# Stable carbon isotope gradients in benthic foraminifera as proxy for organic carbon fluxes in the Mediterranean Sea

Marc Theodor, Gerhard Schmiedl, Frans Jorissen, Andreas Mackensen

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Marc Theodor, Gerhard Schmiedl, Frans Jorissen, Andreas Mackensen. Stable carbon isotope gradients in benthic foraminifera as proxy for organic carbon fluxes in the Mediterranean Sea. Biogeo-SCIENCES, 2016, 20, pp.6385-6404. 10.5194/bg-13-6385-2016. insu-03682684

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# Stable carbon isotope gradients in benthic foraminifera as proxy for organic carbon fluxes in the Mediterranean Sea

Marc Theodor<fnref n="1" />, Gerhard Schmiedl<fnref n="1" />, Frans Jorissen<fnref n="2" />, and Andreas Mackensen<fnref n="3" />

*Correspondence to:* Marc Theodor (marc.theodor@uni-hamburg.de)

Received: 6 June 2016 – Published in Biogeosciences Discuss.: 9 June 2016

Revised: 26 October 2016 – Accepted: 8 November 2016 – Published: 30 November 2016

**Abstract.** We have determined stable carbon isotope ratios of epifaunal and shallow infaunal benthic foraminifera in the Mediterranean Sea to relate the inferred gradient of pore water $\delta^{13}C_{DIC}$ to varying trophic conditions. This is a prerequisite for developing this difference into a potential transfer function for organic matter flux rates. The data set is based on samples retrieved from a well-defined bathymetric range (400–1500 m water depth) of sub-basins in the western, central, and eastern Mediterranean Sea. Regional contrasts in organic matter fluxes and associated $\delta^{13}C_{DIC}$ of pore water are recorded by the $\delta^{13}C$ difference ($\Delta\delta^{13}C_{Umed-Epi}$) between the shallow infaunal *Uvigerina mediterranea* and epifaunal species (*Planulina ariminensis*, *Cibicidoides pachydermus*, *Cibicides lobatulus*). Within epifaunal taxa, the highest $\delta^{13}C$ values are recorded for *P. ariminensis*, providing the best indicator for bottom water $\delta^{13}C_{DIC}$. In contrast, *C. pachydermus* reveals minor pore water effects at the more eutrophic sites. Because of ontogenetic trends in the $\delta^{13}C$ signal of *U. mediterranea* of up to 1.04‰, only tests larger than 600 µm were used for the development of the transfer function. The recorded differences in the $\delta^{13}C$ values of *U. mediterranea* and epifaunal taxa ($\Delta\delta^{13}C_{Umed-Epi}$) range from –0.46 to –2.13‰, with generally higher offsets at more eutrophic sites. The measured $\delta^{13}C$ differences are related to site-specific differences in microhabitat, depth of the principal sedimentary redox boundary, and TOC content of the ambient sediment. The $\Delta\delta^{13}C_{Umed-Epi}$ values reveal a consistent relation to $C_{org}$ fluxes estimated from satellite-derived sur-

face water primary production in open-marine settings of the Alboran Sea, Mallorca Channel, Strait of Sicily, and southern Aegean Sea. In contrast, $\Delta\delta^{13}C_{Umed-Epi}$ values in areas affected by intense resuspension and riverine organic matter sources of the northern to central Aegean Sea and the canyon systems of the Gulf of Lion suggest higher $C_{org}$ fluxes compared to the values based on recent primary production. Taking regional biases and uncertainties into account, we establish a first $\Delta\delta^{13}C_{Umed-Epi}$-based transfer function for $C_{org}$ fluxes for the Mediterranean Sea.

## 1 Introduction

The stable isotope composition of benthic foraminifera is used in a wide range of paleoceanographic applications. The $\delta^{18}O$ signal of benthic foraminifera provides information on bottom water temperature and salinity, and has been applied to estimate global ice volume changes (e.g., Shackleton and Opdyke, 1973; Adkins et al., 2002; Marchitto et al., 2014). The benthic foraminiferal $\delta^{13}C$ signal is mainly used for the reconstruction of changes in deep-sea circulation, bottom water oxygen concentrations, and organic carbon fluxes to the sea floor (Curry and Lohmann, 1982; Zahn et al., 1986; McCorkle and Emerson, 1988; Mackensen and Bickert, 1999; Pahnke and Zahn, 2005). Recently, more quantitative approaches have been applied to the reconstruction of past changes in deep-water oxygenation (Stott et al., 2000;

<footnote n="1">¹Center for Earth System Research and Sustainability, Institute of Geology, University of Hamburg, Bundesstrasse 55, 20146 Hamburg, Germany</footnote>

<footnote n="2">²CNRS, UMR 6112, LPG-BIAF, Recent and Fossil Bio-Indicators, Université d'Angers, 2 Boulevard Lavoisier, 49045 Angers CEDEX, France</footnote>

<footnote n="3">³Alfred Wegener Institute, Helmholtz Centre for Polar and Marine Research, Am Alten Hafen 26, 27568 Bremerhaven, Germany</footnote>
Schmiedl and Mackensen, 2006; Hoogakker et al., 2015). There have also been attempts to use multispecies $\delta^{13}C$ records to reconstruct past organic carbon fluxes (Zahn et al., 1986; Schilman et al., 2003; Kuhnt et al., 2008). However, all of these studies lack a regional calibration based on living specimens and modern organic carbon flux data.

The $\delta^{13}C$ gradient of pore water dissolved inorganic carbon (DIC) in the uppermost surface sediment is directly related to the flux and decomposition rates of organic matter (McCorkle and Emerson, 1988; McCorkle et al., 1990; Holsten et al., 2004). With increasing depth in the sediment, more $^{13}C$ depleted organic matter ($\delta^{13}C$ around -18 to -23‰, e.g., Mackensen, 2008) is remineralized by microbial activity (McCorkle et al., 1985). This process results in $\delta^{13}C_{DIC}$ pore water depletions of up to -4‰ relative to the bottom water signal (McCorkle and Emerson, 1988; McCorkle et al., 1990; Holsten et al., 2004). The preferential release of $^{12}C$ to the pore water stops when no more organic matter (OM) is remineralized, which mostly coincides with the total consumption of electron acceptors, of which oxygen, nitrate, and sulfate are the most energy-efficient ones (McCorkle and Emerson, 1988; McCorkle et al., 1990; Koho and Pina-Ochoa, 2012, Hoogakker et al., 2015).

The $\delta^{13}C_{DIC}$ pore water gradient is reflected in the $\delta^{13}C$ signal of benthic foraminifera from defined microhabitats on and below the sediment-water interface (Grossman, 1984a, b; McCorkle et al., 1990, 1997; Rathburn et al., 1996; Mackensen and Licari, 2004; Schmiedl et al., 2004; Fontanier et al., 2006). Although benthic foraminifera can migrate through the sediment (Linke and Lutze, 1993; Ohga and Kitzazato, 1997) and living individuals may occur across a relatively wide depth interval, the $\delta^{13}C$ of a species exhibits relatively little scatter, and all specimens tend to reflect the same calcification depth (Mackensen and Douglas, 1989; McCorkle et al., 1990, 1997; Mackensen et al., 2000; Schmiedl et al., 2004). The study of McCorkle and Emerson (1988) has shown that the difference between $\delta^{13}C_{DIC}$ of bottom water and $\delta^{13}C_{DIC}$ of pore water at the depth in the sediment where oxygen approaches zero is directly related to the oxygen content of the bottom water mass. Based on this observation, the $\delta^{13}C$ difference of epifaunal (e.g., *Cibicidoides*) and deep infaunal (*Globobulimina*) taxa was used as proxy for the quantification of past changes in deep-water oxygenation (Schmiedl and Mackensen, 2006; Hoogakker et al., 2015). In well-oxygenated bottom waters, enhanced organic matter fluxes and decomposition rates result in steepening $\delta^{13}C_{DIC}$ gradients in the uppermost sediment, which is then reflected by the $\delta^{13}C$ difference between epifaunal and shallow infaunal (e.g., *Uvigerina*) species (Zahn et al., 1986; Mackensen et al., 2000; Brückner and Mackensen, 2008). A simple relation between observed $\delta^{13}C$ gradients and organic matter fluxes is obscured by the ability of infaunal species to shift their microhabitat in response to changing trophic conditions (Schmiedl and Mackensen, 2006; Theodor et al., 2016). Interspecific differences in the $\delta^{13}C$ composition of benthic

foraminifera are further influenced by species-specific “vital effects”, which can be as large as 1‰ (Schmiedl et al., 2004; McCorkle et al., 2008; Brückner and Mackensen, 2008) and are a reflection of metabolic processes and test calcification rates (McConnaughey, 1989a, b). Of minor impact, but still traceable, is the influence of carbonate ion concentration and alkalinity gradients in pore waters (Bemis et al., 1998). Finally, significant ontogenetic $\delta^{13}C$ trends have been documented for certain taxa, particularly for the genera *Uvigerina* and *Bolivina* (Schmiedl et al., 2004; Schumacher et al., 2010; Theodor et al., 2016).

