Rapid prey evolution and the dynamics of two-predator food webs

Traits affecting ecological interactions can evolve on the same time scale as population and community dynamics, creating the potential for feedbacks between evolutionary and ecological dynamics. Theory and experiments have shown in particular that rapid evolution of traits conferring defense against predation can radically change the qualitative dynamics of a predator–prey food chain. Here, we ask whether such dramatic effects are likely to be seen in more complex food webs having two predators rather than one, or whether the greater complexity of the ecological interactions will mask any potential impacts of rapid evolution. If one prey genotype can be well-defended against both predators, the dynamics are like those of a predator–prey food chain. But if defense traits are predator-specific and incompatible, so that each genotype is vulnerable to attack by at least one predator, then rapid evolution produces distinctive behaviors at the population level: population typically oscillate in ways very different from either the food chain or a two-predator food web without rapid prey evolution. When many prey genotypes coexist, chaotic dynamics become likely. The effects of rapid evolution can still be detected by analyzing relationships between prey abundance and predator population growth rates using methods from functional data analysis.     

Microbial food chains and food webs

Mathematical models for simple microbial food chains and food webs in continuous culture are developed and analyzed. A model for competition of two microbial species for a single scarce resource is also presented as a degenerate case of the food web model. Two models for food chains are developed. The first is based on a model of microbial growth (Monod's) that is widely mentioned and used at the present time. The second is based on a generalization of that model that recent experimental results on microbial food chains seem to require. Experimental data for microbial food webs are almost entirely lacking but a tentative model having what are felt to be the right properties is developed and analyzed. The results obtained from these models seem to be consistent in most circumstances with current ecological thinking on community dynamics.     

Keystone species and food webs

Different species are of different importance in maintaining ecosystem functions in natural communities. Quantitative approaches are needed to identify unusually important or influential, ‘keystone’ species particularly for conservation purposes. Since the importance of some species may largely be the consequence of their rich interaction structure, one possible quantitative approach to identify the most influential species is to study their position in the network of interspecific interactions. In this paper, I discuss the role of network analysis (and centrality indices in particular) in this process and present a new and simple approach to characterizing the interaction structures of each species in a complex network. Understanding the linkage between structure and dynamics is a condition to test the results of topological studies, I briefly overview our current knowledge on this issue. The study of key nodes in networks has become an increasingly general interest in several disciplines: I will discuss some parallels. Finally, I will argue that conservation biology needs to devote more attention to identify and conserve keystone species and relatively less attention to rarity.     

Omnivory and stability of food webs

The role and prevalence of omnivory, defined as feeding on more than one trophic level, are critical to understand food web structure and dynamics. Whether omnivory stabilizes or destabilizes food webs depends on the assumptions of theoretical models. Recently, Tanabe and Namba [Tanabe, K., Namba, T., 2005. Omivory creates chaos in simple food web models. Ecology 86, 3411–3414] found that omnivory can create chaos in a simple food web model with linear functional responses and 12 model parameters. In this paper, first we numerically examined bifurcation diagrams with all the parameters as bifurcation parameters, including self-limitation of the intermediate consumer and predator. Chaos spontaneously appears when the intraguild predator’s consumption rates are low for nutrient-rich intraguild prey and high for nutrient-poor basal resource and the intraguild prey reproduces efficiently feeding on the basal resource. Second, we investigated effects of the addition of a species into the basic model food web which exhibits chaos. The additional species is assumed to consume only one of the basal resource, intermediate consumer, or omnivorous predator. Consequences of the addition greatly depend on the trophic level on which the additional species feeds. While the increased diversity of predators feeding on the intermediate consumer stabilizes the web, the increased diversity of prey feeding on the basal resource induces collapse of the food web through exploitative competition for the basal resource. The food chain with the top predator feeding on the omnivorous predator is highly unstable unless the mortality of the top predator is extremely low. We discuss the possibility of real-world chaos and the reason why stability of food webs strongly depends on the topological structure of the webs. Finally, we consider the implications of our results for food web theory and resource management.     

