Omnivory and the stability of food webs

The ecological concept of omnivory, feeding at more than a single trophic level, is formulated as an intermediate stage between any two of three classical three-dimensional species interaction systems-tritrophic chain, competition, and polyphagy. It is shown that omnivory may be either stabilizing or destabilizing, depending, in part, on the conditions of the parent systems from which it derives. It is further conjectured that the tritrophic to competition gradient cannot be entirely stable, that there must be an instability at some level of intermediate omnivory.     

Examining the potential effects of species aggregation on the network structure of food webs

One of the key measures that have been used to describe the topological properties of complex networks is the “degree distribution”, which is a measure that describes the frequency distribution of number of links per node. Food webs are complex ecological networks that describe the trophic relationships among species in a community, and the topological properties of empirical food webs, including degree distributions, have been examined previously. Previously, the “niche model” has been shown to accurately predict degree distributions of empirical food webs, however, the niche model-generated food webs were referenced against empirical food webs that had their species grouped together based on their taxonomic and/or trophic relationships (aggregated food webs). Here, we explore the effects of species aggregation on the ability of the niche model to predict the total- (sum of prey and predator links per node), in- (number of predator links per node), and out- (number of prey links per node) degree distributions of empirical food webs by examining two food webs that can be aggregated at different levels of resolution. The results showed that (1) the cumulative total- and out-degree distributions were consistent with the niche model predictions when the species were aggregated, (2) when the species were disaggregated (i.e., higher resolution), there were mixed conclusions with regards to the niche model's ability to predict total- and out-degree distributions, (3) the model's ability to predict the in-degree distributions of the two food webs was generally inadequate. Although it has been argued that universal functional form based on the niche model could describe the degree distribution patterns of empirical food webs, we believe there are some limitations to the model's ability to accurately predict the structural properties of food webs.     

Benthos as the basis for arctic lake food webs

Plankton have traditionally been viewed as the basis for limnetic food webs, with zooplankton acting as a gateway for energy passing between phytoplanktonic primary producers and fish. Often, benthic production has been considered to be important primarily in shallow systems or as a subsidy to planktonic food web pathways. Stable isotope food web analyses of two arctic lakes (NE14 and I minus) in the Toolik Lake region of Alaska indicate that benthos are the primary source of carbon for adults of all species of benthic and pelagic fish present. We found no effect of turbidity, which may suppress benthic algae by shading, on food web structure. Even though Secchi transparency varied from 10.2 m in NE14 to 0.55–2.6 m in I minus, food webs in both lakes were based upon benthos, had four trophic levels, and culminated with omnivorous lake trout. We suggest that the importance of benthos in the food webs of these lakes is due to their extreme oligotrophy, resulting in planktonic resources that are insufficient for the support of planktivorous consumers.     

Food webs in river networks

Food webs and river drainages are both hierarchical networks and complex adaptive systems. How does living within the second affect the first? Longitudinal gradients in productivity, disturbance regimes and habitat structure down rivers have long interested ecologists, but their effects on food web structure and dynamics are just beginning to be explored. Even less is known about how network structure per se influences river and riparian food webs and their members. We offer some preliminary observations and hypotheses about these interactions, emphasizing observations on upstream–downstream changes in food web structure and controls, and introducing some ideas and predictions about the unexplored question of food web responses to some of the network properties of river drainages.     

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.     

Emergence of non-random structure in local food webs generated from randomly structured regional webs

Previous studies have shown that high-resolution, empirical food webs possess a non-random network structure, typically characterized by uniform or exponential degree distributions. However, the empirical food webs that have been investigated for their structural properties represent local communities that are only a subset of a larger pool of regionally coexisting species. Here, we use a simple model to investigate the effects of regional food web structure on local food webs that are assembled by two simple processes: random immigration of species from a source web (regional food web), and random extinction of species within the local web. The model shows that local webs with non-random degree distributions can arise from randomly structured source webs. A comparison of local webs assembled from randomly structured source webs with local webs assembled from source webs generated by the niche model shows that the former have higher species richness at equilibrium, but have a nonlinear response to changing extinction rates. These results imply that the network structure of regional food webs can play a significant role in the assembly and dynamics of local webs in natural ecosystems. With natural landscapes becoming increasingly fragmented, understanding such structure may be a necessary key to understanding the maintenance and stability of local species diversity.     

