A mechanistic theory for aquatic food chain length

Multiple hypotheses propose an ostensibly disparate array of drivers of food chain length (FCL), with contradictory support from natural settings. Here we posit that the magnitude of vertical energy flux in food webs underlies several drivers of FCL. We show that rising energy flux fuels top-heavy biomass pyramids, promoting omnivory, thereby reducing FCL. We link this theory to commonly evaluated hypotheses for environmental drivers of FCL (productivity, ecosystem size) and demonstrate that effects of these drivers should be context-dependent. We evaluate support for this theory in lake and marine ecosystems and demonstrate that ecosystem size is the most important driver of FCL in low-productivity ecosystems (positive relationship) while productivity is most important in large and high-productivity ecosystems (negative relationship). This work stands in contrast to classical hypotheses, which predict a positive effect of productivity on FCL, and may help reconcile the contradictory nature of published results for drivers of FCL.