The complexity of factors influencing the stable isotope composition of deep-sea benthic foraminifera and differences between species in different depths in the sediment motivates isotopic studies on living foraminifera in relation to their biology and microhabitat. In particular, combined ecological and biogeochemical studies on a statistically relevant number of sites and on live specimens from areas with well-defined environmental gradients are required for the establishment of reference data sets and transfer functions that could then be used for a more quantitative assessment of organic matter fluxes. The Mediterranean Sea is particularly suitable for such a study because the present deep-sea environments are characterized by systematically high oxygen contents along a gradient of trophic differences. In all basins, subsurface water masses are highly oxygenated with $O_2$ concentrations of greater than $160 \mu mol kg^{-1}$ due to frequent replenishment of intermediate water in the Levantine Sea and deep water in the Gulf of Lion, Adriatic Sea, and Aegean Sea (Wüst, 1961; Lascaratos et al., 1999; Pinardi and Masetti, 2000; Tanhua et al., 2013; Pinardi et al., 2015). The inflow of nutrients with Atlantic surface waters causes an overall west-east gradient in primary production, from values of about $225 \ g \ C \ m^{-2} \ yr^{-1}$ in the Alboran Sea to about $40 \ g \ C \ m^{-2} \ yr^{-1}$ in the extremely nutrient-depleted oligotrophic Levantine Basin (Bosc et al., 2004; López-Sandoval et al., 2011; Puyo-Pay et al., 2011; Huertas et al., 2012; Tanhua et al., 2013, Gogou et al., 2014). In areas influenced by nutrient input of larger rivers and Black Sea outflow, primary production can be locally enhanced, for example, leading to a trend of decreasing primary production values along a north-south transect in the Aegean Sea (Lykousis et al., 2002; Skliris et al., 2010). In addition, resuspension and lateral transport of organic matter can lead to locally enhanced food availability in submarine canyons and isolated basins (Puig and Palanques, 1998; Danovaro et al., 1999; Heussner et al., 2006; Canals et al., 2013).

In this study, we have compiled a data set on the stable carbon isotope composition of living and dead individuals of three epifaunal species (*Cibicidoides pachydermus*, *Planulina ariminensis*, *Cibicides lobatulus*), and one shallow infaunal species (*Uvigerina mediterranea*) from 19 Mediterranean sites. The sites are located in a well-defined depth interval (between 400 and 1500 m) and represent a wide range of trophic conditions. Adjusted for ontogenetic effects,
<figure><img src="image_3.png" /><figcaption>**Figure 1.** Location of the study areas in the Mediterranean Sea and regional bathymetric maps with locations of sample sites in the (a) Mallorca Channel, (b) Alboran Sea, (c) Gulf of Lion and Spanish slope off Barcelona, (d) Strait of Sicily, and (e) Aegean Sea.</figcaption></figure>

the $\Delta\delta^{13}C_{Umed-Epi}$ signal was compared to the microhabitat of *U. mediterranea*, the depth of the main redox boundary, TOC content, and organic carbon flux rates calculated from satellite-derived primary production or (if available) flux measurements from sediment trap studies. Major objective of this study is the development and evaluation of a transfer function for organic matter fluxes applicable to the quantification of past trophic changes in the Mediterranean Sea.

## 2 Material and methods

This study is based on a compilation of new and published isotope data of multicorer samples retrieved from various Mediterranean sub-basins covering a water depth range of 424 to 1466 m (Table 1). The study areas include the Alboran Sea and the Mallorca Channel (R/V *Meteor* cruise M69/1 in August 2006, Hübscher et al., 2010; data published in Theodor et al., 2016); the Gulf of Lion, Spanish slope off Barcelona, and Strait of Sicily (M40/4 in February 1998, Hieke et al., 1999; this study and data published in Schmiedl et al., 2004); and the Aegean Sea (M51/3 in November 2001, Hemleben et al., 2003; this study) (Fig. 1). For each station, the sediment color change from yellowish brown to greenish gray was used as an indicator for the change in redox potential from positive to negative values, which serves as an approximation of oxygen consumption and penetration in the surface sediment (Lyle, 1983; Schmiedl et al., 2000).

The upper 10 cm of the sediment were commonly sliced into 0.5 to 1 cm intervals (in the Aegean Sea into coarser intervals below 3 cm) and all samples were subsequently preserved in rose-bengal-stained alcohol (1.5 g rose bengal per 1 L of 96% ethanol) in order to stain cytoplasm of living or recently living foraminifera (Walton, 1952; Bernhard, 2000). In the laboratory, the sediment samples were wet sieved over a 63 µm sieve and, after drying at 40°C, dry sieved over a 150 µm (Aegean Sea samples) or 125 µm (remaining samples) mesh, respectively. From the coarse fraction of the different down-core intervals, stained individuals of selected epifaunal and shallow infaunal taxa were counted and the median living depths (MLDs; Theodor et al., 2016) were calculated as references for the respective microhabitat preferences. Only tests with at least three subsequent brightly red-colored chambers were considered as living. The low number of stained individuals of epifaunal taxa impeded analyses, except at Site 540B, where stained tests of *C. pachydermus* were available. Likewise, stained tests of *U. mediterranea* were absent at Sites 586 and 589.