Energetics of microbial food webs

The energetic demand of microorganisms in natural waters and the flux of energy between microorganisms and metazoans has been evaluated by empirical measurements in nature, in microcosms and mesocosms, and by simulation models. Microorganisms in temperate and tropical waters often use half or more of the energy fixed by photosynthesis. Most simulations and some experimental results suggest significant energy transfer to metazoans, but empirical evidence is mixed. Considerations of the range of growth yields of microorganisms and the number of trophic transfers among them indicate major energy losses within microbial food webs. Our ability to verify and quantify these processes is limited by the variability of assimilation efficiency and uncertainty about the structure of microbial food webs. However, even a two-step microbial chain is a major energy sink. As an energetic link to metazoans, the detritus food web is inefficient, and its significance may have been overstated. There is not enough bacterial biomass associated with detritus to support metazoan detritivores. Much detritus is digestible by metazoans directly. Thus, metazoans and bacteria may to a considerable degree compete for a common resource. Microorganisms, together with metazoans, are important to the stability of planktonic communities through their roles as rapid mineralizers of organic matter, releasing inorganic nutrients. The competition for organic matter and the resultant rapid mineralization help maintain stable populations of phytoplankton in the absence of advective nutrient supply. At temperatures near O °C, bacterial metabolism is suppressed more than is the rate of photosynthesis. As a result, the products of the spring phytoplankton bloom in high-temperate latitudes are not utilized rapidly by bacteria. At temperatures below 0°C microbial food webs are neither energy sinks or links: they are suppressed. Because the underlying mechanism of low-temperature inhibition is not known, we cannot yet generalize about this as a control of food web processes. Microorganisms may operate on several trophic levels simultaneously. Therefore, the realism of the trophic level concept and the reality of the use of ecological efficiency calculations in ecosystem models is questionable.     

The structure of food webs

For nonrandom models of species interaction there is a precipitous decrease in stability as connectance increases. However, the range of stability for different models of the same connectance is large; stability also depends on how the species interactions are organized. Systems with species feeding on more than one trophic level (omnivores) are likely to be unstable, the extent depending on the number and position of the omnivores. For systems of equal connectance, those that are completely compartmentalized are less likely to be stable than those that are not.     

Integrating ecosystem engineering and food webs

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

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.     

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.     

Making connections in food webs

Patterns in food web structure have provided an important, though contentious, testing ground for ideas about the population dynamics and energetics of multispecies systems. One of the most debated of these patterns is the apparent decrease in food web connectance as the number of species in a web Increases. Several contrasting mechanisms that might determine food web connectance have been suggested. These mechanisms, in combination with new, food web data, suggest that the conventional pattern, and explanations for it, may well be open to dispute. The true nature of the relationship between connectance and species number has implications for the explanation of other web patterns and for theories of food web structure, but a general explanation remains elusive.     

Aggregation and disaggregation of microbial food webs

Models of the microbial food web have generally used compartments aggregated by general body size and gross taxonomy. It has been assumed that these also reflect guilds or holons. Generally, results of simulation or analysis based on this structure have been reasonably well validated. Herein I summarize why the aggregations may be justified and what may be learned from disaggregation.     

Multispecies food webs with diffusion

Using Liapunov's direct method, in this paper, it has been shown that the general Lotka-Volterra food web is stable without and with diffusion under each case of homogeneous reservoir and flux boundary conditions. However, for a three species food web with Holling's functional response the above general result regarding stability is not necessarily true. In such a case, conditions and regions for non-linear stability, without and with diffusion, have been derived. It is shown that such an otherwise unstable system may become stable with diffusion at least in a subregion of the positive octant of the state space.     

Parasites entangled in food webs

Food webs are a fundamental concept in ecology in which parasites have been virtually ignored. In a recent article, Lafferty et al. address this imbalance, finding that the inclusion of parasites in food webs could be of greater importance to ecosystem stability than was previously thought. Furthermore, the bottom of the food chain is perhaps no longer the most dangerous place to be.     

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.