Food web complexity weakens size-based constraints on the pyramids of life

Marine ecosystems are generally expected to have bottom-heavy trophic structure (more plants than animals) due to size-based constraints arising from increased metabolic requirements and inefficient energy transfer. However, size-based (allometric) approaches are often limited to confined trophic-level windows where energy transfer is predicted by size alone and are constrained to a balance between bottom-up and top-down control at steady state. In real food webs, energy flow is more complex and imbalances in top-down and bottom-up processes can also shape trophic structure. We expand the size-based theory to account for complex food webs and show that moderate levels of food web connectance allow for inverted trophic structure more often than predicted, especially in marine ecosystems. Trophic structure inversion occurs due to the incorporation of complex energy pathways and top-down effects on ecosystems. Our results suggest that marine ecosystems should be top-heavy, and observed bottom-heavy trophic structure may be a result of human defaunation of the ocean that has been more extreme than presently recognized.