Coexistence of Photosynthetic Marine Microorganisms, Viruses and Grazers: Towards Integration in Ocean Ecosystem Models

Photosynthetic microorganisms are responsible for primary production at the base of the marine food web and influence global biogeochemistry. Their growth is balanced by mortality processes, including zooplankton grazing and viral lysis. These predators coexist despite competing for the same microorganisms. Here, we develop a community model of photosynthetic microorganisms, grazers and viruses that incorporates elemental quotas and is suitable for ocean ecosystem models. We evaluate the extent to which coexistence is facilitated by: (i) explicit infected phytoplankton; (ii) heterogeneity in susceptibility to viral infection; and (iii) higher-order mortality for the predators. We show a trade-off between the virus latent period and virulence in facilitating coexistence. The latent period generates oscillations that reduce the growth rate of the free virus, promoting coexistence. Heterogeneity in susceptibility supports coexistence through resource partitioning, while higher-order mortality widens the coexistence regime. The model outcomes are sensitive to viral life history traits, including the percentage of infected cells and the balance between virally- and zooplankton-induced mortality. Leveraging algebraic model equilibria, we identify parameter combinations that yield realistic ecological properties in simplified epipelagic environments. Our models suggest that efforts to embed virus dynamics in ocean ecosystem models should include moderate to strong resistance to viral infection.