Evolutionary ecology of Early Paleocene planktonic foraminifera: size, depth habitat and symbiosis

The carbon stable isotope (delta C-13) composition of the calcitic tests of planktonic foraminifera has an important role as a geochemical tracer of ocean carbon system changes associated with the Cretaceous/Paleogene (K/Pg) mass extinction event and its aftermath. Questions remain, however, about the extent of delta C-13 isotopic disequilibrium effects and the impact of depth habitat evolution on test calcite delta C-13 among rapidly evolving Paleocene species, and the influence this has on reconstructed surface-to-deep ocean dissolved inorganic carbon (DIC) gradients. A synthesis of new and existing multispecies data, on the relationship between delta C-13 and delta O-18 and test size, sheds light on these issues. Results suggest that early Paleocene species quickly radiated into a range of depths habitats in a thermally stratified water column. Negative 680 gradients with increasing test size in some species of Praemurica suggest either ontogenetic or ecotypic dependence on calcification temperature that may reflect depth/light controlled variability in symbiont photosynthetic activity. The pattern of positive delta C-13 test-size correlations allows us to (1) identify metabolic disequilibrium delta C-13 effects in small foraminifera tests, as occur in the immediate aftermath of the K/Pg event, (2) constrain the timing of evolution of foraminiferal photosymbiosis to 63.5 Ma, similar to 0.9 Myr earlier than previously suggested, and (3) identify the apparent loss of symbiosis in a late-ranging morphotype of Praemurica. These findings have implications for interpreting delta C-13 DIC gradients at a resolution appropriate for incoming highly resolved K/Pg core records.