For stable isotope measurements, stained tests (and unstained tests if no stained tests were available) of three epifaunal species (*C. pachydermus*, *P. ariminensis*, *C. lobatulus*) and one shallow infaunal species (*U. mediterranea*) were selected and each test was measured using an optical micrometer with an accuracy of 10 µm. In total, 2 stained and 63 unstained epifaunal tests as well as 155 stained and 197 unstained tests of *U. mediterranea* were measured. Individual numbers of tests measured were 1–6 for *C. pachydermus*, 1–5 for *P. ariminensis*, 1–5 for *C. lobatulus*, and 1–8 for *U. mediterranea*. The stable carbon and oxygen isotope mea-
<table><caption>**Table 1.** Position, water depth, median living depth (MLD) of *Uvigerina mediterranea*, geochemical, primary production (PP), and Corg flux values of the investigated multicorer sites. Annual PP values are averages for the year prior to sampling after data from the GlobColour project. Corg fluxes were calculated after Betzer et al. (1984) and the MLD after Theodor et al. (2016).</caption><thead><tr><th>Site</th><th>Latitude</th><th>Longitude</th><th>Station depth (m)</th><th>MLD<sub>Umed</sub> (cm)</th><th>Redox boundary depth (cm)</th><th>TOC (%)</th><th>PP (Antoine and Morel, 1996) (g C m<sup>-2</sup> a<sup>-1</sup>)</th><th>PP (Uitz et al., 2008) (g C m<sup>-2</sup> a<sup>-1</sup>)</th><th>Corg flux (Antoine and Morel, 1996) (g C m<sup>-2</sup> a<sup>-1</sup>)</th><th>Corg flux (Uitz et al., 2008) (g C m<sup>-2</sup> a<sup>-1</sup>)</th></tr></thead><tbody><tr><td>537</td><td>37°02.14′ N</td><td>13°11.33′ E</td><td>472</td><td>0.83</td><td>2.75</td><td>0.560</td><td>173.06</td><td>120.18</td><td>12.26</td><td>7.33</td></tr><tr><td>540A</td><td>42°27.69′ N</td><td>03°25.64′ E</td><td>911</td><td>0.43</td><td>2.25</td><td></td><td>203.97</td><td>160.16</td><td>10.22</td><td>7.27</td></tr><tr><td>540B</td><td>42°25.70′ N</td><td>03°41.34′ E</td><td>812</td><td>1.22</td><td>7</td><td>0.750</td><td>193.99</td><td>151.44</td><td>10.24</td><td>7.22</td></tr><tr><td>540C</td><td>41°21.04′ N</td><td>03°01.36′ E</td><td>721</td><td>0.97</td><td>4.25</td><td>0.650</td><td>179.74</td><td>138.14</td><td>9.91</td><td>6.84</td></tr><tr><td>585</td><td>36°39.60′ N</td><td>25°55.72′ E</td><td>708</td><td>2.25</td><td>21</td><td>0.430</td><td>151.13</td><td>105.86</td><td>7.85</td><td>4.75</td></tr><tr><td>586</td><td>36°34.32′ N</td><td>25°57.91′ E</td><td>424</td><td>1.00</td><td>18</td><td>0.408</td><td>151.13</td><td>105.86</td><td>10.83</td><td>6.56</td></tr><tr><td>589</td><td>36°45.19′ N</td><td>26°35.38′ E</td><td>584</td><td>2.13</td><td>14.5</td><td>0.698</td><td>150.87</td><td>105.59</td><td>8.84</td><td>5.34</td></tr><tr><td>592</td><td>37°47.65′ N</td><td>26°15.72′ E</td><td>1148</td><td>0.38</td><td>16</td><td>0.630</td><td>151.46</td><td>110.29</td><td>5.81</td><td>3.72</td></tr><tr><td>595</td><td>38°15.63′ N</td><td>25°06.17′ E</td><td>662</td><td>0.56</td><td>19</td><td></td><td>159.63</td><td>114.32</td><td>8.84</td><td>5.52</td></tr><tr><td>596</td><td>38°57.32′ N</td><td>24°45.20′ E</td><td>884</td><td>0.41</td><td>30</td><td>0.730</td><td>160.50</td><td>116.00</td><td>7.43</td><td>4.70</td></tr><tr><td>599</td><td>39°45.36′ N</td><td>24°05.61′ E</td><td>1084</td><td>0.47</td><td>16.5</td><td>0.579</td><td>195.88</td><td>138.51</td><td>8.66</td><td>5.31</td></tr><tr><td>601</td><td>40°05.22′ N</td><td>24°36.62′ E</td><td>977</td><td>0.27</td><td>6</td><td>0.750</td><td>206.68</td><td>145.42</td><td>9.97</td><td>6.07</td></tr><tr><td>602</td><td>40°13.03′ N</td><td>24°15.39′ E</td><td>1466</td><td>0.78</td><td>4</td><td>0.820</td><td>236.78</td><td>164.09</td><td>9.36</td><td>5.58</td></tr><tr><td>338</td><td>36°15.03′ N</td><td>03°24.98′ W</td><td>732</td><td>0.55</td><td>1.75</td><td>0.832</td><td>294.00</td><td>207.05</td><td>19.64</td><td>11.98</td></tr><tr><td>339</td><td>36°18.30′ N</td><td>03°08.39′ W</td><td>849</td><td>0.81</td><td>2.25</td><td>0.766</td><td>280.09</td><td>197.86</td><td>16.71</td><td>10.24</td></tr><tr><td>347</td><td>36°27.90′ N</td><td>02°55.50′ W</td><td>629</td><td>0.63</td><td>1.5</td><td>0.835</td><td>273.71</td><td>192.02</td><td>19.53</td><td>11.85</td></tr><tr><td>394</td><td>38°53.39′ N</td><td>02°38.40′ E</td><td>646</td><td>1.28</td><td>8</td><td></td><td>171.05</td><td>124.55</td><td>9.90</td><td>6.33</td></tr><tr><td>395</td><td>38°57.70′ N</td><td>02°31.51′ E</td><td>834</td><td>0.81</td><td>7</td><td>0.463</td><td>170.54</td><td>125.07</td><td>8.40</td><td>5.42</td></tr><tr><td>396</td><td>39°09.60′ N</td><td>02°28.78′ E</td><td>562</td><td>0.88</td><td>10</td><td>0.403</td><td>167.82</td><td>123.45</td><td>10.52</td><td>6.82</td></tr><tr><td>Canyon Ø</td><td>42°27.60′ N</td><td>03°29.80′ E</td><td>920</td><td>1.50</td><td>4</td><td>0.870</td><td></td><td></td><td>19.7*</td><td>19.7*</td></tr><tr><td>Canyon (Feb)</td><td>42°27.60′ N</td><td>03°29.80′ E</td><td>920</td><td>0.49</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Canyon (Aug)</td><td>42°27.60′ N</td><td>03°29.80′ E</td><td>920</td><td>2.50</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Slope Ø</td><td>42°25.60′ N</td><td>03°42.00′ E</td><td>800</td><td>1.81</td><td>11</td><td>0.720</td><td></td><td></td><td>12.8*</td><td>12.8*</td></tr><tr><td>Slope (Feb)</td><td>42°25.60′ N</td><td>03°42.00′ E</td><td>800</td><td>3.21</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Slope (Aug)</td><td>42°25.60′ N</td><td>03°42.00′ E</td><td>800</td><td>0.41</td><td></td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>

<footnote>* Schmiedl et al. (2000).</footnote>
<figure><img src="image_4.png" /><figcaption>**Figure 2.** (a) The δ¹³C of epifaunal species (*Cibicidoides pachydermus*, *Cibicides lobatulus*, *Planulina ariminensis*) for each investigated site. Each symbol represents a single measurement. Red symbols mark relocated or fossil tests that have not been used to calculate δ¹³Cₚ. Green circles show δ¹³Cₚ values used as an approximation of the δ¹³C of bottom water DIC. Details on the selection of tests and procedure for the estimation of δ¹³Cₚ values are discussed in Sect. 4.1. (b) The δ¹³Cₚ vs. water depth shows a wider scattering for the Aegean Sea than for the western Mediterranean Sea. Colored lines in the background indicate water mass end members of the Mediterranean Sea after Pierre (1999).</figcaption></figure>

surements were performed at the Alfred Wegener Institute, Helmholtz Centre for Polar and Marine Research at Bremerhaven with two Finnigan MAT 253 stable isotope ratio mass spectrometers coupled to automatic carbonate preparation devices (Kiel IV). The mass spectrometers were calibrated via international standard NBS 19 to the PDB scale, with results given in δ notation vs. VPDB. Based on an internal laboratory standard (Solnhofen limestone) measured over a 1-year period together with samples, the precision of stable isotope measurements was better than 0.06 and 0.08‰ for carbon and oxygen, respectively. The δ¹³C difference between epifaunal and shallow infaunal taxa was calculated as a proxy for the difference in δ¹³C in DIC of bottom and shallow pore water. For *U. mediterranea*, this procedure was restricted to measurements from the size fraction greater than 600 µm in order to minimize ontogenetic effects (Schmiedl et al., 2004; Theodor et al., 2016).

Total organic carbon (TOC) concentration in the surface sediment was measured with a Carlo Erba 1500 CNS analyzer with a precision of 0.02 % on weighted sample splits in tin capsules. Before measurement, CaCO₃ was removed from these weighted samples by adding 1N HCl. The TOC values of Sites 596, 601, and 602 were taken from Möbius et al. (2010a, b). Bottom water oxygen concentrations are based on CTD measurements stored in the MedAtlas data set.

Primary productivity values in surface waters of the year preceding the sampling at each site are based on satellite data of the GlobColour project, and were calculated with the algorithms of Antoine and Morel (1996) as well as Uitz et al. (2008). If available, these estimates were compared with nearby direct primary productivity and export flux measurements. The export fluxes down to the sea floor were estimated according to the function of Betzer et al. (1984) adapted by Felix (2014).

## 3 Results

Benthic foraminiferal δ¹³C values of our samples cover a range of more than 3‰, with higher average values of epifaunal species than shallow infaunal *Uvigerina mediterranea* (Table 2). The epifaunal species *Cibicidoides pachydermus*, *Cibicides lobatulus*, and *Planulina ariminensis* show average values between 1.90‰ at Site 586 (southern Aegean Sea) and -0.16‰ at Site 347 (Mallorca Channel) (Table 2, Fig. 2). The highest average epifaunal δ¹³Cₚ values are in the southern and central Aegean Sea (Sites 586, 595), while further to the north at Site 602 (northern Aegean Sea) the average δ¹³Cₚ value of 0.87‰ is among the lowest measured. At Site 540B in the Gulf of Lion, the average δ¹³Cₚ value of 1.01‰ is in good agreement with 1.00‰ measured by
<figure><img src="image_5.png" /></figure>

**Figure 3.** Correlation between δ¹³C_Umed and δ¹³C_Epi difference (Δδ¹³C_Umed-Epi) and size classes of *U. mediterranea*. For better clarity and due to the large number of measured data (Table 1), the linear regressions for each site are given, showing clear ontogenetic trends in δ¹³C_Umed due to size-independent δ¹³C_Epi values. The shown data are from live (rose-bengal-stained) and dead (unstained) individuals of *U. mediterranea* as well as for the western Mediterranean Sea (left) and Aegean Sea (right). Dashed lines represent already published data (Schmiedl et al., 2004; Theodor et al., 2016).

Schmiedl et al. (2004) at the same site. Size-dependent measurements did not reveal any ontogenetic trend in the δ¹³C signal of the epifaunal taxa (Table S1 in the Supplement).

For *U. mediterranea*, δ¹³C_Umed values vary between -1.41 and 0.85‰ for stained tests and between -1.52 and 1.77‰ for unstained tests (Table 1). The highest average values are recorded in the southern Aegean Sea, with 0.58 and 1.11‰ for stained and unstained tests, respectively. The lowest average values are recorded for the northern Aegean Sea, with -0.98 and -1.13‰ for stained and unstained tests, respectively. The variability at a single site reaches 1.38‰ in stained (Site 537) and 2.21‰ in unstained tests (Site 586). The ontogenetic δ¹³C_Umed trends are generally comparable in the western Mediterranean Sea and the Strait of Sicily, with 0.11 ± 0.03‰ 100 µm⁻¹ for stained and 0.07 ± 0.03‰ 100 µm⁻¹ for unstained tests, except for Site 396 that shows an anomalous negative trend (Table 3, Fig. 3).

