Fate of calcifying tropical symbiont-bearing large benthic foraminifera: living sands in a changing ocean.

Concerns regarding the response of calcifiers in future warmer and more acidic oceans have been raised in many studies. Tropical large benthic foraminifera (LBF) are important carbonate producers that reside in coral reefs worldwide. Similar to corals, these organisms live in symbioses with microalgae, which promote high calcification rates. The contribution of LBFs to reef sediments is under threat due to climate change. In this review, we synthesize research conducted on the effects of increased temperature and acidification on these organisms, and assess the potential impacts on reef carbonate production. A meta-analysis of all available experimental data (18 publications, 84 individual experiments) on the effects of ocean warming and acidification on LBF holobiont health was performed using log-transformed response ratios (LnRR) comparing present-day ambient and projected future scenarios. For the latter, we used Representative Concentration Pathway 8.5 from the Intergovernmental Panel on Climate Change, which projects changes of +4 °C and -0.3 pH units by the year 2100. Overall, a general negative trend on holobiont growth was observed across most species of LBFs in response to both stressors. The only exception was the hyaline species (porous CaCO3 test composed of interlocking microcrystals) that have diatom symbionts. Species in this group appear resilient to future ocean acidification scenarios. Differences in the response of LBF species to warming and acidifying oceans may be due to (1) differences in the carbonate species' use in formation of the CaCO3 skeleton (CO2 vs. CO3(2-)), (2) varied responses of the symbiont types (diatom, dinoflagellate, rhodophyte) to stressors, or (3) the degree of nutritional dependence of the host to its symbiont. We also summarize current estimates of carbonate production by LBFs to provide a context of their contribution to reefs. Finally, we outline major gaps in knowledge in addressing the potential for LBF species persistence in a changing ocean.