Oceanic upwelling systems are important for the Earth's atmospheric and marine carbon budget (Toggweiler & Sarmiento, 1985;Watson & Naveira Garabato, 2006). In upwelling regions, relatively cold, nutrient-rich and carbon-dioxide-(CO 2 ) bearing waters from below the thermocline typically characterize the surface-water hydrography. Such conditions allow for enhanced marine primary productivity and, thus, removal of the upwelled CO 2 from the surface-ocean through organic-matter export into the deep sea. The efficiency of this so-called biological pump is of particular importance in the Eastern Equatorial Pacific (EEP) where coastal and equatorial upwelling support >10% of the biological production in the present-day oceans (Pennington et al., 2006). Across the Plio-Pleistocene transition-a time interval that is highly relevant for the understanding of anthropogenic warming because it comprises climatic boundary conditions and atmospheric CO 2 levels as they are expected in the near future (e.g., Dowsett et al., 2013; Martinez-Boti et al., 2015;Robinson et al., 2008)-major shifts in marine export production occurred in the global oceans (Lawrence et al., 2013, and references therein). However, modes and drivers of these changes within the EEP upwelling regime are not entirely clear.Since the early Pliocene, the Earth underwent a transition from a warmer climate state and Anthropocene-like atmospheric CO 2 (>400 parts per million by volume [ppmv]) lacking large ice sheets in the Northern Hemisphere to a progressively cooler climate state and pre-industrial-like atmospheric CO 2 (<280 ppmv) during the Pleistocene with a stronger response of the climate/cryosphere system to orbital forcing (Lisiecki & Raymo, 2005; Martinez-Boti et al., 2015;Seki et al., 2010). Over the past ∼5 Myr and thus this transitional period, a close coupling between biological production and sea-surface temperatures in the equatorial upwelling system of the East Pacific Ocean has been documented on glacial-interglacial