In the Aegean Sea, the ontogenetic δ¹³C_Umed trends are approximately 50‰ steeper with an increase of 0.16 ± 0.04‰ 100 µm⁻¹ for stained tests. Unstained tests reveal a higher variability and a less steep slope of 0.10 ± 0.07‰ 100 µm⁻¹ (Table 3, Fig. 3). In order to avoid bias due to ontogenetic effects, only δ¹³C values of *U. mediterranea* tests larger than 600 µm were used for comparison with δ¹³C_Epi values.

The calculated Δδ¹³C_Umed-Epi values for stained tests range from -0.64‰ in the Gulf of Lion (slope site) and -0.74‰ (Site 585) to -1.29‰ in the western Mediterranean Sea (Sites 347 and 540A) to -1.85‰ in the northern Aegean Sea (Site 602) (Table 2). Due to the wider scattering of the δ¹³C values of unstained tests, Δδ¹³C_Umed-Epi values range from -0.61‰ (Site 589) to -2.0‰ (Site 602) in the Aegean Sea and from -0.55‰ (Site 540B) to -1.06‰ (Site 339) in the western Mediterranean Sea and the Strait of Sicily (Table 2). The magnitude of Δδ¹³C_Umed-Epi values exhibits a
<table><caption>**Table 2.** Average stable carbon isotope composition of selected benthic foraminifera with standard deviations. Bold values of epifaunal species were applied to estimate δ¹³C<sub>Epi</sub>. Also given are values for *Uvigerina mediterranea* tests larger than 600 µm and the difference of this species compared to the average epifaunal stable carbon isotope ratios (Δδ¹³C<sub>Umed-Epi</sub>).</caption><thead><tr><th>Site</th><th>δ¹³C<sub>Pari</sub><br>(‰ VPDB)</th><th>SD (‰)</th><th>δ¹³C<sub>Cpac</sub><br>(‰ VPDB)</th><th>SD (‰)</th><th>δ¹³C<sub>Clob</sub><br>(‰ VPDB)</th><th>SD (‰)</th><th>δ¹³C<sub>Epi</sub><br>(‰ VPDB)</th><th>SD (‰)</th></tr></thead><tbody><tr><td>537</td><td></td><td></td><td><strong>1.11</strong></td><td>0.32</td><td></td><td></td><td>1.11</td><td>0.32</td></tr><tr><td>540A</td><td><strong>1.08</strong></td><td>0.14</td><td>0.76</td><td>0.17</td><td></td><td></td><td>1.08</td><td>0.14</td></tr><tr><td>540B</td><td><strong>0.99</strong></td><td></td><td><strong>1.01</strong></td><td>0.13</td><td></td><td></td><td>1.01</td><td>0.11</td></tr><tr><td>540C</td><td>0.76</td><td></td><td><strong>1.01</strong></td><td>0.09</td><td></td><td></td><td>1.01</td><td>0.09</td></tr><tr><td>585</td><td><strong>1.32</strong></td><td>0.24</td><td></td><td></td><td></td><td></td><td>1.32</td><td>0.24</td></tr><tr><td>586</td><td><strong>1.90</strong></td><td>0.15</td><td></td><td></td><td></td><td></td><td>1.90</td><td>0.15</td></tr><tr><td>589</td><td><strong>1.34</strong></td><td>0.11</td><td></td><td></td><td></td><td></td><td>1.34</td><td>0.11</td></tr><tr><td>592</td><td><strong>1.30</strong></td><td>0.23</td><td></td><td></td><td></td><td></td><td>1.30</td><td>0.23</td></tr><tr><td>595</td><td><strong>1.87</strong></td><td>0.15</td><td></td><td></td><td></td><td></td><td>1.87</td><td>0.15</td></tr><tr><td>596</td><td><strong>0.96</strong></td><td>0.04</td><td></td><td></td><td></td><td></td><td>0.96</td><td>0.04</td></tr><tr><td>599</td><td><strong>1.76</strong></td><td>0.12</td><td></td><td></td><td></td><td></td><td>1.76</td><td>0.12</td></tr><tr><td>601</td><td></td><td></td><td><strong>0.47</strong></td><td>0.06</td><td></td><td></td><td>1.02</td><td>0.06</td></tr><tr><td>602</td><td><strong>0.87</strong></td><td></td><td>0.31</td><td>0.20</td><td></td><td></td><td>0.87</td><td></td></tr><tr><td>338</td><td><strong>1.22</strong></td><td></td><td>0.64</td><td>0.21</td><td>0.92</td><td></td><td>1.22</td><td></td></tr><tr><td>339</td><td><strong>1.22</strong></td><td>0.11</td><td>0.86</td><td></td><td>-0.12</td><td></td><td>1.22</td><td>0.11</td></tr><tr><td>347</td><td><strong>1.16</strong></td><td>0.07</td><td>0.82</td><td>0.06</td><td>-0.16</td><td></td><td>1.16</td><td>0.07</td></tr><tr><td>394</td><td></td><td></td><td>1.52</td><td>0.01</td><td><strong>0.98</strong></td><td></td><td>1.28</td><td></td></tr><tr><td>395</td><td></td><td></td><td>1.54</td><td></td><td><strong>0.80</strong></td><td>0.02</td><td>1.1</td><td>0.02</td></tr><tr><td>396</td><td><strong>1.22</strong></td><td>0.22</td><td>1.76</td><td></td><td>0.92</td><td></td><td>1.22</td><td>0.22</td></tr><tr><td>Canyon Ø</td><td></td><td></td><td>0.52</td><td>0.04</td><td></td><td></td><td>0.80</td><td>0.07</td></tr><tr><td>Slope Ø</td><td></td><td></td><td>0.39</td><td>0.09</td><td></td><td></td><td>1.00</td><td>0.06</td></tr></tbody></table>