Food webs: Ordering species according to body size yields high degree of intervality

Food webs, the networks describing “who eats whom” in an ecosystem, are nearly interval, i.e. there is a way to order the species so that almost all the resources of each consumer are adjacent in the ordering. This feature has important consequences, as it means that the structure of food webs can be described using a single (or few) species' traits. Moreover, exploiting the quasi-intervality found in empirical webs can help build better models for food web structure. Here we investigate which species trait is a good proxy for ordering the species to produce quasi-interval orderings. We find that body size produces a significant degree of intervality in almost all food webs analyzed, although it does not match the maximum intervality for the networks. There is also a great variability between webs. Other orderings based on trophic levels produce a lower level of intervality. Finally, we extend the concept of intervality from predator-centered (in which resources are in intervals) to prey-centered (in which consumers are in intervals). In this case as well we find that body size yields a significant, but not maximal, level of intervality. These results show that body size is an important, although not perfect, trait that shapes species interactions in food webs. This has important implications for the formulation of simple models used to construct realistic representations of food webs.     

Biodiversity maintenance in food webs with regulatory environmental feedbacks

Although the food web is one of the most fundamental and oldest concepts in ecology, elucidating the strategies and structures by which natural communities of species persist remains a challenge to empirical and theoretical ecologists. We show that simple regulatory feedbacks between autotrophs and their environment when embedded within complex and realistic food-web models enhance biodiversity. The food webs are generated through the niche-model algorithm and coupled with predator-prey dynamics, with and without environmental feedbacks at the autotroph level. With high probability and especially at lower, more realistic connectance levels, regulatory environmental feedbacks result in fewer species extinctions, that is, in increased species persistence. These same feedback couplings, however, also sensitize food webs to environmental stresses leading to abrupt collapses in biodiversity with increased forcing. Feedback interactions between species and their material environments anchor food-web persistence, adding another dimension to biodiversity conservation. We suggest that the regulatory features of two natural systems, deep-sea tubeworms with their microbial consortia and a soil ecosystem manifesting adaptive homeostatic changes, can be embedded within niche-model food-web dynamics.     

Adaptive omnivory and species coexistence in tri-trophic food webs

The commonness of omnivory in natural communities is puzzling, because simple dynamic models of tri-trophic systems with omnivory are prone to species extinction. In particular, the intermediate consumer is frequently excluded by the omnivore at high levels of enrichment. It has been suggested that adaptive foraging by the omnivore may facilitate coexistence, because the intermediate consumer should persist more easily if it is occasionally dropped from the omnivore's diet. We explore theoretically how species permanence in tri-trophic systems is affected if the omnivore forages adaptively according to the "diet rule", i.e., feeds on the less profitable of its two prey species only if the more profitable one is sufficiently rare. We show that, compared to systems where omnivory is fixed, adaptive omnivory may indeed facilitate 3-species persistence. Counter to intuition, however, facilitation of 3-species coexistence requires that the intermediate consumer is a more profitable prey than the basal resource. Consequently, adaptive omnivory does not facilitate persistence of the intermediate consumer but enlarges the persistence region of the omnivore towards parameter space where a fixed omnivore would be excluded by the intermediate consumer. Overall, the positive effect of adaptive omnivory on 3-species persistence is, however, small. Generally, whether omnivory is fixed or adaptive, 3-species permanence is most likely when profitability (=conversion efficiency into omnivores) is low for basal resources and high for intermediate consumers.     

