Phytoplankton community structure is shaped both by the bottom-up influences of the physical-chemical environment and by the top-down impacts of food webs. Emergent patterns in the contemporary ocean can thus be “null hypotheses” of future changes if underlying structuring relationships remain intact but only shift spatially. To provide such a context for the California Current Ecosystem (CCE) and adjacent open-ocean ecosystems, we used a combination of digital epifluorescence microscopy and flow cytometry to investigate variability of phytoplankton biomass, composition and size structure across gradients of ecosystem richness, as represented by total autotrophic carbon (AC). Biomass of large micro-sized (>20 µm) phytoplankton increases as a power function with system richness. Nano-sized cells (2-20 µm) increase at a lower rate at low AC, and level off at high AC. Pico-sized cells (<2-µm) do not clearly dominate at low AC and decline significantly at high AC, neither predicted by competition theory. This study provides several new insights into structural relationships and mechanisms in the CCE: 1) diatoms and dinoflagellates co-dominate the micro-phytoplankton size class throughout the range of system richness; 2) nano-phytoplankton co-dominate biomass in oligotrophic (low AC) waters, suggesting widespread mixotrophy rather than direct competition with pico-phytoplankton for nutrients; and 3) the pico-phytoplankton decline at high AC impacts small eukaryotes as well as photosynthetic bacteria, consistent with a broad stimulation of grazing pressure on all bacterial-sized cells in richer systems. Observed variability in heterotrophic bacteria and nano-flagellate grazers with system richness is consistent with this mechanism.