<table><caption>**Table 2. Continued.**</caption><thead><tr><th>Site</th><th>δ¹³C<sub>Umed</sub><br>stained<br>(‰ VPDB)</th><th>SD (‰)</th><th>δ¹³C<sub>Umed</sub><br>unstained<br>(‰ VPDB)</th><th>SD (‰)</th><th>δ¹³C<sub>Umed</sub><br>stained<br>(> 600 µm)<br>(‰ VPDB)</th><th>SD (‰)</th><th>δ¹³C<sub>Umed</sub><br>unstained<br>(> 600 µm)<br>(‰ VPDB)</th><th>SD (‰)</th><th>Δδ¹³C<sub>Umed-Epi</sub><br>stained<br>(> 600 µm)<br>(‰)</th><th>SD (‰)</th><th>Δδ¹³C<sub>Umed-Epi</sub><br>unstained<br>(> 600 µm)<br>(‰)</th><th>SD (‰)</th></tr></thead><tbody><tr><td>537</td><td>0.17</td><td>0.38</td><td>-0.88</td><td>0.16</td><td>0.35</td><td>0.26</td><td>-0.82</td><td>0.03</td><td>-0.76</td><td>0.58</td><td>-1.93</td><td>0.35</td></tr><tr><td>540A</td><td>-0.46</td><td>0.21</td><td></td><td></td><td>-0.21</td><td>0.09</td><td></td><td></td><td>-1.29</td><td>0.30</td><td></td><td></td></tr><tr><td>540B</td><td>0.13</td><td>0.32</td><td>0.19</td><td>0.35</td><td>0.27</td><td>0.23</td><td>0.46</td><td>0.10</td><td>-0.74</td><td>0.34</td><td>-0.55</td><td>0.21</td></tr><tr><td>540C</td><td>-0.14</td><td>0.30</td><td>0.06</td><td>0.32</td><td>0.05</td><td>0.26</td><td>0.28</td><td>0.32</td><td>-0.97</td><td>0.41</td><td>-0.74</td><td>0.47</td></tr><tr><td>585</td><td>0.58</td><td>0.22</td><td>0.50</td><td>0.47</td><td>0.58</td><td>0.22</td><td>0.50</td><td>0.22</td><td>-0.74</td><td>0.46</td><td>-0.82</td><td>0.46</td></tr><tr><td>586</td><td></td><td></td><td>0.95</td><td>0.46</td><td></td><td></td><td>1.11</td><td>0.31</td><td></td><td></td><td>-0.79</td><td>0.46</td></tr><tr><td>589</td><td></td><td></td><td>0.51</td><td>0.46</td><td></td><td></td><td>0.73</td><td>0.39</td><td></td><td></td><td>-0.61</td><td>0.50</td></tr><tr><td>592</td><td>-0.14</td><td>0.02</td><td>0.15</td><td>0.25</td><td>-0.12</td><td>0.00</td><td>0.24</td><td>0.20</td><td>-1.42</td><td>0.23</td><td>-1.06</td><td>0.43</td></tr><tr><td>595</td><td>0.09</td><td>0.53</td><td>0.67</td><td>0.41</td><td>0.37</td><td>0.31</td><td>0.77</td><td>0.41</td><td>-1.49</td><td>0.46</td><td>-1.09</td><td>0.56</td></tr><tr><td>596</td><td>-0.38</td><td>0.38</td><td>-0.43</td><td>0.34</td><td>-0.23</td><td>0.33</td><td>-0.27</td><td>0.27</td><td>-1.19</td><td>0.37</td><td>-1.23</td><td>0.31</td></tr><tr><td>599</td><td>0.03</td><td>0.26</td><td>0.25</td><td>0.45</td><td>0.12</td><td>0.20</td><td>0.41</td><td>0.28</td><td>-1.63</td><td>0.32</td><td>-1.35</td><td>0.40</td></tr><tr><td>601</td><td>-0.53</td><td>0.27</td><td>-0.47</td><td>0.38</td><td>-0.34</td><td>0.14</td><td>-0.37</td><td>0.35</td><td>-1.36</td><td>0.20</td><td>-1.39</td><td>0.41</td></tr><tr><td>602</td><td>-1.11</td><td>0.31</td><td>-1.09</td><td>0.27</td><td>-0.98</td><td>0.32</td><td>-1.13</td><td>0.26</td><td>-1.85</td><td>0.32</td><td>-2.00</td><td>0.26</td></tr><tr><td>338</td><td>-0.05</td><td>0.26</td><td>0.29</td><td>0.37</td><td>0.07</td><td>0.28</td><td>0.55</td><td>0.23</td><td>-1.15</td><td>0.28</td><td>-0.67</td><td>0.23</td></tr><tr><td>339</td><td>0.02</td><td>0.46</td><td>0.06</td><td>0.20</td><td>0.22</td><td>0.28</td><td>0.16</td><td>0.19</td><td>-0.99</td><td>0.39</td><td>-1.06</td><td>0.30</td></tr><tr><td>347</td><td>-0.19</td><td>0.25</td><td>0.02</td><td>0.17</td><td>-0.13</td><td>0.13</td><td>0.41</td><td>0.00</td><td>-1.29</td><td>0.20</td><td>-0.75</td><td>0.07</td></tr><tr><td>394</td><td>0.58</td><td>0.31</td><td>0.61</td><td>0.23</td><td>0.64</td><td>0.26</td><td>0.71</td><td>0.13</td><td>-0.64</td><td>0.27</td><td>-0.58</td><td>0.14</td></tr><tr><td>395</td><td>0.47</td><td>0.30</td><td>0.53</td><td>0.21</td><td>0.53</td><td>0.27</td><td>0.63</td><td>0.13</td><td>-0.57</td><td>0.27</td><td>-0.46</td><td>0.13</td></tr><tr><td>396</td><td>0.66</td><td>0.22</td><td>-0.64</td><td>0.60</td><td>0.72</td><td>0.19</td><td>-0.91</td><td>0.42</td><td>-0.50</td><td>0.29</td><td>-2.13</td><td>0.52</td></tr><tr><td>Canyon Ø</td><td>-0.32</td><td>0.29</td><td>-0.32</td><td>0.27</td><td>-0.17</td><td>0.20</td><td>-0.21</td><td>0.26</td><td>-0.97</td><td>0.27</td><td>-1.01</td><td>0.33</td></tr><tr><td>Slope Ø</td><td>0.26</td><td>0.30</td><td></td><td></td><td>0.33</td><td>0.26</td><td></td><td></td><td>-0.67</td><td>0.32</td><td></td><td></td></tr></tbody></table>

relation with trophic conditions at each site, revealing higher values at more eutrophic sites.

The median living depth of the shallow infaunal *U. mediterranea* (MLD<sub>Umed</sub>) is used here to describe its micro-habitat and generally increases at sites with deep main re-

dox boundaries, at least in the western Mediterranean Sea. The deepest MLD<sub>Umed</sub> are 2.13 and 2.25 cm in the southern Aegean Sea, while the shallowest depths of 0.27 and 0.38 cm are recorded in the central and northern Aegean Sea, respectively (Table 1). In the Gulf of Lion, the MLD<sub>Umed</sub> is between
<table><caption>**Table 3.** Linear regressions of ontogenetic trends of δ¹³C<sub>Umed</sub>. The measured number of stained and unstained tests as well as the significance values are added.</caption><thead><tr><th colspan="5">U. mediterranea stained</th></tr><tr><th>Site</th><th>n</th><th>Linear fit</th><th>R<sup>2</sup></th><th>p value</th></tr></thead><tbody><tr><td>537</td><td>24</td><td>Y = 0.001379 × X - 1.810017</td><td>0.67</td><td>1.1065 × 10<sup>-6</sup></td></tr><tr><td>540A</td><td>23</td><td>Y = 0.001007 × X - 1.770373</td><td>0.70</td><td>5.746 × 10<sup>-7</sup></td></tr><tr><td>540B</td><td>14</td><td>Y = 0.001257 × X - 1.7208851</td><td>0.54</td><td>0.0027</td></tr><tr><td>540C</td><td>46</td><td>Y = 0.000943 × X - 1.639236</td><td>0.55</td><td>3.769 × 10<sup>-6</sup></td></tr><tr><td>585</td><td>3</td><td>Y = -0.00224 × X + 1.222667</td><td>1.00</td><td>0.0082</td></tr><tr><td>592</td><td>2</td><td>Y = 0.00034 × X - 1.6535</td><td>1.00</td><td>n/a</td></tr><tr><td>595</td><td>10</td><td>Y = 0.002013 × X - 3.012139</td><td>0.75</td><td>0.0012</td></tr><tr><td>596</td><td>7</td><td>Y = 0.001822 × X - 2.490764</td><td>0.60</td><td>0.0401</td></tr><tr><td>599</td><td>10</td><td>Y = 0.001600 × X - 2.789560</td><td>0.49</td><td>0.0289</td></tr><tr><td>601</td><td>11</td><td>Y = 0.001322 × X - 2.497314</td><td>0.70</td><td>0.0013</td></tr><tr><td>602</td><td>15</td><td>Y = 0.001143 × X - 2.709099</td><td>0.41</td><td>0.0102</td></tr><tr><td>338</td><td>10</td><td>Y = 0.001498 × X - 2.265059</td><td>0.72</td><td>0.0020</td></tr><tr><td>339</td><td>12</td><td>Y = 0.001527 × X - 2.323114</td><td>0.48</td><td>0.0124</td></tr><tr><td>347</td><td>7</td><td>Y = 0.001126 × X - 2.119201</td><td>0.68</td><td>0.0232</td></tr><tr><td>394</td><td>19</td><td>Y = 0.000968 × X - 1.680654</td><td>0.27</td><td>0.0221</td></tr><tr><td>395</td><td>23</td><td>Y = 0.001509 × X - 2.135640</td><td>0.40</td><td>0.0012</td></tr><tr><td>396</td><td>20</td><td>Y = 0.000789 × X - 1.304866</td><td>0.39</td><td>0.0034</td></tr><tr><td>Canyon aug</td><td>7</td><td>Y = 0.000516 × X - 1.297794</td><td>0.45</td><td>0.1015</td></tr><tr><td>Canyon (Feb)</td><td>21</td><td>Y = 0.000701 × X - 1.634263</td><td>0.43</td><td>0.0012</td></tr><tr><td>Slope (Aug)</td><td>6</td><td>Y = 0.000671 × X - 1.207976</td><td>0.34</td><td>0.2244</td></tr><tr><td>Slope (Feb)</td><td>14</td><td>Y = 0.001223 × X - 1.518849</td><td>0.48</td><td>0.0060</td></tr></tbody><thead><tr><th colspan="5">U. mediterranea unstained</th></tr><tr><th>Site</th><th>n</th><th>Linear fit</th><th>R<sup>2</sup></th><th>p value</th></tr></thead><tbody><tr><td>537</td><td>7</td><td>Y = 0.000408 × X - 2.169793</td><td>0.33</td><td>0.1784</td></tr><tr><td>540B</td><td>16</td><td>Y = 0.001017 × X - 1.457536</td><td>0.80</td><td>2.4803 × 10<sup>-6</sup></td></tr><tr><td>540C</td><td>9</td><td>Y = 0.000938 × X - 1.343878</td><td>0.53</td><td>0.0270</td></tr><tr><td>585</td><td>4</td><td>Y = 0.001610 × X - 1.910093</td><td>0.85</td><td>0.0808</td></tr><tr><td>586</td><td>29</td><td>Y = 0.001555 × X - 2.035859</td><td>0.48</td><td>2.9156 × 10<sup>-5</sup></td></tr><tr><td>589</td><td>25</td><td>Y = 0.001612 × X - 1.917381</td><td>0.58</td><td>1.0482 × 10<sup>-5</sup></td></tr><tr><td>592</td><td>28</td><td>Y = 0.001001 × X - 1.826222</td><td>0.48</td><td>4.5201 × 10<sup>-5</sup></td></tr><tr><td>595</td><td>36</td><td>Y = 0.000841 × X - 1.740275</td><td>0.17</td><td>0.0130</td></tr><tr><td>596</td><td>37</td><td>Y = 0.001065 × X - 2.004262</td><td>0.30</td><td>0.0005</td></tr><tr><td>599</td><td>12</td><td>Y = 0.001031 × X - 2.201211</td><td>0.31</td><td>0.0600</td></tr><tr><td>601</td><td>21</td><td>Y = 0.000312 × X - 1.871316</td><td>0.04</td><td>0.3927</td></tr><tr><td>602</td><td>14</td><td>Y = -0.000427 × X - 1.697763</td><td>0.12</td><td>0.2159</td></tr><tr><td>338</td><td>10</td><td>Y = 0.001343 × X - 1.735480</td><td>0.90</td><td>3.0586 × 10<sup>-5</sup></td></tr><tr><td>339</td><td>9</td><td>Y = 0.000456 × X - 1.408000</td><td>0.31</td><td>0.1197</td></tr><tr><td>347</td><td>10</td><td>Y = 0.000615 × X - 1.400530</td><td>0.71</td><td>0.0023</td></tr><tr><td>394</td><td>22</td><td>Y = 0.000573 × X - 1.221329</td><td>0.32</td><td>0.0060</td></tr><tr><td>395</td><td>15</td><td>Y = 0.000544 × X - 1.301955</td><td>0.33</td><td>0.0256</td></tr><tr><td>396</td><td>17</td><td>Y = -0.001682 × X - 1.020989</td><td>0.50</td><td>0.0016</td></tr><tr><td>Canyon (Aug)</td><td>36</td><td>Y = 0.000584 × X - 1.469912</td><td>0.28</td><td>0.0009</td></tr></tbody></table>

