A unifying theory for top-heavy ecosystem structure in the ocean

Size generally dictates metabolic requirements, trophic level, and consequently, ecosystem structure, where inefficient energy transfer leads to bottom-heavy ecosystem structure and biomass decreases as individual size (or trophic level) increases. However, many animals deviate from simple size-based predictions by either adopting generalist predatory behavior, or feeding lower in the trophic web than predicted from their size. Here we show that generalist predatory behavior and lower trophic feeding at large body size increase overall biomass and shift ecosystems from a bottom-heavy pyramid to a top-heavy hourglass shape, with the most biomass accounted for by the largest animals. These effects could be especially dramatic in the ocean, where primary producers are the smallest components of the ecosystem. This approach makes it possible to explore and predict, in the past and in the future, the structure of ocean ecosystems without biomass extraction and other impacts.