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2012, Trends in Ecology & Evolution
Glossary Assembly: the set of processes by which a food web is rebuilt after disturbance or the creation of new habitat. Bioenergetics: the flow and transformation of energy in and between living organisms and between living organisms and their environment. Cascade model: a food-web model which assumes hierarchical feeding along a single niche axis, with each species allocated a probability of feeding on taxa below it in the hierarchy. Community web: a food web intended to include all species and trophic links that occur within a defined ecological community. 'Species' in this sense might involve various degrees of aggregation or division of biological species. Compartmentalisation: the property by which one subset of species within a food web operates, to varying degrees, independently of other parts. Ecological stoichiometry: how the balance of multiple chemical elements within organisms influences their interactions in food webs. Ecosystem function: the physical, chemical, and biological processes or attributes that contribute to the self-maintenance of the ecosystem; including energy flow, nutrient cycling, filtering, buffering of contaminants, and regulation of populations. Interaction strength: a measure of how much a predator alters population size, biomass, or production of its prey. In food webs these are associated with energy flows while noting that predators can affect prey non-trophically. Niche model: a food-web model which assigns each consumer a feeding distribution on the niche axis that can overlap with itself (cannibalism) and permits trophic loops but still generates realistic patterns whereby species tend to feed in a linear hierarchy. Quantitative food web: a food web where the interaction strengths or trophic flows are quantified. Scale-independence and/or dependence: the degree to which attributes of a food web change with food-web size. Stability: measures of the ability to and/or speed with which a food web regains its structure following a disturbance (resilience) or resists change in food web structure (resistance).
Ecology, 2009
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.
Physical Review E, 2006
In the last three decades, researchers have tried to establish universal patterns about the structure of food webs. Recently was proposed that the exponent η characterizing the efficiency of the energy transportation of the food web had a universal value (η = 1.13). Here we establish a lower bound and an upper one for this exponent in a general spanning tree with the number of trophic species and the trophic levels fixed. When the number of species is large the lower and upper bounds are equal to 1, implying that the result η = 1.13 is due to finite size effects. We also evaluate analytically and numerically the exponent η for hierarchical and random networks. In all cases the exponent η depends on the number of trophic species K and when K is large we have that η → 1. Moreover, this result holds for any number M of trophic levels. This means that food webs are very efficient resource transportation systems.
Trends in Ecology & Evolution, 2012
Given the unprecedented rate of species extinctions facing the planet, understanding the causes and consequences of species diversity in ecosystems is of paramount importance. Ecologists have investigated both the influence of environmental variables on species diversity and the influence of species diversity on ecosystem function and stability. These investigations have largely been carried out without taking into account the overarching stabilizing structures of food webs that arise from evolutionary and successional processes and that are maintained through species interactions. Here, we argue that the same large-scale structures that have been purported to convey stability to food webs can also help to understand both the distribution of species diversity in nature and the relationship between species diversity and food web stability. Specifically, the allocation of species diversity to slow energy channels within food webs results in the skewed distribution of interactions strengths that has been shown to confer stability to complex food webs. We end by discussing the processes that might generate and maintain the structured, stable and diverse food webs observed in nature.
The American Naturalist, 1999
2009
The robustness of ecosystems to species losses is a central question in ecology, given the current pace of extinctions and the many species threatened by human impacts, including habitat destruction and climate change. Robustness from the perspective of secondary extinctions has been addressed in the context of food webs to consider the complex network of species interactions that underlie responses to perturbations. In-silico removal experiments have examined the structural properties of food webs that enhance or hamper the robustness of ecosystems to species losses, with a focus on the role of hubs, the most connected species. Here we take a different approach and focus on the role of the connections themselves. We show that trophic links can be divided into functional and redundant based on their contribution to robustness. The analysis of empirical webs shows that hubs are not necessarily the most important species as they may hold many redundant links. Furthermore, the fraction of functional connections is high and constant across systems regardless of size and interconnectedness. The main consequence of this scaling pattern is that ecosystem robustness can be considerably reduced by species extinctions even when these do not result in any secondary extinctions. This introduces the possibility of tipping points in the collapse of ecosystems.