Food partitioning by lake-dwelling triclads and glossiphoniid leeches: field and laboratory experiments

The triclads Polycelis tenuis and Dugesia polychroa and the glossiphoniid leeches Glossiphonia complanata and Helobdella stagnalis are abundant on the stony shores of productive British lakes. All species are food limited and there is considerable overlap in the diets of these triclads and leeches. This paper investigates interactions between the two groups using field and laboratory experiments to try to identify the mechanism of their co-existence. Triclad and leech numbers were manipulated inside experimental enclosures, mathced by controls, erected on the stony shore of an eutrophic English lake. Increasing the numbers of P. tenuis and D. polychroa prior to the reproductive season in spring resulted in a significant decrease in the numbers and body size of G. complanata and H. stagnalis compared with control populations in the summer months, and vice versa. However, increases and decreases were temporary with a readjustment of numbers and body size to control levels in the autumn after reproduction had ceased. It is suggested that increasing the numbers of either group elevated the severity of both intra- and interspecific competition for food. The condition of prey may, in part, determine the strength of competition, and this was examined in laboratory experiments in which different densities and ratios of P. tenuis and H. stagnalis were offered either live of recently crushed Asellus aquaticus. In monospecific controls, growth rates of P. tenuis were greater when fed on crushed than live Asellus, but there was no significant difference in the growth of H. stagnalis fed either live or crushed prey. In mixed cultures of predators, P. tenuis and H. stagnalis were the superior competitors when fed on crushed and live Asellus, respectively. However, when competitive pressure was low, at low densities of predators, the presence of H. stagnalis in mixed cultures fed on live prey was beneficial to the growth of P. tenuis. These results are explained in terms of the greater ability of triclads to detect damaged prey, leaking body fluids, due to their sophisticated chemosensory system, and the ability of leeches to capture live prey due to the presence of suckers. It is concluded that co-existence of the two groups in British lakes is assisted by the partitioning of food on a live or damaged basis.     

Effects of dynamics on ecological networks

Ecological food webs define the feeding patterns of interacting species. The architecture of such networks may be affected by dynamical processes operating within them, ultimately influencing the capacity of the networks to persist. As yet relatively little is known about these effects. We compared the architecture of ecological networks with a fixed number of species, constructed in four contrasting ways: (I) topological networks, which required only that species had prey to eat; (II) persistent networks, in which species had also to persist under a simple model of population dynamics; (III) assembled networks, built up by sequential addition of species with dynamical persistence at each step in the sequence; (IV) evolved networks where, in addition to dynamical persistence, body size of species was determined by a simple mutation-selection process. Dynamics had fundamental effects on architecture, the networks of classes II, III and IV being restricted to a small number of trophic levels, in contrast to the non-dynamic, topological class I networks. Class III assembled networks tended to have fewer trophic levels and a more pyramidal biomass distribution than networks of classes II and IV. In evolved class IV networks, the smallest consumers converged to similar body sizes, whereas larger consumers evolved more slowly and did not show such convergence. The results indicate that dynamics affect the architecture of food webs, and that assumptions about simultaneous arrival, sequential arrival and evolution lead to different outcomes. Sequential assembly was shown to have a special property of finding rare sets of persistent species in a small number of steps, suggesting that the rarity of stable communities is not a serious problem in the development of complex communities.     

Evidence for the existence of a robust pattern of prey selection in food webs

Food webs aim to provide a thorough representation of the trophic interactions found in an ecosystem. The complexity of empirical food webs, however, is leading many ecologists to focus dynamic ecosystem studies on smaller microcosm or mesocosm studies based upon community modules, which comprise three to five species and the interactions likely to have ecological relevance. We provide here a structural counterpart to community modules. We investigate food-web 'motifs' which are n-species connected subgraphs found within the food web. Remarkably, we find that the over- and under-representation of three-species motifs in empirical food webs can be understood through comparison to a static food-web model, the niche model. Our result conclusively demonstrates that predation upon species with some 'characteristic' niche value is the prey selection mechanism consistent with the structural properties of empirical food webs.     

Serological tracers of meiofaunal food webs

Serological methods utilizing taxon-specific antibodies were used to identify trophic connections in a salt marsh of South Carolina (U. S. A.). The incorporation of meiofauna within the benthic invertebrate food web was detected with these methods when microscopial examinations of predator stomachs revealed nothing but amorphous material and detritus. Measurements of soluble prey proteins in both predator guts and surficial sediments provided data to quantify the trophic connections. Difficulties with data interpretation limit the utility of serological methods for quantifying predation in the field.     

