Effects of intraspecific variation in a native species' phenology on its coexistence with non-native plants

Intraspecific trait variation is ubiquitous and is likely to influence species coexistence. Despite theoretical progress, empirical work on the effects of intraspecific variation on the dynamics of competing species is rare. This is because of the formidable empirical requirements necessary to link intraspecific variation in species' functional traits with intraspecific variation in the demographic and competitive rates that mediate coexistence. Here we partially overcome these challenges to determine how intraspecific variation in reproductive phenology in a native Californian annual plant species Lasthenia californica affects its ability to coexist with two non-native species Bromus madritensis and Lactuca serriola that display contrasting phenological patterns. Using data from a field experiment, we empirically parameterize a model of competitive population dynamics, accounting for the effects of intraspecific phenological trait variation on the native species' response to both intra- and interspecific competition. We find that intraspecific variation in phenology drives differences in the native species' response to competition. Moreover, simulations of the parameterized model show that this variation improves the competitive performance of the native species. This occurs because of the effects of nonlinear averaging mediated by a nonlinear, concave-up competition function that is a general feature of competition across a wide range of taxa. While intraspecific variation improves competitive performance, we also find that the magnitude of the benefit is predicted to be insufficient to prevent competitive exclusion against the non-native species with early phenogy Bromus. Against the second non-native species with later phenology Lactuca, intraspecific variation is predicted to result in coexistence where competitive exclusion would otherwise occur, but we could not rule out alternative qualitative outcomes for this interaction.