Proceedings of The National Academy of Sciences, 2002
Feeding relationships can cause invasions, extirpations, and population fluctuations of a species to dramatically affect other species within a variety of natural habitats. Empirical evidence suggests that such strong effects rarely propagate through food webs more than three links away from the initial perturbation. However, the size of these spheres of potential influence within complex communities is generally unknown. Here, we show for that species within large communities from a variety of aquatic and terrestrial ecosystems are on average two links apart, with >95% of species typically within three links of each other. Species are drawn even closer as network complexity and, more unexpectedly, species richness increase. Our findings are based on seven of the largest and most complex food webs available as well as a food-web model that extends the generality of the empirical results. These results indicate that the dynamics of species within ecosystems may be more highly interconnected and that biodiversity loss and species invasions may affect more species than previously thought.
Artificial Life and Robotics, 2009
Ecology, 1983
The structure of 40 real food webs , represent ing aquatic and terrestrial communities fro~ all latitudes, is found to be markedly affected by the degree of variability of (he physical environme nt. In particular, food webs in flu ctuating ecosystems are characterized by a significantl y lower conn e cta!l~e than webs representative of more constant systems. This is interpreted within the context of stabil Ity theory as a means 10 optimize feedi ng in the face of increasing distu rbance. It is sh own further that the nature of the habitat itself imposes addition al constraints on food web structu re in interti dal, pelagic, estuarine, and forest ecosystems .
Ecology Letters, 2010
Food web structure plays an important role when determining robustness to cascading secondary extinctions. However, existing food web models do not take into account likely changes in trophic interactions (‘rewiring’) following species loss. We investigated structural dynamics in 12 empirically documented food webs by simulating primary species loss using three realistic removal criteria, and measured robustness in terms of subsequent secondary extinctions. In our model, novel trophic interactions can be established between predators and food items not previously consumed following the loss of competing predator species. By considering the increase in robustness conferred through rewiring, we identify a new category of species – overlap species – which promote robustness as shown by comparing simulations incorporating structural dynamics to those with static topologies. The fraction of overlap species in a food web is highly correlated with this increase in robustness; whereas species richness and connectance are uncorrelated with increased robustness. Our findings underline the importance of compensatory mechanisms that may buffer ecosystems against environmental change, and highlight the likely role of particular species that are expected to facilitate this buffering.
Ecological Modelling, 2007
Ecological network analysis allows for an investigation of the structural and functional interconnectedness in ecosystems. Typically, these interactions are seen to comprise a food web of "who eats whom", but more generally applies to the transfer of energy-matter within the biotic and abiotic ecosphere. This web of transactions can be depicted as a digraph or an adjacency matrix in which the presence of direct transactions are represented as a 1 and no transactions as 0. Each transaction between system components leads to an overall network structural pattern. These structures cluster into different categories or regimes based on their cyclic nature. This paper demonstrates threshold effects of the placement or removal of links, such that certain changes essentially keep the structure in the same regime whereas others shift it to another regime in a non-linear manner.
