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The chemical composition of fossil foraminiferal shells (tests) is widely used as tracers for past ocean chemistry. It is, therefore, important to understand how different (trace) elements are transported and incorporated into these tests from adjacent seawater. The elemental distribution within the walls of foraminiferal tests might be used to differentiate between proposed transport mechanisms. Here, the microdistribution of Mg, Mn, Na, S, and Sr in tests of three species of foraminifera, known to have contrasting test chemistry, is investigated by a combination of electron probe microanalysis (EPMA) and nanoscale secondary ion mass spectrometry (nanoSIMS), micro-X-ray fluorescence (µXRF), and micro-X-ray absorption near-edge structure (µXANES) analyses. The three investigated species are the symbiont-barren Ammonia sp. T6 and Bulimina marginata, which precipitate a low-Mg calcite test, and the symbiont-bearing species Amphistegina lessonii, which produces a test with intermediate Mg content. Because all analyzed tests were formed under controlled and identical laboratory conditions, the observed distributions of elements are not due to environmental variability but are a direct consequence of the processes involved in calcification or, in the case of A. lessonii, possibly symbiont activity. Despite some variability in elemental microdistribution between specimens from a given species, our combined dataset shows species-specific distributions of the elements (e.g., peak heights and/or band-widths) and also a systematic colocation of Mg, Na, S, and Sr for all three species, suggesting a coupled or simultaneous uptake, transport, and incorporation of these elements during chamber addition. The observed trace element patterns generally reflect a laminar calcification model, suggesting that heterogeneity of these elements is intrinsically linked to chamber addition. Although the incorporation of redox-sensitive Mn depends on the Mn concentration of the culture medium, the Mn distribution observed in Ammonia sp. suggests that Mn transport is similarly linked to laminar calcification dynamics. However, for B. marginata, Mn banding was sometimes anticorrelated with Mg banding, suggesting that (bio)availability, uptake, and transport of Mn differ from those for van Dijk et al. Chemical Heterogeneities in Foraminiferal Shells Ammonia sp. Our results from symbiont-bearing A. lessonii suggest that the activity of symbionts (i.e., photosynthesis/respiration) may influence the incorporation of Mn owing to alternation of the chemistry in the microenvironment of the foraminifera, an important consideration in the development of this potential proxy for past oxygenation of the oceans.

Abstract. Shell chemistry of foraminiferal carbonate proves to be useful in reconstructing past ocean conditions. A new addition to the proxy toolbox is the ratio of sulfur (S) to calcium (Ca) in foraminiferal shells, reflecting the ratio of SO42- to CO32- in seawater. When comparing species, the amount of SO42- incorporated, and therefore the S∕Ca of the shell, increases with increasing magnesium (Mg) content. The uptake of SO42- in foraminiferal calcite is likely connected to carbon uptake, while the incorporation of Mg is more likely related to Ca uptake since this element substitutes for Ca in the crystal lattice. The relation between S and Mg incorporation in foraminiferal calcite therefore offers the opportunity to investigate the timing of processes involved in Ca and carbon uptake. To understand how foraminiferal S∕Ca is related to Mg∕Ca, we analyzed the concentration and within-shell distribution of S∕Ca of three benthic species with different shell chemistry: Ammonia tepida, Bulimina marginata and Amphistegina lessonii. Furthermore, we investigated the link between Mg∕Ca and S∕Ca across species and the potential influence of temperature on foraminiferal S∕Ca. We observed that S∕Ca is positively correlated with Mg∕Ca on a microscale within specimens, as well as between and within species. In contrast, when shell Mg∕Ca increases with temperature, foraminiferal S∕Ca values remain similar. We evaluate our findings in the light of previously proposed biomineralization models and abiological processes involved during calcite precipitation. Although all kinds of processes, including crystal lattice distortion and element speciation at the site of calcification, may contribute to changes in either the amount of S or Mg that is ultimately incorporated in foraminiferal calcite, these processes do not explain the covariation between Mg∕Ca and S∕Ca values within specimens and between species. We observe that groups of foraminifera with different calcification pathways, e.g., hyaline versus porcelaneous species, show characteristic values for S∕Ca and Mg∕Ca, which might be linked to a different calcium and carbon uptake mechanism in porcelaneous and hyaline foraminifera. Whereas Mg incorporation might be controlled by Ca dilution at the site of calcification due to Ca pumping, S is linked to carbonate ion concentration via proton pumping. The fact that we observe a covariation of S and Mg within specimens and between species suggests that proton pumping and Ca pumping are intrinsically coupled across multiple scales.

Large benthic foraminifera (LBF) are marine calcifying protists that commonly harbor algae as symbionts. These organisms are major calcium carbonate producers and important contributors to primary production in the photic zones. Light is one of the main known factors limiting their distribution, and species of this group developed specific mechanisms that allow them to occupy different habitats across the light gradient. Operculina ammonoides (Gronovius, 1781) is a planispiral LBF that has two main shell morphotypes, thick involute and flat evolute. Earlier studies suggested morphologic changes with variation in water depth and presumably light. In this study, specimens of the two morphotypes were placed in the laboratory under artificial low light and near the sea floor at depths of 15 m, 30 m, and 45 m in the Gulf of Aqaba-Eilat for 23 days. Differences in growth and symbionts content were evaluated using weight, size, and chlorophyll a. Our results show that O. ammonoides exhibit morphological plasticity when constructing thinner chambers after relocation to low light conditions, and adding more weight per area after relocation to high light conditions. In addition, O. ammonoides exhibited chlorophyll content adaptation to a certain range of light conditions, and evolute specimens that were acclimatized to very low light did not survive relocation to a high light environment, possibly due to photo-oxidative stress.

