical evidence suggests that for most taxa local species richness has increased as successful invasions outpace extinctions at this scale. This pattern suggests that food webs, which represent feeding interactions at the local scale, may be increasing in species richness. Knowledge of how food web structure relates to invasive species establishment and the effect of successful invaders on subsequent food web structure remains an unknown but potentially important aspect of global change. Here we explore the effect of food web topology on invasion success in model food webs to develop hypotheses about how the distribution of biodiversity across trophic levels affects the success of invasion at each trophic level. Our results suggest a connectance (C) based framework for predicting invasion success in food webs due to the way that C constrains the number of species at each trophic level and thus the number of potential predators and prey for an invader at a given trophic level. We use the relationship between C and the proportion of species at each trophic level in 14 well studied food webs to make the following predictions; 1) the success of basal invaders will increase as C increases due to the decrease in herbivores in high C webs, 2) herbivore invasion success will decrease as C increases due to the decrease in the proportion of basal species and increase in intermediate species and omnivores in high C webs. 3) Top predator invasion success will increase as C increases due to the increase in intermediate prey species. However, it is not clear how the relative influence of trophic structure compares to empirically known predictors of invasion success such as invader traits, propagule pressure, and resource availability.     

Will a large complex system be productive?

While the relationship between food web complexity and stability has been well documented, how complexity affects productivity remains elusive. In this study, we combine food web theory and a data set of 149 aquatic food webs to investigate the effect of complexity (i.e. species richness, connectance, and average interaction strength) on ecosystem productivity. We find that more complex ecosystems tend to be more productive, although different facets of complexity have contrasting effects. A higher species richness and/or average interaction strength increases productivity, whereas a higher connectance often decreases it. These patterns hold not only between realized complexity and productivity, but also characterize responses of productivity to simulated declines of complexity. Our model also predicts a negative association between productivity and stability along gradients of complexity. Empirical analyses support our predictions on positive complexity-productivity relationships and negative productivity-stability relationships. Our study provides a step forward towards reconciling ecosystem complexity, productivity and stability.     

Estimating morphologically determined connectance and structure for food webs of freshwater invertebrates

1. Connectance is a parameter of central importance in determining food-web structure, but the processes determining its value remain unclear. In evaluating possible explanations it is useful to know what patterns, and values, of connectance occur in food webs assembled at random from a set of species in a regional species pool; i.e. where the number of links is determined by the morphological features of the species present, not by the immediate effects of energetics or stability on the particular web. 2. This study examines, by means of laboratory experiments, the occurrence of potential feeding interactions among a set of freshwater invertebrate species randomly selected from different freshwater sites in a geographical region. The results from pairwise feeding trials are used to construct two ‘theoretical’ food webs, in which the patterns and values of connectance are examined. 3. Analyses of these webs indicate that their structure is consistent with the observed values in previously documented ‘real’ webs. Directed connectance values of 0.12–0.16 (or less) suggest that the assembled webs are no more connected than many freshwater webs from natural systems. The number of links per species increases curvilinearly with the number of species, during web assembly, consistent with recent hypotheses. 4. These results also indicate that quantifying, and understanding the determinants of, trophic generalism or specialism does have implications for understanding how connectance is constrained in real webs. Freshwater invertebrates seem to be relatively generalist, and freshwater food webs perhaps correspondingly highly connected. Such arguments have implications for interpreting other aspects of food-web structure in these systems, and for parameterizing models that are based on connectance.     

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.     

The structure of food webs along river networks

Do changes in the species composition of riverine fish assemblages along river networks lead to predictable changes in food‐web structure? We assembled empirical “fish‐centered” river food webs for three rivers located along a latitudinal gradient in the South Saskatchewan River Basin (SSRB) that differ in land‐use impacts and geomorphology but flow through similar mountain, foothill, and prairie physiographic regions. We then calculated 17 food‐web properties to determine whether the nine river food webs differed according to physiographic region or river sub‐basin. There were no statistically significant differences in the 17 food‐web properties calculated among the rivers. In contrast, fish species richness, connectance, the proportion of herbivores, and the proportion of cannibals changed longitudinally along the river network. Our results suggest that regional changes in river geomorphology and physicochemistry play an important role in determining longitudinal variation in food‐web properties such as fish species richness and connectance. In contrast, the overall structure of river food webs may be relatively similar and insensitive to regional influences such as zoogeography. Further explorations of river and other food webs would greatly illuminate this suggestion.     

水生生态系统食物网复杂性与多样性的关系

探索食物网的复杂结构是生态学的中心问题之一。基于构建的黄河口海草床食物网并耦合实际食物网的数据集，整理了包含河口、湖泊、海洋和河流四种水生生态系统类型的48个实际食物网案例。以食物网的节点数反映食物网多样性，物种之间的营养链接数、链接密度和连通度来表示食物网的复杂性，采用营养缩尺模型描述水生生态系统食物网的复杂性特征与节点数的普适性规律。结果表明：所涉及的48个水生生态系统食物网的多样性和复杂性跨度较大，其中，节点数的分布范围为4-124，链接数为3-1830，链接密度为0.75-15.71，连通度为0.06-0.25。不同类型水生生态系统间的连通度存在显著性差异（P=0.01），节点数、链接数、链接密度不存在显著性差异。各类型生态系统的食物网链接数、链接密度均随节点数的增加而增加（R²=0.92，P<0.001和R²=0.82，P<0.001）。湖泊生态系统的连通度随节点数的变化不明显，围绕在0.20附近；而其他3种类型生态系统的食物网连通度随节点数的增加而降低（R²=0.06-0.41，P<0.001）。对全球尺度的水生食物网多样性和复杂性的定量化研究对于提升对食物网的复杂结构的科学认识，从系统尺度探究多样性和复杂性的关系提供数据支撑。     

