Losers in the ‘Rock-Paper-Scissors’ game: The role of non-hierarchical competition and chaos as biodiversity sustaining agents in aquatic systems

Processes occurring within small areas (patch-scale) that influence species richness and spatial heterogeneity of larger areas (landscape-scale) have long been an interest of ecologists. This research focused on the role of patch-scale deterministic chaos arising in phytoplankton assemblages characteristic of “Rock-Paper-Scissors” population dynamics (i.e., competitively non-hierarchical). We employed a simple 2-patch model configuration with lateral mixing and through-flow, and tested the robustness of species richness at the scale of the landscape and spatial heterogeneity. Three different assemblages were used that in a dimensionless box model configuration exhibited chaotic behavior. Our results showed that when a spatial dimension was added to the model configuration, and when all species were shared between patches (i.e., no invading populations), chaos-induced species richness and spatial heterogeneity were quickly reduced with the onset of mixing. While assemblages in each patch were comprised of exactly the same species, they differed in their proportional population densities due to differing stages of succession and the incidence of alternative assemblage structures. Even at very low mixing rates (0.001 d⁻¹), which produced low passive migration rates (0.1% of the total biomass per day), the incidence of high richness and heterogeneity decreased by ∼80%. Interestingly, this sensitivity was not the same for the three assemblages tested. Declines in species richness and spatial heterogeneity associated with mixing were greater in assemblages comprised of competitively dissimilar species (based on the area occupied in the resource-tradeoff space defined by the R* model). The underlying mechanisms may involve the degree to which nutrient dynamics are altered with the arrival of immigrants. Our findings suggest that in partially to well-mixed aquatic systems, the roles of patch-scale non-hierarchical competition and chaos as factors maintaining species richness and spatial heterogeneity may be limited. However, in aquatic systems that experience periods of very low mixing, or even disconnection, non-hierarchical competition and chaos might indeed contribute significantly to biodiversity.