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Larger benthic foraminifera (LBF) are unicellular eukaryotic calcifying organisms and an important component of tropical and subtropical modern and ancient oceanic ecosystems. They are major calcium carbonate producers and important contributors to primary production due to the photosynthetic activity of their symbiotic algae. Studies investigating the response of LBF to seawater carbonate chemistry changes are therefore essential for understanding the impact of climate changes and ocean acidification (OA) on shallow marine ecosystems. In this study, calcification, respiration, and photosynthesis of the widespread diatom-bearing LBF Operculina ammonoides were measured in laboratory experiments that included manipulation of carbonate chemistry parameters. pH was altered while keeping dissolved inorganic carbon (DIC) constant, and DIC was altered while keeping pH constant. The results show clear vulnerability of O. ammonoides to low pH and CO 3 2− under constant DIC conditions, and no increased photosynthesis or calcification under high DIC concentrations. Our results call into question previous hypotheses, suggesting that mechanisms such as the degree of cellular control on calcification site pH/DIC and/or enhanced symbiont photosynthesis in response to OA may render the hyaline (perforate and calcitic-radial) LBF to be less responsive to OA than porcelaneous LBF. In addition, manipulating DIC did not affect calcification when pH was close to present seawater levels in a model encompassing the total population size range. In contrast, larger individuals (>1,200 μm, >1 mg) were sensitive to changes in DIC, a phenomenon we attribute to their physiological requirement to concentrate large quantities of DIC for their calcification process. Plain Language Summary Large benthic foraminifera (LBF) are unicellular marine organisms that build calcium carbonate shells and live in association with oxygen-producing photosymbiotic algae. They are a significant component of tropical and subtropical marine ecosystems and play an important role in the chemical and biological balance of the oceans. Studies investigating the response of these organisms to ocean acidification are essential for understanding the impact of future ocean acidification on shallow subtropical costal systems. In this study, growth rates and oxygen production of a widespread LBF species were measured in laboratory experiments that manipulated key carbonate chemistry parameters. The results show clear vulnerability of this species to lower pH, regardless of the levels of dissolved inorganic carbon (which is the sum of all inorganic carbon in seawater). This finding calls into question recent suggestions that differences in the vulnerability of foraminiferal species relates to shell structure and type of symbionts. Rather, our results highlight that these factors do not necessarily offer protection against ocean acidification. Overall, this study contributes to our understanding of the impact of increasing carbon dioxide and ocean acidification on a key group...

<p><strong>Abstract.</strong> Shell chemistry of foraminiferal carbonate proves to be useful in reconstructing past ocean conditions. A new addition to the proxy toolbox is the ratio of sulfur (S) to calcium (Ca) in foraminiferal shells, reflecting the ratio of SO<sub>4</sub><sup>2−</sup> to CO<sub>3</sub><sup>2−</sup> in seawater. When comparing species, the amount of SO<sub>4</sub><sup>2−</sup> incorporated, and therefore the S/Ca of the shell, increases with increasing magnesium (Mg) content. The uptake of SO<sub>4</sub><sup>2−</sup> in foraminiferal calcite is likely coupled to carbon uptake, while the incorporation of Mg is more likely related to Ca uptake since this element substitutes Ca in the crystal lattice. The relation between S and Mg incorporation in foraminiferal calcite therefore offers the opportunity to investigate the timing of processes involved in Ca and carbon uptake. To understand how foraminiferal S/Ca is related to Mg/Ca, we analyzed the concentration and within-shell distribution of S/Ca of three benthic species with different shell chemistry: <i>Ammonia tepida</i>, <i>Bulimina marginata</i> and <i>Amphistegina lessonii</i>. Furthermore, we investigated the link between Mg/Ca and S/Ca across species and the potential influence of temperature on foraminiferal S/Ca. We observed that S/Ca is positively correlated with Mg/Ca on microscale within specimens, as well as between and within species. In contrast, when shell Mg/Ca increases with temperature, foraminiferal S/Ca values remain similar. We evaluate our findings in the light of previously proposed biomineralization models and abiological processes involved during calcite precipitation. Although all kinds of processes, including crystal lattice distortion and element speciation at the site of calcification, may contribute to changes in the amount of S and Mg that is ultimately incorporated in foraminiferal calcite, these processes do not explain the consistent co-variation between Mg/Ca and S/Ca values. We observe that groups of foraminifera with different calcification pathways, e.g. hyaline versus porcelaneous species, show characteristic values for S/Ca and Mg/Ca, which might be linked to a different calcium and carbon uptake mechanism in porcelaneous and hyaline foraminifera. Whereas Mg incorporation is linked to the Ca-pump, S is linked to carbonate ion concentration via proton pumping. The fact that we observe coupled behavior of S and Mg, within specimens and between species suggests that proton pumping and Ca pumping are intrinsically coupled across scales.</p>