Record of Foraminifera test composition throughout the Phanerozoic.

Marine calcifiers produce calcareous structures (e.g. shells, skeletons or tests) and are therefore sensitive to ocean chemistry. Nevertheless, the long-term evolutionary consequences of marine carbonate changes are not well understood. This article compares calcareous and non-calcareous responses to ocean chemistry changes throughout the Phanerozoic Eon (541 million years ago to present). To accomplish this, we calculated proportional wall-type diversity, origination rates and extinction rates for 2282 benthic foraminiferal genera. Calcareous origination and extinction rates fluctuated throughout the Palaeozoic Era (541-251.9 million years ago), but during the Mesozoic Era (251.9-66 million years ago), calcareous origination and extinction rates stabilized following the evolution of pelagic calcifiers. Despite variations in Cenozoic Era (66-0 million years ago) foraminifera diversity, calcareous wall types maintained around 77% proportional diversity. Although calcareous wall-type extinction rates decline during the Mesozoic and Cenozoic, Phanerozoic foraminifera wall-type changes during individual events are largely contingent upon contemporaneous conditions rather than overarching trends. Of the Big Five mass extinction events, calcareous wall-type proportions only decreased at the end-Permian (73% to 26% diversity) and end-Triassic (56% to 50% diversity). These results suggest long-term ocean chemistry changes were not the main driver of foraminiferal wall-type diversity through time.