Competition and allelopathy with resource storage: two resources

Allelopathy is added to a familiar mathematical model of competition between two species for two essential resources in a chemostat environment. Both species store the resources, and each produces a toxin that induces mortality in the other species. The corresponding model without toxins displays outcomes of competitive exclusion independent of initial conditions, competitive exclusion that depends on initial conditions (bistability), and globally stable coexistence, depending on tradeoffs between competitors in growth requirements and consumption of the resources. Introducing toxins that act only between, and not within species, can destabilize coexistence leading to bistability or other multiple attractors. Invasibility of the missing species into a resident׳s semitrivial equilibrium is related to competitive outcomes. Mutual invasibility is necessary and sufficient for a globally stable coexistence equilibrium, but is not necessary for coexistence at a locally stable equilibrium. Invasibility of one semitrivial equilibrium but not the other is necessary but not sufficient for competitive exclusion independent of initial conditions. Mutual non-invasibility is necessary but not sufficient for bistability. Numerical analysis suggests that when competitors display bistability in the absence of toxin production, increases in the overall magnitude of resource supply cause bistability to arise over a larger range of supply ratios between the two resources. When competitors display coexistence in the absence of toxin production, increases in overall resource supply destabilize coexistence and produce bistability or other configurations of multiple attractors over large ranges of supply ratios. The emergence of multiple attractors at high resource supplies suggests that blooms of harmful algae producing allelopathic toxins could be difficult to predict under such rich conditions.