Chaotic mixotroph dynamics arise with nutrient loading: Implications for mixotrophy as a harmful bloom forming mechanism

Mixotrophic nutrition, in which organisms combine aspects of heterotrophic and autotrophic nutrition, is a common characteristic amongst the phytoplankton, and especially amongst harmful algae bloom (HAB) species. To explore how the addition of a mixotrophic population influences a theoretical plankton system under different enrichment levels, we employ a previously published numerical plankton model that reproduces seasonal succession dynamics. Through this work we found potential for chaos to occur as the system was enriched. Focusing on occurrence of chaos in the mixotroph population, a novel finding, we conducted a bifurcation analysis which demonstrated a period doubling route to chaos. Further, we demonstrate that the occurrence of chaotic dynamics in the model system is not an artifact of the mixotroph life-history trait values used in our initial simulations, rather chaos was found to occur under a wide range of mixotroph trait value combinations. Through our trait analysis, we also found that regardless of whether or not the system was chaotic for a specific combination of values, variation in the mixotroph population peak population density could be high between consecutive years. These findings have implications that are important for the management of nutrients in aquatic systems and HABs. Results shed light on why it may be difficult to predict the occurrence of mixotrophic HABs based solely on environmental variables, when the year-to-year peak density of the mixotroph is chaotic or highly variable with enrichment. These results also demonstrate how initiation of a mixotrophic HAB might occur with enrichment, due to erratic population spikes, when broadcast allelopathic or toxic chemicals are at low concentrations, a novel bloom initiation mechanism.