An empiricist's guide to modern coexistence theory for competitive communities

While most ecological theories have historically invoked niche differences as the primary mechanism allowing species coexistence, we now know that species coexistence in competitive communities actually depends on the balance of two opposing forces: niche differences (ND) that determine how species limit their own growth rate versus that of their competitor, and relative fitness differences (RFD) that establish competitive hierarchies among species. Several different empirical methods have been proposed for measuring ND and RFD in order to make predictions about coexistence of species, yet it remains unclear which method(s) are appropriate for a given empirical study and whether or not those methods actually yield the same information. Here we summarize and compare five different empirical methods, with the aim of providing a practical guide for empiricists who want to predict coexistence among species. These include two phenomenological methods that estimate ND and RFD based on observing competitive interactions among species; two mechanistic methods that estimate ND and RFD based solely on information about species’ resource requirements; and a fifth method that does not yield ND and RFD but describes the impacts of those forces within communities. Based on the specific requirements, limitations, and assumptions of each approach, we offer a series of decision steps that can be used to determine which method(s) are best for a given study system. In particular, we show there are important tradeoffs between mechanistic methods, which require detailed understanding of species niches and physiology but are more tractable experimentally, and phenomenological methods which do not require this detailed information but can be impractical for some study designs. Importantly, we show that although each method can be used to estimate ND and RFD, the methods do not always yield the same values. Therefore we caution against future syntheses that compile these estimates from different empirical studies. Finally, we highlight several areas where modern coexistence theory could benefit from additional empirical work.