The Equilibrium Conundrum.

The idea that natural systems tend to be at equilibrium dates back to the origin of the field of ecology and continues to underlie most ecological theory. However, empirical evidence for equilibrium dynamics in nature and in experiments is surprisingly elusive. Here, we address this conundrum by first exploring the history of equilibrium in ecological theory and the evidence for equilibrium dynamics in natural systems. We then search the literature to quantify how empiricists deal with equilibrium in their research and address barriers to integrating the concept of equilibrium into empirical work by providing step-by-step instructions for determining whether a population is at equilibrium. Next, we lay out three ways that equilibrium is embedded in theory, and for each, outline when meeting the equilibrium assumption in empirical tests is critical for scientific inference, and when it may be possible to relax this assumption. And finally, we present concrete steps that empiricists and theoreticians can each take in order to meet in the middle when it comes to equilibrium. We hope that this paper will stimulate new discussions from researchers from across the theory-empirical divide about this longstanding issue.