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The latitudinal temperature gradient is a fundamental state parameter of the climate system tied to the dynamics of heat transport and radiative transfer. Thus, it is a primary target for temperature proxy reconstructions and global climate models. However, reconstructing the latitudinal temperature gradient in past climates remains challenging due to the scarcity of appropriate proxy records and large proxy–model disagreements. Here, we develop methods leveraging an extensive compilation of planktonic foraminifera δ 18 O to reconstruct a continuous record of the latitudinal sea-surface temperature (SST) gradient over the last 95 million years (My). We find that latitudinal SST gradients ranged from 26.5 to 15.3 °C over a mean global SST range of 15.3 to 32.5 °C, with the highest gradients during the coldest intervals of time. From this relationship, we calculate a polar amplification factor (PAF; the ratio of change in >60° S SST to change in global mean SST) of 1.44 ± 0.15. Our results are closer to model predictions than previous proxy-based estimates, primarily because δ 18 O-based high-latitude SST estimates more closely track benthic temperatures, yielding higher gradients. The consistent covariance of δ 18 O values in low- and high-latitude planktonic foraminifera and in benthic foraminifera, across numerous climate states, suggests a fundamental constraint on multiple aspects of the climate system, linking deep-sea temperatures, the latitudinal SST gradient, and global mean SSTs across large changes in atmospheric CO 2 , continental configuration, oceanic gateways, and the extent of continental ice sheets. This implies an important underlying, internally driven predictability of the climate system in vastly different background states. 

Abstract Spines and rhizopodia play an important role in the feeding behavior, symbiont ecology, shell geochemistry, and density and drag of planktonic foraminifera. However, there are few empirical data on planktonic foraminifera in situ, and these delicate structures are disturbed on capture. Here, we report spine and rhizopod measurements from underwater images obtained in the California Current System near La Jolla, California by Zooglider, a new autonomous zooplankton-sensing glider. Across all observed species, we find that spine length and flexibility correlate with test size and that spines increase the effective prey encounter volume of spinose foraminifera by two to three orders of magnitude. Our data also yielded several novel observations regarding hastigerinid foraminifera (Hastigerinella digitata and Hastigerina pelagica), a group of unusually large planktonic foraminifera that are abundant in our dataset below 250 m. First, the effective encounter volume of hastigerinid foraminifera can be very large: our largest specimen occupies almost 40 cm3 (about the size of a golf ball), while the median specimen occupies 5.3 cm3 (about the size of a cherry). Second, the majority of hastigerinid foraminifera in our dataset have asymmetric bubble capsules, which are most frequently oriented with their bubbles on the upward side of the test, consistent with the hypothesis that the bubble capsule is positively buoyant. Third, 16% of hastigerinid foraminifera in our dataset have dispersed bubble capsules with detached bubbles distributed along the spines and rhizopodia, consistent with a regular source of natural disturbance. Taken together, our observations suggest that hastigerinid foraminifera play a larger role as mesopelagic predators in the California Current System than previously recognized.