In ecosystems, species interact with other species directly and through abiotic factors in multiple ways, often forming complex networks of various types of ecological interaction. Out of this suite of interactions, predator–prey interactions have received most attention.

The aim of this paper is to provide new and testable ideas on how to understand and model ecosystems in which many different types of ecological interaction operate simultaneously.

we propose that food webs are structured among two main axes of organization: a vertical (classic) axis representing trophic position and a new horizontal ‘ecological stoichiometry’ axis representing decreasing palatability of plant parts and detritus for herbivores and detrivores and slower turnover times.