Heterogeneity in plant-soil feedbacks and resident population dynamics affect mutual invasibility

1. Understanding the mechanisms governing coexistence is a central goal in ecology and has implications for conserving and restoring communities, yet the high diversity in many plant communities is difficult to explain. Theory suggests that plant-soil feedbacks (PSF) can lead to frequency-dependent coexistence by suppressing conspecifics more than heterospecifics, potentially helping to explain high-diversity plant communities. In addition, species-specific population dynamics, including the rate at which individuals are replaced in a population or population turnover rate, may influence coexistence outcomes. 2. We have created a rigorous test of the coexistence predictions of theory by generating a soil heterogeneity experiment in the field and testing for mutual invasibility by establishing resident populations, then experimentally invading them. Experimental tests of mutual invasibility can demonstrate coexistence because, if species are able to invade one another's populations when at low density, they should exhibit long-term coexistence. We use pairs of congeners in this experiment that coexist at small spatial scales, sometimes within cm, at our field site. 3. We demonstrate that invader individuals established better in congener's soils than in conspecific soils, consistent with plant-soil feedback mediated coexistence. This effect was often mediated by competition with established resident plants. 4. Further, we show that soil heterogeneity interacted with the population turnover rate of the resident population to influence invasibility (P < 0.10), consistent with the theoretical prediction that a plant's population dynamics will interact with heterogeneity to influence coexistence. 5. Synthesis. Plant-soil feedbacks (PSF) can in theory lead to frequency-dependent coexistence, and reciprocally, negative feedback effects in glasshouse experiments are often consistent with this prediction. We provide the first field test of mutual invasibility structured by PSF, demonstrating that PSF can lead to coexistence when they create a patchy, or heterogeneous, soil environment. This work suggests that understanding the influence of PSF on diversity necessitates understanding the spatial scale at which soil heterogeneity emerges in the field. Thus, high diversity might be maintained in plant communities by heterogeneity created by plants' influence on the soil, and this outcome depends strongly on population dynamics.