It's about (taking up) space: Discreteness of individuals and the strength of spatial coexistence mechanisms

One strand of modern coexistence theory (MCT) partitions invader growth rates (IGR) to quantify how different mechanisms contribute to species coexistence, highlighting fluctuation-dependent mechanisms. A general conclusion from the classical analytic MCT theory is that coexistence mechanisms relying on temporal variation (such as the temporal storage effect) are generally less effective at promoting coexistence than mechanisms relying on spatial or spatiotemporal variation (primarily growth-density covariance). However, the analytic theory assumes continuous population density, and IGRs are calculated for infinitesimally rare invaders that have infinite time to find their preferred habitat and regrow, without ever experiencing intraspecific competition. Here we ask if the disparity between spatial and temporal mechanisms persists when individuals are, instead, discrete and occupy finite amounts of space. We present a simulation-based approach to quantifying IGRs in this situation, building on our previous approach for spatially non-varying habitats. As expected, we found that spatial mechanisms are weakened; unexpectedly, the contribution to IGR from growth-density covariance could even become negative, opposing coexistence. We also found shifts in which demographic parameters had the largest effect on the strength of spatial coexistence mechanisms. Our substantive conclusions are statements about one model, across parameter ranges that we subjectively considered realistic. Using the methods developed here, effects of individual discreteness should be explored theoretically across a broader range of conditions, and in models parameterized from empirical data on real communities.