0.43 and 0.49 cm in the axis of the Lacaze-Duthiers Canyon and around 1.22 cm on the open slope (Table 1, Fig. 4a). The depth of the sediment color change, which marks the shift in redox potential and thus oxygen penetration, ranges from 2.25 cm in the Gulf of Lion (Site 540A) to as much as 30 cm in the central Aegean Sea (Site 596) (Table 1, Fig. 4b).

The measured TOC contents of the surface sediment range from 0.41 % (Site 586, southern Aegean Sea) and 0.58 % (Site 537, Strait of Sicily) to a maximum of 0.82 % (Site 602, northern Aegean Sea) (Table 1, Fig. 4c). The Δδ¹³C<sub>Umed-Epi</sub> and the MLD<sub>Umed</sub> (Fig. 4a) as well as the main redox bound-
<figure><img src="image_6.png" /><figcaption>**Figure 4.** The δ¹³C difference between live *Uvigerina mediterranea* and epifaunal taxa (Δδ¹³C<sub>Umed-Epi</sub>) plotted against (a) MLD of *U. mediterranea*, (b) depth of redox boundary in the sediment, and (c) TOC content of the sediment. The MLD error bars for the canyon and slope sites in the Gulf of Lion reflect the seasonal MLD contrasts of *U. mediterranea* between February and August 1997 (Schmiedl et al., 2004).</figcaption></figure>

ary depth (Fig. 4b) show good correspondence, whereas the link to percentage of TOC is less distinct (Fig. 4c).

The estimated values for annual primary production (PP) range from 106 to 294 g C m⁻² a⁻¹. Application of the different algorithms of Antoine and Morel (1996) and Uitz et al. (2008) resulted in an average offset of 54 g C m⁻² a⁻¹, with PP values consistently higher when applying the algorithm of Antoine and Morel (1996). The highest PP values occur in the Alboran Sea (274–294 vs. 192–207 g C m⁻² a⁻¹ according to Uitz et al., 2008) and the northern Aegean Sea (196–237 and 139–164 g C m⁻² a⁻¹, respectively), while the lowest PP values occur in the southern and central Aegean Sea (151–161 and 106–116 g C m⁻² a⁻¹, respectively) (Table 1).

# 4 Discussion

## 4.1 Stable carbon isotope signal of epifaunal foraminifera in relation to surrounding water masses

The δ¹³C of *Cibicoides pachydermus*, *Cibicides lobatus*, and *Planulina ariminensis* seems to reflect the δ¹³C<sub>DIC</sub> of the ambient bottom water since these species prefer an epifaunal microhabitat (Lutze and Thiel, 1989; Kitazato, 1994; Schmiedl et al., 2000). Comparison with published water δ¹³C<sub>DIC</sub> measurements confirms that δ¹³C<sub>Epi</sub> values are a possible bottom water proxy for the Mediterranean Sea (Pierre, 1999; Schmiedl et al., 2004; Theodor et al., 2016). Further, our new data corroborate previous observations that ontogenetic effects in the δ¹³C<sub>Epi</sub> signal of these taxa are lacking (Corliss et al., 2002; Franco-Fraguas et al., 2011; Theodor et al., 2016) (Table S1).

Because of the lack of stained epifaunal tests at most sites, unstained tests were integrated into the analysis. For empty
tests, a shift to higher δ¹³Cₚₐᵢ values due to potential disso-
lution effects should be considered (Edgar et al., 2013). In
addition, reworked or allochthonous tests can bias the results
as documented for the δ¹³Cₚₐ₇χ of Site 396 in the Mal-
lorca Channel. At this site, fossil tests have been admixed
in the surface sediment as indicated by heavy δ¹⁸O values
of greater than 4.0‰ (Table S1). In the Alboran Sea (Sites
339 and 347), we measured interspecific epifaunal δ¹³C dif-
ferences of up to 1.4‰. This variability is a result of im-
plausibly low δ¹³Cₖₐβ values, probably due to a relocation
from shallower depths closer to the coast. These unrealistic
δ¹³Cₚₐ₇χ and δ¹³Cₖₐβ values were omitted for δ¹³Cₚₐᵢ es-
timation. In order to minimize these biases, a large number of
tests were measured, which was possible for C. pachydermus
and P. ariminensis, showing commonly 0.3–0.5‰ higher
δ¹³C values for the latter species (Table 2, Fig. 2a). Despite
the aforementioned uncertainties, data of C. lobatulus were
used to estimate δ¹³Cₚₐᵢ at the Mallorca Channel Sites 394
and 395, when no tests of other species were available for
analysis (Theodor et al., 2016). For proper δ¹³Cₚₐᵢ estima-
tion of Sites 394 and 395, the difference between δ¹³Cₚₐᵢ and
δ¹³Cₖₐβ (Δδ¹³Cₚₐᵢ-ₖₐβ = 0.30‰) at Site 396 was added to
the δ¹³Cₖₐβ values (Table 2, Fig. 2a).

The δ¹³C offset between *C. pachydermus* and *P. ariminensis* is not constant and appears to increase at sites with deep main redox boundaries. This suggests a connection with increasing organic matter availability and the varying offsets can be attributed to slight differences in their microhabitat (Table 2, Fig. 2a). While *P. ariminensis* is a strictly epifaunal species, living attached on surfaces or above the sediment (Lutze and Thiel, 1989), *C. pachydermus* commonly lives at or slightly below the sediment-water interface (Rathburn and Corliss, 1994; Schmiedl et al., 2000; Licari and Mackensen, 2005). A very shallow infaunal microhabitat of *C. pachydermus* is corroborated by slightly lower δ¹³C values relative to bottom water δ¹³C_DIC suggesting pore water influence (Schmiedl et al., 2004; Fontanier et al., 2006). In order to compensate for potential pore water effects in the δ¹³C signal of the epifaunal species, the highest δ¹³Cₚₐᵢ values, mostly of *P. ariminensis*, should be selected for further comparison with shallow infaunal δ¹³C_Umed signals. This strategy could not always be realized, either due to the lack of *P. ariminensis* (Sites 537, 601, canyon, and slope) or when lower δ¹³C values were recorded for *P. ariminensis* relative to *C. pachydermus* (Site 540C). In these cases, bottom water δ¹³C_DIC measurements (canyon, slope; data from Schmiedl et al., 2004), the addition of the Δδ¹³Cₚₐᵢ-ₖₐ₇χ value of the nearby Site 602 (for correction of Site 601) or the δ¹³Cₚₐ₇χ values (Sites 537, 540C) were used, accepting possible deviations of δ¹³Cₚₐᵢ from bottom water δ¹³C_DIC (Table 2).

