From the Neutral Theory to a Comprehensive and Multiscale Theory of Ecological Equivalence

The neutral theory of biodiversity assumes that coexisting organisms are equally able to survive, reproduce, and disperse (ecological equivalence), but predicts that stochastic fluctuations of these abilities drive diversity dynamics. It predicts remarkably well many biodiversity patterns, although substantial evidence for the role of niche variation across organisms seems contradictory. Here, we discuss this apparent paradox by exploring the meaning and implications of ecological equivalence. We address the question whether neutral theory provides an explanation for biodiversity patterns and acknowledges causal processes. We underline that ecological equivalence, although central to neutral theory, can emerge at local and regional scales from niche-based processes through equalizing and stabilizing mechanisms. Such emerging equivalence corresponds to a weak conception of neutral theory, as opposed to the assumption of strict equivalence at the individual level in strong conception. We show that this duality is related to diverging views on hypothesis testing and modeling in ecology. In addition, the stochastic dynamics exposed in neutral theory are pervasive in ecological systems and, rather than a null hypothesis, ecological equivalence is best understood as a parsimonious baseline to address biodiversity dynamics at multiple scales.