The mechanistic basis for higher-order interactions and non-additivity in competitive communities

Motivated by both analytical tractability and empirical practicality, community ecologists have long treated the species pair as the fundamental unit of study. This notwithstanding, the challenge of understanding more complex systems has repeatedly generated interest in the role of so-called higher-order interactions (HOIs) imposed by species beyond the focal pair. Here we argue that HOIs - defined as non-additive effects of density on per capita growth - are best interpreted as emergent properties of phenomenological models (e.g. Lotka-Volterra competition) rather than as distinct 'ecological processes' in their own right. Using simulations of consumer-resource models, we explore the mechanisms and system properties that give rise to HOIs in observational data. We demonstrate that HOIs emerge under all but the most restrictive of assumptions, and that incorporating non-additivity into phenomenological models improves the quantitative and qualitative accuracy of model predictions. Notably, we also observe that HOIs derive primarily from mechanisms and system properties that apply equally to single-species or pairwise systems as they do to more diverse communities. Consequently, there exists a strong mandate for further recognition of non-additive effects in both theoretical and empirical research.