**1. Introduction**

Planktonic foraminifera are important marine calcifiers, and the ongoing change in the oceanic carbon system makes it essential to understand the influence of environmental factors on the biomineralization of their shells [1]. Although shell weight is a prominent, easily measured feature of foraminiferal tests that has direct implications for the carbon cycle and carbonate budgets, it is currently not being widely recorded or discussed because its exact environmental meaning is unclear. The amount of calcite deposited by planktonic foraminifera during calcification has been hypothesized to reflect a range of environmental factors. Through the years, changes in planktonic foraminifera shell weights have been linked to different biotic and abiotic parameters such as dissolution [2], carbonate ion concentration [3], optimum growth conditions [4], phosphate concentrations [5], temperature [6], or salinity [1], and thus the various studies have used foraminifera shell weights each time as a different

proxy. Recent studies have shown that planktonic foraminifera can alter their shell mass according to ambient seawater density [7] and that the degree of this alteration in time is a function of latitude [8].

Planktonic organisms are able to biosynthesize out of equilibrium with their ambient environment by maintaining chemical gradients. However, as passive floaters, they must always retain equilibrium with the seawater in order to remain afloat at certain depths. It can thus be argued that plankton physiology is more sensitive to the physical rather than chemical characteristics of seawater. Because planktonic organisms lack active floatation devices, their only inert way to counterbalance seawater buoyancy changes and remain at certain depths is to modify their shell mass [9]. Shell biomineralization must thus be a function of both chemical and physical seawater properties. Although the effects of ocean chemistry on plankton have been extensively studied, there is currently a lack of literature on the effects of physical oceanic properties such as buoyancy, density, or pressure, which very likely affect foraminifera physiology and morphology [10]. Here, we examine the shell mass of the planktonic foraminifera species *Globigerina bulloides* (NCBI:txid69025) from a sediment core of the northeastern tropical Atlantic Ocean through the two most recent climatic cycles. After assessing the preservation of the foraminifera tests, we report consistent shell weights, and thus steady foraminiferal calcification, independent of atmospheric *p*CO2, in agreement with a Pliocene Caribbean record [11]. We thus attribute this consistency to the stability of the hydrological conditions over time at the tropics, because the tropical environment is strongly associated with the notion of physical and chemical stability [12,13].

The stability in the foraminifera shell weights is briefly interrupted midway through Termination II, where elevated weights are recorded. In order to understand this feature, we used standard geochemical analyses (δ <sup>18</sup>O, Mg/Ca) for a set of samples that we combined with high resolution X-ray computed tomography (CT) to evaluate potential changes in the thickness of foraminifera shells [14] along with other biometric characteristics. The geochemical analyses confirmed a relationship between shell mass and water density, which is further supported by the µCT data that indicate clay contamination as the cause of the elevated shell weights. The µCT analysis also allowed the determination of cell volumes, volume normalized weights (i.e., shell density), or porosities, and has proven a valuable tool in the study of foraminifera shells. The present record provides new evidence on the response of planktonic calcifiers to ocean acidification that will help to better constrain the role that shell mass variations have on the sedimentary calcite budget and the carbon cycle.

#### **2. Location and Oceanographic Setting**

GeoB 8502 (19◦13.27′ N, 18◦56.04′ W) is a Tropical North Atlantic pelagic site at the lower reaches of the Cap Timiris Canyon, approximately 250 km offshore the Mauritanian coast (Figure 1). The canyon head abuts the Tamanrasset River System, which, although not discharging under present-day climate conditions, ranks among the largest river systems worldwide. The mouth of this potential river system is located off Cap Timiris, but its flow pathways are at present covered by extensive Saharan sand dunes [15]. Presently, the Senegal River is the northernmost active drainage system of West Africa. Core GeoB 8502-2 was retrieved from 2956 m water depth on the lower Northwestern (NW) African continental rise and consists of levee sediments that are predominantly hemipelagic deposits.

Modern climate over the NW African margin is governed by the dynamics of the West African Monsoon, which is associated with the seasonal latitudinal shifts of the intertropical convergence zone (ITCZ) [16]. In winter, the equatorward displacement of the ITCZ (5◦ N) causes a southward shift of dry subtropical air masses and is associated with the development of strong easterly Saharan Air Layer winds. The southward shift of the ITCZ and wind development cause dust transport from the Sahara [17]. The dust plume is generally located between 15 and 25◦ N along an E–W axis over the tropical Atlantic Ocean [18]. During boreal summer, dry subtropical air is shifted northward as the ITCZ is located around 20◦ N (Figure 1). This represents the onset of the rainy season (summer monsoon), with heavy rainfall and changes in atmospheric circulation [19]. Trade winds

from the southern hemisphere, loaded with water vapor, penetrate north to the West African continent. The moisture-laden air spreads over the ocean and continent, permitting heavy rains over the area.

As part of the Eastern Boundary Current system, the Mauritanian upwelling region is one of the major upwelling areas in the Atlantic Ocean [20]. Along the NW African margin, the temporal dynamics of the coastal upwelling is driven basically by the intensity of the northeast trade-winds, itself dependent on the seasonal intertropical convergence zone (ITCZ) migration [21,22] on a perennial basis, producing cold nutrient-rich surface waters with modern sea surface temperatures (SSTs) as low as 16 ◦C. The studied area is under the influence of the major return branch of the subtropical gyre, the Canary Current (CC), flowing southward along the north-west African coast, then becoming the North Equatorial Current (NEC) when turning southwestward and leaving the African continent. Further south, the westward flowing North Equatorial Counter Current (NECC) is encountered, which transports low salinity water in the area and is known to show a strong seasonal cycle position with maximum velocities when the ITCZ is located at the northernmost position and weak velocities in northern spring [22].

The main water masses encountered in the upwelling region are the Tropical Surface Water (TSW), the North and South Atlantic Central Waters (NACW and SACW), and the Antarctic Intermediate Water (AAIW) [22]. Because of the Cape Verde frontal zone (CVFZ), which is the transition boundary zone between NACW and SACW, the upwelling is fed by two different subsurface water masses depending on latitude [23]. The CVFZ is located at about 20◦ N off Africa oriented southwestward to about 16◦ N in the central tropical Atlantic. The front is associated with a convergence at the coast between the CC conveying NACW southward and a northward flow of SACW. With SACW and NACW occupying the same density range, the front is density-compensated and results in a multitude of intrusions, filaments, and lenses [24]. The NACW is warmer and more saline compared with the SACW. Both central water masses appear in the permanent pycnocline between depths of 150 m and 600 m at temperatures greater than about 8 ◦C, below which lies the AAIW [22]. At greater depths, the core sediments are currently bathed in the carbonate saturated North Atlantic Deep Water (NADW) and may have remained so during the glacials [25].

**Figure 1.** Location of the studied core GeoB 8502-2 along the north-western African margin compared with the atmospheric and oceanographic regional settings. The Cap Timiris Canyon pathway is marked. The arrows show the pathway of the modern dominant surface currents in the area. Red and blue bands identify the migration domains of the summer and winter intertropical convergence zone (ITCZ), respectively. The dashed red band close to the core position denotes the Cape Verde frontal zone (CVFZ) location. The Tamanrasett River paleodrainage valley is indicated, as suggested by [15].

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