Abstract
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.
Highlights
Size generally dictates metabolic requirements, trophic level, and ecosystem structure, where inefficient energy transfer leads to bottom-heavy ecosystem structure and biomass decreases as individual size increases
Size-based predation leads to welldefined ecosystem structure in a linear size spectrum, where biomass decreases with size due to inefficient energy transfer[4,7,12,13,14]
predator-prey mass ratio (PPMR) is defined as the mean ratio of mass at trophic level n divided by the mass at trophic level n−1
Summary
Size generally dictates metabolic requirements, trophic level, and ecosystem structure, where inefficient energy transfer leads to bottom-heavy ecosystem structure and biomass decreases as individual size (or trophic level) increases. 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. Using a simple model, we show how the inclusion of large generalist predators and gigantic secondary consumers could substantially increase total biomass and reduces mean trophic level In marine ecosystems, these animals can invert the trophic structure to a hyperboloid (hourglass) rather than a traditional pyramid shape as biomass is more concentrated in large animals[20,21,22]. The effects of predators that feed well below prey sizes predicted through size-spectra theory or that feed on a wide range of prey sizes provide an alternative perspective to previous explanations of high top predator biomass in pristine marine ecosystems
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