Abstract. The biogeochemical impact of coccolithophores is defined not only by their
overall abundance in the oceans but also by wide ranges in physiological
traits such as cell size, degree of calcification and carbon production
rates between different species. Species' sensitivity to environmental
forcing has been suggested to relate to their cellular PIC : POC (particulate inorganic carbon : particulate organic carbon) ratio and
other physiological constraints. Understanding both the short-term and
longer-term adaptive strategies of different coccolithophore lineages, and
how these in turn shape the biogeochemical role of the group, is therefore
crucial for modeling the ongoing changes in the global carbon cycle. Here we
present data on the phenotypic evolution of a large and heavily calcified
genus Helicosphaera (order Zygodiscales) over the past 15 million years (Myr), at two
deep-sea drill sites in the tropical Indian Ocean and temperate South
Atlantic. The modern species Helicosphaera carteri, which displays ecophysiological adaptations
in modern strains, was used to benchmark the use of its coccolith morphology
as a physiological proxy in the fossil record. Our results show that, on the single-genotype level, coccolith morphology
has no correlation with growth rates, cell size or PIC and POC production
rates in H. carteri. However, significant correlations of coccolith morphometric
parameters with cell size and physiological rates do emerge once multiple
genotypes or closely related lineages are pooled together. Using this
insight, we interpret the phenotypic evolution in Helicosphaera as a global, resource-limitation-driven selection for smaller cells, which appears to be a common
adaptive trait among different coccolithophore lineages, from the warm and
high-CO2 world of the middle Miocene to the cooler and low-CO2
conditions of the Pleistocene. However, despite a significant decrease in
mean coccolith size and cell size, Helicosphaera kept a relatively stable PIC : POC ratio (as inferred
from the coccolith aspect ratio) and thus highly conservative
biogeochemical output on the cellular level. We argue that this supports its
status as an obligate calcifier, like other large and heavily calcified
genera such as Calcidiscus and Coccolithus, and that other adaptive strategies, beyond
size adaptation, must support the persistent, albeit less abundant,
occurrence of these taxa. This is in stark contrast with the ancestral
lineage of Emiliania and Gephyrocapsa, which not only decreased in mean size but also displayed
much higher phenotypic plasticity in their degree of calcification while becoming
globally more dominant in plankton communities.