The size structure of phytoplankton assemblages strongly influences energy transfer through the food web and carbon cycling in the ocean. We determined the macroevolutionary trajectory in the median size of dinoflagellate cysts to compare with the macroevolutionary size change in other plankton groups. We found the median size of the dinoflagellate cysts generally decreases through the Cenozoic. Diatoms exhibit an extremely similar pattern in their median size over time, even though species diversity of the two groups has opposing trends, indicating that the macroevolutionary size change is an active response to selection pressure rather than a passive response to changes in diversity. The changes in the median size of dinoflagellate cysts are highly correlated with both deep ocean temperatures and the thermal gradient between the surface and deep waters, indicating the magnitude and frequency of nutrient availability may have acted as a selective factor in the macroevolution of cell size in the plankton. Our results suggest that climate, because it affects stratification in the ocean, is a universal abiotic driver that has been responsible for macroevolutionary changes in the size structure of marine planktonic communities over the past 65 million years of Earth's history.cell size ͉ climate change ͉ dinoflagellates ͉ evolution ͉ food webs M arine phytoplankton are a polyphyletic group of unicellular or colonial photoautotrophs (1) that range in size from Ͻ1 m to Ͼ1 mm in equivalent spherical diameter, corresponding to Ͼ8 orders of magnitude variation in cell volume. Because cell size influences nutrient uptake kinetics, photosynthesis, respiration, growth, and sinking rates (2-5), as well as genome size and rate of evolution (6, 7), the size structure of phytoplankton assemblages strongly inf luences energy transfer through the food web and tempo of evolution in the sea (8-10). Based on analyses of fossil records over the Cenozoic, macroevolutionary changes in cell size have been reported for marine diatoms (11), coccolithophorids (12), and the amoeboid planktonic foraminifera (13-15); however, the underlying causes of such changes remain unclear. Broadly, hypotheses accounting for macroevolutionary trends fall into two groups: (i) specific biotic or abiotic forcings unique to each taxon or (ii) taxonspecific responses to a universal abiotic factor. Here, based on an analysis of the fossil record, we report that, like the other plankton groups examined, dinoflagellates exhibit an active macroevolutionary change in size in concert with changes in the thermal contrast between the surface and deep ocean. These results strongly suggest that a universal abiotic driver related to climate is responsible for macroevolutionary changes in the size structure of marine planktonic communities over the past 65 million years of Earth's history.Dinoflagellates are a group of unicellular eukaryotic flagellates characterized by a longitudinal and a transverse flagellum and are often armored with cellulosic plates. There are Ϸ2,...