Predicting the effects of temperature on food web connectance

Few models concern how environmental variables such as temperature affect community structure. Here, we develop a model of how temperature affects food web connectance, a powerful driver of population dynamics and community structure. We use the Arrhenius equation to add temperature dependence of foraging traits to an existing model of food web structure. The model predicts potentially large temperature effects on connectance. Temperature-sensitive food webs exhibit slopes of up to 0.01 units of connectance per 1°C change in temperature. This corresponds to changes in diet breadth of one resource item per 2°C (assuming a food web containing 50 species). Less sensitive food webs exhibit slopes down to 0.0005, which corresponds to about one resource item per 40°C. Relative sizes of the activation energies of attack rate and handling time determine whether warming increases or decreases connectance. Differences in temperature sensitivity are explained by differences between empirical food webs in the body size distributions of organisms. We conclude that models of temperature effects on community structure and dynamics urgently require considerable development, and also more and better empirical data to parameterize and test them.     

Food webs robustness to biodiversity loss: the roles of connectance, expansibility and degree distribution

We analyse the robustness of food webs against species loss by considering the influence of several structural factors of the networks, such as connectance, degree distribution and expansibility. The last concept refers to the absence of structural bottlenecks in the food web, whose removal separate the network into large isolate clusters. In theory networks with identical connectance can display different expansibility characteristics. Using the spectral scaling method we studied 17 food networks and classified them as good expansion (GE) and not-GE networks. The combination of GE properties and degree distribution of species permitted the classification of food webs into six different classes. These classes characterize the differences in robustness of food webs to species loss. While the webs having uniform degree distributions and displaying GE properties are the most robust to species loss, the presence of bottlenecks and skewed distribution of the number of links per species make food webs very vulnerable to primary removal of species.     

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.     

Using trophic hierarchy to understand food web structure

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

Parasites in food webs: the ultimate missing links

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

Connectance indicates the robustness of food webs when subjected to species loss

Many ecologists are concerned that biodiversity loss from human impact on natural ecosystems could compromise ecosystem stability. A relationship between diversity and stability was proposed by MacArthur [MacArthur, R.H., 1955. Fluctuation of animal populations and a measure of community stability. Ecology 36, 533–536.]. Current thinking (for example, McCann, K., 2000. The diversity–stability debate. Nature 405, 228–233.) acknowledges that interaction pattern among species, rather than species richness per se, is one element of this relationship. Dunne et al. [Dunne, J.A., Williams, R.J., Martinez, N.D., 2002a. Network structure and biodiversity loss in food webs: robustness increases with connectance. Ecol. Lett. 5, 558–567.] showed that the robustness of 16 food webs is correlated with their connectance. Connectance is one measure of interaction pattern. Robustness relates to the maintenance of network integrity and so has consequences for stability; the loss of integrity must have ecosystem-wide implications. This paper tests the hypothesis that changes in a food web's connectance indicate changes in its robustness. It concludes that any change in connectance with species loss, but especially large, negative changes, constitutes a decrease in robustness. Estimation of the change in connectance could support interpretation of monitoring data on species composition, acting as an indicator of food web robustness and, indirectly, of ecosystem stability. It could assist managers to understand the implications of biodiversity loss caused by human intervention in ecosystems, and could assist either choice of intervention or amelioration of impacts.     

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.     

Connectance and parasite diet breadth in flea-mammal webs

The number of links in webs of species interactions, which lies at the heart of the biodiversity-stability debate, has given rise to controversy during the last 20 yr. Studies exploring these web properties have mainly focused on symmetric webs where each species can potentially feed on any other species; asymmetric webs such as host-parasite webs, where one set of species feed on another set of species, have been overlooked. However, food webs are incomplete without parasites and the study of parasite-host sub-web properties deserves attention. Here, using a large database involving 33 regional interaction webs between mammals and their flea parasites, we found a negative relationship between species richness and host-parasite connectance. We suggest that some phylogenetic constraints on flea diet may explain our observed patterns because we found that parasite diet breadth, measured as host taxonomic diversity, was invariant along our host richness gradient. We found that the slope of the logarithmic relationship between the number of realized links and species richness is lower than slope values reported for food webs. We suggest that connectance may not respond to increasing species richness as rapidly in host-parasite webs as in predator-prey food webs due to stronger coevolutionary requirements.     

Climate change alters the structure of arctic marine food webs due to poleward shifts of boreal generalists

Climate-driven poleward shifts, leading to changes in species composition and relative abundances, have been recently documented in the Arctic. Among the fastest moving species are boreal generalist fish which are expected to affect arctic marine food web structure and ecosystem functioning substantially. Here, we address structural changes at the food web level induced by poleward shifts via topological network analysis of highly resolved boreal and arctic food webs of the Barents Sea. We detected considerable differences in structural properties and link configuration between the boreal and the arctic food webs, the latter being more modular and less connected. We found that a main characteristic of the boreal fish moving poleward into the arctic region of the Barents Sea is high generalism, a property that increases connectance and reduces modularity in the arctic marine food web. Our results reveal that habitats form natural boundaries for food web modules, and that generalists play an important functional role in coupling pelagic and benthic modules. We posit that these habitat couplers have the potential to promote the transfer of energy and matter between habitats, but also the spread of pertubations, thereby changing arctic marine food web structure considerably with implications for ecosystem dynamics and functioning.