The applied δ¹³Cₚₐᵢ values are related to different
Mediterranean water masses (Fig. 2b). The δ¹³Cₚₐᵢ values
of the Gulf of Lion and the Spanish continental slope off
Barcelona are around 1.0‰ matching the δ¹³C_DIC signa-
ture of upper western Mediterranean deep water (WMDW

(Pierre, 1999). Likewise, the slightly higher δ¹³Cₚₐᵢ values of
1.1‰ in the Strait of Sicily fall in the range of δ¹³C_DIC val-
ues of intermediate waters from the eastern Mediterranean
Sea and reflect the transitional setting of this area. In con-
trast, the δ¹³Cₚₐᵢ values of the Mallorca Channel and the
Alboran Sea are even higher than those recorded for the east-
ern Mediterranean Sea (Fig. 2b). This inconsistent isotope
pattern likely reflects a shift in deep-water formation in the
eastern Mediterranean during the 1990s, the so-called East-
ern Mediterranean Transient (EMT; Roether et al., 2007).
The EMT was accompanied by an enhanced deep-water for-
mation in the Aegean Sea and also fostered a complete re-
newal of WMDW during the mid-2000s (Schroeder et al.,
2006, 2008). Unfortunately, the imprint of WMDW change
on δ¹³C_DIC of the water mass was not documented, but it
should have affected the sites sampled after this transition,
i.e., during *Meteor* cruise M69/1 in 2006.

The broad range of recorded δ¹³Cₚₐᵢ values of 0.87 to 1.95‰ in the Aegean Sea reflects the strong small-scale oceanographic differences of this region, including presence of various small isolated basins (Figs. 1, 2b). The comparatively high δ¹³Cₚₐᵢ values of the shallower sites indicate intensified vertical convection at sites of subsurface water formation, which recently resumed after the stagnation phase of 1994–2000 (Androulidakis et al., 2012), although the main deep-water formation area is restricted to the Cretan Sea (Roether et al., 1996; Lascaratos et al., 1999). Reduced replenishment of bottom waters at greater depth of isolated basins (Zervakis et al., 2003; Velaoras and Lascaratos, 2005) is accompanied by relatively low δ¹³C_DIC and accordingly low δ¹³Cₚₐᵢ values in these environments.

## 4.2 Biological and environmental effects on the stable carbon isotope signal of *Uvigerina mediterranea*

Size-dependent changes in the δ¹³C signal of *Uvigerina mediterranea* are attributed to ontogenetic effects. Small tests are depleted in ¹³C, while larger tests are closer to δ¹³C_DIC of the ambient pore water (Fig. 3). Relatively low δ¹³C_Umed values of small tests suggest stronger metabolic fractionation in younger individuals (Schmiedl et al., 2004; McCorkle et al., 2008; Schumacher et al., 2010; Theodor et al., 2016). A linear ontogenetic increase of 0.11‰ 100 µm⁻¹ was observed at all sites of the western Mediterranean Sea, while a steeper slope of 0.16‰ 100 µm⁻¹ was recorded in the Aegean Sea (Fig. 3). In addition, the δ¹³C_Umed values of small individuals from the Aegean Sea were of order 1‰ lower compared to those from the western Mediterranean Sea.

Differences in ontogenetic δ¹³C slopes of the related
species *U. peregrina* have been attributed to its highly oppor-
tunistic response to regional contrasts in organic matter quan-
tity and quality, and seasonality of supply (Theodor et al.,
2016). Obviously, similar effects are also operational in on-
togenetic δ¹³C trends of *U. mediterranea*. In the Aegean Sea,
this species appears to respond to strong seasonal contrasts
in organic matter fluxes (Siokou-Frangou et al., 2002) result-
ing in particularly high metabolic activity and low δ¹³C<sub>Umed</sub>
values in young individuals. A steepening of the δ¹³C<sub>Umed</sub>
slopes from the north to the south Aegean Sea probably oc-
curs for the same reasons as *U. peregrina* in the western
Mediterranean Sea. Because of the higher number of mea-
sured tests, this shift of the slope angles is more obvious
in unstained than stained tests (Fig. 3). Although the num-
ber of sites was larger than in Theodor et al. (2016), a sim-
ilar trend in δ¹³C<sub>Umed</sub> is not recognizable for the western
Mediterranean Sea. This may express lower differences in
the seasonal food supply between the sites or the total higher
input of organic matter compared to the Aegean Sea.

The δ¹³C<sub>Umed</sub> of unstained individuals from 5 cm sedi-
ment depth in the western Mediterranean Sea and Strait of
Sicily are on average 0.1 to 0.2‰ lower than those of stained
specimens in the topmost centimeter. This adds to previous
observations of Theodor et al. (2016) suggesting the influ-
ence of the Suess effect (Keeling, 1979; Quay et al., 1992)
in living individuals while it is absent in some 100-year-old
specimens. The Suess effect reduces δ¹³C values in the at-
mosphere and oceans due to the anthropogenic release of
isotopically light CO₂ out of fossil fuels. A similar effect
was not seen in the Aegean Sea since live and dead individ-
uals were selected from the same sediment depth and thus
had only minor age differences (Table 2, Fig. 3). The only
exception is Site 595 in the central Aegean Sea, where the
deviation is even higher (0.5–0.7‰) when compared to the
western Mediterranean Sea. Since this signal is restricted to
only one site, it is probably due to relocation of fossil tests
by the effects of bioturbation or lateral sediment transport.

Under well-oxygenated conditions, the pore water
δ¹³C<sub>DIC</sub> gradient depends on the organic matter fluxes and
associated decomposition rates of organic matter in the sur-
face sediment (McCorkle and Emerson, 1988; McCorkle et
al., 1985, 1990; Holsten et al., 2004). Organic matter fluxes
also control the depth of the oxygenated layer (Rutgers van
der Loeff, 1990) and thus the microhabitat range of infau-
nal foraminifera (Corliss, 1985; Jorissen et al., 1995; Koho
et al., 2008; Koho and Pina-Ochoa, 2012). Subsurface wa-
ters in the Mediterranean Sea are well ventilated resulting
in bottom water oxygen concentrations above 4.1 mL L⁻¹ at
all sites in our study (MedAtlas, 1997). The δ¹³C signal of
*U. mediterranea* appears particularly suitable to monitor the
pore water δ¹³C<sub>DIC</sub> signal in the near-surface sediment be-
cause it seems to be less influenced by species-specific vital
effects (McConnaughy, 1989a, b) when compared to other
shallow infaunal taxa (for example, *U. peregrina*) (Schmiedl
et al., 2004; Theodor et al., 2016).

In this study, the deviation of δ¹³C<sub>Umed</sub> from bottom water
δ¹³C<sub>DIC</sub> (reflected as higher Δδ¹³C<sub>Umed-Epi</sub> values; Fig. 4)
suggests exponential relations with the MLD of *U. mediter-
ranea*, the depth of the oxygenated layer, and the TOC con-
tent of the surface sediment. At the more oligotrophic to
mesotrophic sites of the Mallorca Channel, the Gulf of Lion,

the Spanish slope off Barcelona, and the southern Aegean
Sea, relatively low Δδ¹³C<sub>Umed-Epi</sub> values correspond to a rel-
atively thick oxygenated layer and low TOC contents. The
rather deep position of the redox boundary, exceeding 10 cm
at some sites, enables *U. mediterranea* to inhabit a rela-
tively wide microhabitat range. In contrast, relatively high
Δδ¹³C<sub>Umed-Epi</sub> values at the more mesotrophic to eutrophic
sites of the Alboran Sea coincide with relatively thin oxy-
genated layers and higher TOC contents. Here, the micro-
habitat range of *U. mediterranea* is compressed because of
limited pore water oxygen (Fig. 4).

When comparing sites within the central and northern
Aegean Sea, the foraminiferal stable isotope difference and
the biogeochemical and ecological characteristics lack a con-
sistent relation (Fig. 4). In these areas, strongly negative
Δδ¹³C<sub>Umed-Epi</sub> values do not systematically correspond to
maximum TOC contents and the shallowest redox bound-
ary (Fig. 4). The reasons for this absence of a clear rela-
tion between Δδ¹³C<sub>Umed-Epi</sub> and environmental parameters
within this area cannot be unraveled with our data. It may
be related to the high variability in oceanographic and bio-
geochemical conditions of the bottom water in the isolated
basins that are characterized by focusing of organic-rich sed-
imentary material (Lykousis et al., 2002; Giresse et al., 2003;
Poulos, 2009) and/or temporarily intermittent replenishment
of deep waters on seasonal to decadal timescales (Zervakis
et al., 2003; Velaoras and Lascaratos, 2005; Androulidakis
et al., 2012). The first possibility can increase the supply of
refractory C<sub>org</sub>, recorded by higher TOC contents, and in-
fluence the foraminiferal microhabitat depths, but has minor
effects on the δ¹³C<sub>DIC</sub> pore water gradient. Latter possibil-
ity refers to local ventilation events, which exchange aged
bottom water with comparatively low δ¹³C<sub>DIC</sub> signature by
surface waters enhanced in ¹³C<sub>DIC</sub>. This may also push the
pore water gradient towards stronger differences, explaining
the more negative Δδ¹³C<sub>Umed-Epi</sub> values, compared to the re-
maining sites with similar conditions (Fig. 4).

