Diversity in neotropical wet forests during the Cenozoic is linked more to atmospheric CO2 than temperature

Models generally predict a response in species richness to climate, but strong climate-diversity associations are seldom observed in long-term (more than 10(6) years) fossil records. Moreover, fossil studies rarely distinguish between the effects of atmospheric CO2 and temperature, which limits their ability to identify the causal controls on biodiversity. Plants are excellent organisms for testing climate-diversity hypotheses owing to their strong sensitivity to CO2, temperature and moisture. We find that pollen morphospecies richness in an angiosperm-dominated record from the Palaeogene and early Neogene (65-20 Ma) of Colombia and Venezuela correlates positively to CO2 much more strongly than to temperature (both tropical sea surface temperatures and estimates of global mean surface temperature). The weaker sensitivity to temperature may be due to reduced variance in long-term climate relative to in higher latitudes, or to the occurrence of lethal or sub-lethal temperatures during the warmest times of the Eocene. Physiological models predict that productivity should be the most sensitive to CO2 within the angiosperms, a prediction supported by our analyses if productivity is linked to species richness; however, evaluations of non-angiosperm assemblages are needed to more completely test this idea.