Marine biodiversification in response to evolving phytoplankton stoichiometry

Diversification of the marine biosphere is intimately linked to the evolution of the biogeochemical cycles of carbon, nutrients, and primary productivity. A meta-analysis of the ratio of carbon-to-phosphorus buried in sedimentary rocks during the past 3 billion years indicates that both food quantity and, critically, food quality increased through time as a result of the evolving stoichiometry (nutrient content) of eukaryotic phytoplankton. Evolving food quantity and quality was primarily a function of broad tectonic cycles that influenced not just carbon burial, but also nutrient availability and primary productivity. Increasing nutrient availability during the middle-to-Late Proterozoic culminated in the production of food (phytoplankton biomass and fresh dead organic matter) with C:P Redfield ratios sufficient to finally promote geologically-rapid biodiversification during the Proterozoic–Phanerozoic transition. This resulted in further, massive nutrient sequestration into biomass that triggered positive feedback via nutrient recycling (bioturbation, mesozooplankton grazing) on phytoplankton productivity. Increasing rates and depths of bioturbation through the Phanerozoic suggest that nutrient recycling continued to increase. Increasing bioturbation and nutrient cycling appear to have been necessary to sustain the primary productivity and “energetics” (biomass, metabolic rates, and physical activity such as predation) of the marine biosphere because of the geologically-slow input of macronutrients like phosphorus from land and the continued sequestration of nutrients into marine and terrestrial biomass.