Understanding what structures ecological communities is vital to answering questions about extinctions, environmental change, trophic cascades, and ecosystem functioning. Optimal foraging theory was conceived to increase such understanding by providing a framework with which to predict species interactions and resulting community structure. Here, we use an optimal foraging model and allometries of foraging variables to predict the structure of real food webs. The qualitative structure of the resulting model provides a more mechanistic basis for the phenomenological rules of previous models. Quantitative analyses show that the model predicts up to 65% of the links in real food webs. The deterministic nature of the model allows analysis of the model's successes and failures in predicting particular interactions. Predacious and herbivorous feeding interactions are better predicted than pathogenic, parasitoid, and parasitic interactions. Results also indicate that accurate prediction and modeling of some food webs will require incorporating traits other than body size and diet choice models specific to different types of feeding interaction. The model results support the hypothesis that individual behavior, subject to natural selection, determines individual diets and that food web structure is the sum of these individual decisions.body size ͉ complexity ͉ connectance E xplaining and predicting community structure is a central part of ecological research. It is vital to answering questions about extinctions (1, 2), environmental change (3), trophic cascades (4), and ecosystem functioning (5, 6). We focus on one of the major components of community structure: the interactions between consumers and resources. Food webs represent communities in terms of species and the feeding links between them, and discovering what determines their structure is a major goal in ecology.There are several different approaches to modeling food webs, each emphasizing different processes by which food web structure might be controlled. For example, dynamic models focus on how structure relates to population dynamics and community stability (2, 7-11). Evolutionary models incorporate the processes that control the formation and expansion of food webs (12, 13). Static models include rules that determine structural attributes of food webs (14-19). These models have developed our thinking about food webs in a number of ways, but they have limitations. The stochastic, and therefore generalized, nature of these models means that predicting the arrangement of links in a particular real food web is difficult. Here, we describe a new approach to modeling food webs that avoids some of these problems through use of the allometries of body size and foraging behavior of individual consumers.The contingency model of optimal foraging predicts the diet that maximises a consumer's rate of energy intake (20). We have shown that this optimal foraging model can predict consumer diet breadths and food web connectance (21). This model of connectance [which we te...

We present a quantitative synthesis of trophic cascades in terrestrial systems using data from 41 studies, reporting 60 independent tests. The studies covered a wide range of taxa in various terrestrial systems with varying degrees of species diversity. We quantified the average magnitude of direct effects of carnivores on herbivore prey and indirect effects of carnivores on plants. We examined how the effect magnitudes varied with type of carnivores in the study system, food web diversity, and experimental protocol. A metaanalysis of the data revealed that trophic cascades were common among the studies. Exceptions to this general trend did arise. In some cases, trophic cascades were expected not to occur, and they did not. In other cases, the direct effects of carnivores on herbivores were stronger than the indirect effects of carnivores on plants, indicating that top-down effects attenuated. Top-down effects usually attenuated whenever plants contained antiherbivore defenses or when herbivore species diversity was high. Conclusions about the strength of top-down effects of carnivores varied with the type of carnivore and with the plant-response variable measured. Vertebrate carnivores generally had stronger effects than invertebrate carnivores. Carnivores, in general, had stronger effects when the response was measured as plant damage rather than as plant biomass or plant reproductive output. We caution, therefore, that conclusions about the strength of top-down effects could be an artifact of the plant-response variable measured. We also found that mesocosm experiments generally had weaker effect magnitudes than open-plot field experiments or observational experiments. Trophic cascades in terrestrial systems, although not a universal phenomenon, are a consistent response throughout the published studies reviewed here. Our analysis thus suggests that they occur more frequently in terrestrial systems than currently believed. Moreover, the mechanisms and strengths of top-down effects of carnivores are equivalent to those found in other types of systems (e.g., aquatic environments).

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Trophic cascades are regarded as important signals for top‐down control of food web dynamics. Although there is clear evidence supporting the existence of trophic cascades, the mechanisms driving this important dynamic are less clear. Trophic cascades could arise through direct population‐level effects, in which predators prey on herbivores, thereby decreasing the abundance of herbivores that impact plant trophic levels. Trophic cascades could also arise through indirect behavioral‐level effects, in which herbivore prey shift their foraging behavior in response to predation risk. Such behavioral shifts can result in reduced feeding time and increased starvation risk, again lowering the impact of herbivores on plants. We evaluated the relative importance of these two mechanisms, using field experiments in an old‐field system composed of herbaceous plants, grasshopper herbivores, and spider predators. We created two treatments, Risk spiders that had their chelicerae glued, and Predation spiders that remained unmanipulated. We then systematically evaluated the impacts of these predator manipulations at behavioral, population, and food web scales in experimental mesocosms. At the behavioral level, grasshoppers did not distinguish between Risk spiders and Predation spiders. Grasshoppers exhibited significant shifts in feeding‐time budget in the presence of spiders vs. when alone. At the grasshopper population level, Risk spider and Predation spider treatments caused the same level of grasshopper mortality, which was significantly higher than mortality in a control without spiders, indicating that the predation effects were compensatory to risk effects. At the food web level, Risk spider and Predation spider treatments decreased the impact grasshoppers had on grass biomass, supporting the existence of a trophic cascade. Moreover, Risk spider and Predation spider treatments produced statistically similar effects, again indicating that predation effects on trophic dynamics were compensatory to risk effects. We conclude that indirect effects resulting from antipredator behavior can produce trophic‐level effects that are similar in form and strength to those generated by direct predation events.

Great uncertainty exists in the global exchange of carbon between the atmosphere and the terrestrial biosphere. An important source of this uncertainty lies in the dependency of photosynthesis on the maximum rate of carboxylation (Vcmax) and the maximum rate of electron transport (Jmax). Understanding and making accurate prediction of C fluxes thus requires accurate characterization of these rates and their relationship with plant nutrient status over large geographic scales. Plant nutrient status is indicated by the traits: leaf nitrogen (N), leaf phosphorus (P), and specific leaf area (SLA). Correlations between Vcmax and Jmax and leaf nitrogen (N) are typically derived from local to global scales, while correlations with leaf phosphorus (P) and specific leaf area (SLA) have typically been derived at a local scale. Thus, there is no global-scale relationship between Vcmax and Jmax and P or SLA limiting the ability of global-scale carbon flux models do not account for P or SLA. We gathered published data from 24 studies to reveal global relationships of Vcmax and Jmax with leaf N, P, and SLA. Vcmax was strongly related to leaf N, and increasing leaf P substantially increased the sensitivity of Vcmax to leaf N. Jmax was strongly related to Vcmax, and neither leaf N, P, or SLA had a substantial impact on the relationship. Although more data are needed to expand the applicability of the relationship, we show leaf P is a globally important determinant of photosynthetic rates. In a model of photosynthesis, we showed that at high leaf N (3 gm−2), increasing leaf P from 0.05 to 0.22 gm−2 nearly doubled assimilation rates. Finally, we show that plants may employ a conservative strategy of Jmax to Vcmax coordination that restricts photoinhibition when carboxylation is limiting at the expense of maximizing photosynthetic rates when light is limiting.