Comparing trophic structure across ecosystems (Extended)

Trophic structure, the partitioning of biomass among organisms at different positions in a food web, varies both within and among ecosystems. However, the causes of this variation are poorly understood. Elton's "pyramid of numbers", where primary producers dominate and consumer densities decrease as trophic levels become more remote from the base of production, applies well to most terrestrial systems. However, many aquatic ecosystems apparently violate Elton's rule with inverted biomass pyramids, or ratios of heterotroph-to-autotroph biomass (H:A) greater than one. In this proposal, we describe synthetic work aimed at understanding differences in trophic structure and the relative strength of bottom-up and top-down inputs between diverse freshwater, marine and terrestrial ecosystems. We will test candidate hypotheses for this variation based on factors known to distinguish food webs in the two habitats, such as nutrient limitation and turnover rates, productivity (quantity) and nutrient stoichiometry (quality). Meta-analysis of local-scale herbivore manipulation experiments will be integrated with theoretical development of food web models, and with larger-scale temporal and spatial patterns from resource gradients. This work will move us closer to a comprehensive trophic-dynamic theory, unified across taxa and ecosystem types. It will also increase our mechanistic understanding of how human impacts, such as eutrophication or predator extirpation, propagate or attenuate in ecosystems through trophic interactions.