Emergence of complexity in evolving niche-model food webs

We have analysed mechanisms that promote the emergence of complex structures in evolving model food webs. The niche model is used to determine predator-prey relationships. Complexity is measured by species richness as well as trophic level structure and link density. Adaptive dynamics that allow predators to concentrate on the prey species they are best adapted to lead to a strong increase in species number but have only a small effect on the number and relative occupancy of trophic levels. The density of active links also remains small but a high number of potential links allows the network to adjust to changes in the species composition (emergence and extinction of species). Incorporating effects of body size on individual metabolism leads to a more complex trophic level structure: both the maximum and the average trophic level increase. So does the density of active links. Taking body size effects into consideration does not have a measurable influence on species richness. If species are allowed to adjust their foraging behaviour, the complexity of the evolving networks can also be influenced by the size of the external resources. The larger the resources, the larger and more complex is the food web it can sustain. Body size effects and increasing resources do not change size and the simple structure of the evolving networks if adaptive foraging is prohibited. This leads to the conclusion that in the framework of the niche model adaptive foraging is a necessary but not sufficient condition for the emergence of complex networks. It is found that despite the stabilising effect of foraging adaptation the system displays elements of self-organised critical behaviour.     

EcoLens: Integration and interactive visualization of ecological datasets

Complex multi-dimensional datasets are now pervasive in science and elsewhere in society. Better interactive tools are needed for visual data exploration so that patterns in such data may be easily discovered, data can be proofread, and subsets of data can be chosen for algorithmic analysis. In particular, synthetic research such as ecological interaction research demands effective ways to examine multiple datasets. This paper describes our integration of hundreds of food-web datasets into a common platform, and the visualization software, EcoLens, we developed for exploring this information. This publicly-available application and integrated dataset have been useful for our research predicting large complex food webs, and EcoLens is favorably reviewed by other researchers. Many habitats are not well represented in our large database. We confirm earlier results about the small size and lack of taxonomic resolution in early food webs but find that they and a non-food-web source provide trophic information about a large number of taxa absent from more modern studies. Corroboration of Tuesday Lake trophic links across studies is usually possible, but lack of links among congeners may have several explanations. While EcoLens does not provide all kinds of analytical support, its label- and item-based approach is effective at addressing concerns about the comparability and taxonomic resolution of food-web data.     

The myth of constant predator: prey ratios

Apparent constancy in the ratio of predator species to prey species has been offered as evidence that ecological communities are structured by interspecific interactions. If significantly different from random expectation, this effect would be one of the few sound pieces of evidence for community structure. The evidence was re-evaluated by using the data from previous studies to form species pools, and forming simulated communities by drawing species at random from these pools (with replacement). Using a correlation coefficient (number of predator species versus number of prey species), and also the statistic used by the original workers (where different), the observed predator:prey correlation was compared to that for the random communities. In five studies, the observed predator:prey ratio was not significantly different from random expectation. In the only two studies where there was significant departure from the null model, it was with more variation in the ratio than expected on a random basis. It is concluded that there is as yet no evidence for near-constant predator:prey ratios.     

Highly resolved early Eocene food webs show development of modern trophic structure after the end-Cretaceous extinction

Generalities of food web structure have been identified for extant ecosystems. However, the trophic organization of ancient ecosystems is unresolved, as prior studies of fossil webs have been limited by low-resolution, high-uncertainty data. We compiled highly resolved, well-documented feeding interaction data for 700 taxa from the 48 million-year-old latest early Eocene Messel Shale, which contains a species assemblage that developed after an interval of protracted environmental and biotal change during and following the end-Cretaceous extinction. We compared the network structure of Messel lake and forest food webs to extant webs using analyses that account for scale dependence of structure with diversity and complexity. The Messel lake web, with 94 taxa, displays unambiguous similarities in structure to extant webs. While the Messel forest web, with 630 taxa, displays differences compared to extant webs, they appear to result from high diversity and resolution of insect–plant interactions, rather than substantive differences in structure. The evidence presented here suggests that modern trophic organization developed along with the modern Messel biota during an 18 Myr interval of dramatic post-extinction change. Our study also has methodological implications, as the Messel forest web analysis highlights limitations of current food web data and models.