Journal of Animal Ecology, 2004
1Recent efforts to understand how the patterning of interaction strength affects both structure and dynamics in food webs have highlighted several obstacles to productive synthesis. Issues arise with respect to goals and driving questions, methods and approaches, and placing results in the context of broader ecological theory.2Much confusion stems from lack of clarity about whether the questions posed relate to community-level patterns or to species dynamics, and to what authors actually mean by the term ‘interaction strength’. Here, we describe the various ways in which this term has been applied and discuss the implications of loose terminology and definition for the development of this field.3Of particular concern is the clear gap between theoretical and empirical investigations of interaction strengths and food web dynamics. The ecological community urgently needs to explore new ways to estimate biologically reasonable model coefficients from empirical data, such as foraging rates, body size, metabolic rate, biomass distribution and other species traits.4Combining numerical and analytical modelling approaches should allow exploration of the conditions under which different interaction strengths metrics are interchangeable with regard to relative magnitude, system responses, and species identity.5Finally, the prime focus on predator–prey links in much of the research to date on interaction strengths in food webs has meant that the potential significance of non-trophic interactions, such as competition, facilitation and biotic disturbance, has been largely ignored by the food web community. Such interactions may be important dynamically and should be routinely included in future food web research programmes.Recent efforts to understand how the patterning of interaction strength affects both structure and dynamics in food webs have highlighted several obstacles to productive synthesis. Issues arise with respect to goals and driving questions, methods and approaches, and placing results in the context of broader ecological theory.Much confusion stems from lack of clarity about whether the questions posed relate to community-level patterns or to species dynamics, and to what authors actually mean by the term ‘interaction strength’. Here, we describe the various ways in which this term has been applied and discuss the implications of loose terminology and definition for the development of this field.Of particular concern is the clear gap between theoretical and empirical investigations of interaction strengths and food web dynamics. The ecological community urgently needs to explore new ways to estimate biologically reasonable model coefficients from empirical data, such as foraging rates, body size, metabolic rate, biomass distribution and other species traits.Combining numerical and analytical modelling approaches should allow exploration of the conditions under which different interaction strengths metrics are interchangeable with regard to relative magnitude, system responses, and species identity.Finally, the prime focus on predator–prey links in much of the research to date on interaction strengths in food webs has meant that the potential significance of non-trophic interactions, such as competition, facilitation and biotic disturbance, has been largely ignored by the food web community. Such interactions may be important dynamically and should be routinely included in future food web research programmes.
Journal of Animal Ecology, 2008
1. Following the development of the relatively successful niche model, several other simple structural food web models have been proposed. These models predict the detailed structure of complex food webs given only two input parameters, the numbers of species and the number of feeding links among them. 2. The models claim different degrees of success but have not been compared consistently with each other or with the empirical data. We compared the performance of five structural models rigorously against 10 empirical food webs from a variety of aquatic and terrestrial habitats containing 25-92 species and 68-997 links. 3. All models include near-hierarchical ordering of species' consumption and have identical distributions of the number of prey of each consumer species, but differ in the extent to which species' diets are required to be contiguous and the rules used to assign feeding links. 4. The models perform similarly on a range of food-web properties, including the fraction of top, intermediate and basal species, the standard deviations of generality and connectivity and the fraction of herbivores and omnivores. 5. For other properties, including the standard deviation of vulnerability, the fraction of cannibals and species in loops, mean trophic level, path length, clustering coefficient, maximum similarity and diet discontinuity, there are significant differences in the performance of the different models. 6. While the empirical data do not support the niche model's assumption of diet contiguity, models which relax this assumption all have worse overall performance than the niche model. All the models underestimate severely the fraction of species that are herbivores and exhibit other important failures that need to be addressed in future research.
Oikos, 2009
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).
Ecology, 2009
Food webs depict who eats whom in communities. Ecologists have examined statistical metrics and other properties of food webs, but mainly due to the uneven quality of the data, the results have proved controversial. The qualitative data on which those efforts rested treat trophic interactions as present or absent and disregard potentially huge variation in their magnitude, an approach similar to analyzing traffic without differentiating between highways and side roads. More appropriate data are now available and were used here to analyze the relationship between trophic complexity and diversity in 59 quantitative food webs from seven studies (14-202 species) based on recently developed quantitative descriptors. Our results shed new light on food-web structure. First, webs are much simpler when considered quantitatively, and link density exhibits scale invariance or weak dependence on food-web size. Second, the ''constant connectance'' hypothesis is not supported: connectance decreases with web size in both qualitative and quantitative data. Complexity has occupied a central role in the discussion of food-web stability, and we explore the implications for this debate. Our findings indicate that larger webs are more richly endowed with the weak trophic interactions that recent theories show to be responsible for food-web stability.
Advances in Ecological Research, 2005
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