The Plio-Pleistocene encompasses large-scale changes in climate and may provide an analog for the function of Earth's ecosystems in a high carbon dioxide (CO 2) world. Pliocene climate was characterized by atmospheric CO 2 levels similar to today, reduced sea surface temperature (SST) gradients, and events such as the restriction of the Central American and Indonesian Seaways (Cane & Molnar, 2001; Fedorov et al., 2013). Despite its relative warmth and stability, the Pliocene is defined by long-term cooling that is potentially related to tectonic plate movement and consequent changes in ocean currents and pCO 2 drawdown, especially in the Indo-Pacific region (Cane & Molnar, 2001). The Plio-Pleistocene transition around 2.7 Ma is marked by the breakdown of Pliocene climate conditions, as strong SST gradients between the high-and low-latitudes and across the Pacific Ocean basin were established, and Northern Hemisphere glaciation intensified (Fedorov et al., 2013). The development of stronger zonal SST gradients in the Pacific Ocean was likely related to gradual thermocline shoaling in the Pliocene and an increase in tropical upwelling that led to strengthened Walker circulation (Ravelo et al., 2004). The interaction between these ocean-atmosphere feedbacks and changes in insolation potentially magnified the climate's sensitivity to obliquity, leading to the 41 kyr pacing of higher amplitude glacial cycles in the early Pleistocene (Ravelo et al., 2004).