4.3 Development of a stable carbon-isotope-based
transfer function for organic carbon fluxes

Our results suggest a close relationship between the δ¹³C
gradient in pore waters of the surface sediment (expressed
as Δδ¹³C<sub>Umed-Epi</sub>) and the OM fluxes to the sea floor, for
open-ocean settings of the western and central Mediterranean
Sea and the southern Aegean Sea (Fig. 5). Based on these
observations, we tested the potential for the development of
a δ¹³C-based transfer function for OM flux rates. In open-
ocean settings, the main food source of deep-sea environ-
ments is the exported OM from the surface layer, where
photosynthetic PP takes place (e.g., Boyd and Trull, 2007;
Bishop, 2009). The majority of produced particulate organic
carbon (POC) is recycled within the photic zone. In the open
Mediterranean Sea, around 4% of the POC is exported out of
the photic zone, which is lower than for other open oceans,
<figure><img src="image_7.png" /><figcaption>**Figure 5.** The δ¹³C difference between live and dead *Uvigerina mediterranea* and epifaunal taxa (Δδ¹³C<sub>Umed-Epi</sub>) against organic carbon flux rates (C<sub>org</sub> flux) calculated from primary productivity in surface waters after Betzer et al. (1984). As in Fig. 4, satellite-derived primary production values of Antoine and Morel (1996) (top) and Uitz et al. (2008) (bottom) were used.</figcaption></figure>

caused by a specific nutrient distribution in the Mediterranean Sea (Moutin and Raimbault, 2002; Gogou et al., 2014). The remineralization of organic matter is intensified, which leads to reduced fluxes to the sea floor.

During transfer from the surface ocean to the deep sea, the amount of exported OM decreases exponentially reflecting microbial decay (Suess, 1980; de la Rocha and Passow, 2007; Packard and Gomez, 2013). Various functions have been developed for the estimation of OM fluxes during sinking of particles through the water column integrating numerous observational data (Suess, 1980; Betzer et al., 1984; Martin et al., 1987; Antia, et al., 2001). The different functions reveal a high variability for the active surface layer, while the results for deeper parts of the water column are within a comparable range (Felix, 2014). In our study (Table 1, Fig. 5), we applied the function of Betzer et al. (1984) for calculating the depth-dependent C<sub>org</sub> fluxes at the different Mediterranean sites us-

ing satellite-derived PP data (Antoine and Morel, 1996; Uitz et al., 2008).

A comparison with direct PP and export flux measurements of sediment trap studies revealed ambiguous results. The PP values calculated after Antoine and Morel (1996) are in a comparable range to PP measurements in the western Mediterranean (Moutin and Raimbault, 2002; Sanchez-Vidal et al., 2004, 2005; Zúñiga et al., 2007, 2008). However, the estimated export fluxes are too high in these areas compared to direct measurements of the referred studies, probably due to the aforementioned high remineralization rate in the Mediterranean Sea. However, the discrepancy in export fluxes is partly compensated by the application of the 21–30 % lower PP values calculated after Uitz et al. (2008). For the Aegean Sea, in contrast, distinctively higher measured PP values have been reported than were estimated (Siokou-Frangou et al., 2002). For the Gulf of Lion, measured OM
export fluxes exceed the predicted values (Heussner et al., 2006), which can be explained by the additional lateral input of organic carbon channeled within the local canyon systems (Schmiedl et al., 2000). In order to compensate for these possible additional Corg fluxes in marginal basin areas, the application of the function of Antoine and Morel (1996) is more promising; hence, a potential overestimation of Corg fluxes in open-ocean areas has to be considered.

For both approaches of PP calculation (Antoine and Morel, 1996; Uitz et al., 2008), the relation between the estimated Corg fluxes and the Δδ¹³C<sub>Umed-Epi</sub> exhibits a complex pattern and, at first instance, lacks a simple and statistically significant correlation (Fig. 5). Particularly, strongly negative Δδ¹³C<sub>Umed-Epi</sub> in the central and northern Aegean Sea suggest high Corg fluxes, which however are not reflected in the estimated PP-based values. The eventual underestimation of Corg fluxes in these more marginal areas is likely caused by additional lateral OM input and the focusing of organic matter in isolated small basins. In fact, the northern and central Aegean Sea experiences high OM input from terrestrial sources through outflow of north Aegean rivers and the Black Sea (Aksu et al., 1999; Tsiaras et al., 2012). In contrast, the measured main redox boundary depth and the TOC contents do not indicate a higher supply in organic matter. However, sediment trap data from the northern Aegean Sea (Lykousis et al., 2002) reveal Corg fluxes of 35–81 g C m⁻² a⁻¹, which are 3–10 times higher than estimated values solely based on PP-based vertical fluxes. Although the high measured values can be partly attributed to the short sampling interval of 2 months in late spring and thus to elevated vertical fluxes during the spring bloom, elevated year-round lateral Corg fluxes can be expected, but of a clearly lower dimension. The measured ratio of primary to reworked OM in the sediment at this site is around 60–70% (Lykousis et al., 2002; Poulos, 2009), which leaves the PP as the main source of the Corg fluxes to the deep sea. Similar results have been derived for canyon systems of the Gulf of Lion where OM resuspension, shelf-to-slope cascading, and channeling results in significantly higher observed than PP-derived estimated Corg fluxes (Heussner et al., 2006; Pusceddu et al., 2010, Pasqual et al., 2010). Even in open slope settings, resuspended OM can significantly contribute to the total Corg flux (McCave et al., 2001; Tesi et al., 2010; Stabholz et al., 2013).

Despite these biases, it appears reasonable to develop a Corg flux transfer function at least for the more open marine settings of the western and central Mediterranean Sea and the southern Aegean Sea (Fig. 6). Here, vertical sinking of PP-derived OM appears to be the main source for Corg fluxes (Pusceddu et al., 2010) explaining the good correlation with the Δδ¹³C<sub>Umed-Epi</sub> values (Fig. 5). Elevated Corg fluxes of the upwelling-affected Alboran Sea (Hernandez-Almeida et al., 2011) are reflected in rather negative Δδ¹³C<sub>Umed-Epi</sub> values while the observed δ¹³C differences in the more oligotrophic regions of the Mallorca Channel, the Spanish slope off Barcelona, the Strait of Sicily, and the southern Aegean

<figure><img src="image_8.png" /><figcaption>**Figure 6.** Correlation of the δ¹³C difference between live *Uvigerina mediterranea* and epifaunal taxa (Δδ¹³C<sub>Umed-Epi</sub>) and organic carbon flux rate (Corg flux) calculated according to Antoine and Morel (1996) and Betzer et al. (1984). Transparent data from the central and northern Aegean Sea and the Gulf of Lion have been removed from the function since PP-based Corg flux values are likely underestimated because of the additional influence of lateral organic matter fluxes on the δ¹³C<sub>Umed</sub> values in these areas.</figcaption></figure>

Sea are lower. So, omitting the data from the northern and central Aegean Sea, and considering sediment trap data from the Gulf of Lion (Heussner et al., 2006) the derived function can be expressed as

$$C_{\text{org}} \text{ flux} = -15.99 \times \Delta\delta^{13}\text{C}_{\text{Umed-Epi}} + 0.34, \quad (1)$$

with a coefficient of determination (R²) of 0.63 and a significance (p) of 0.0021 (Fig. 6). The estimated Corg fluxes can be used to recalculate marine PP, but should be handled carefully due to the highly possible overestimation caused by lateral advection. Especially in more marginal areas, this bias can lead to unreliable recalculated PP values.

The application of this function to unstained *U. mediterranea* tests creates a higher range of uncertainty. The main reason for this inconsistency seems to be the relocation of fossil tests at particular sites, leading to significant contrasts between δ¹³C<sub>Umed</sub> values of stained and unstained tests. For empty *U. mediterranea* tests, marked negative δ¹³C<sub>Umed</sub> outliers appear at Sites 537 and 396, which has already been mentioned in Theodor et al. (2016) for the latter site. In the Alboran Sea (Sites 338 and 347) on the other hand, δ¹³C<sub>Umed</sub> values of unstained tests are about 0.50‰ higher than those of stained tests. Less distinct δ¹³C<sub>Umed</sub> differences between
autochthonous and allochthonous tests may not be detected
so easily. These potential uncertainties have to be consid-
ered in the application of the transfer function to sediment
cores, particularly to down-core records from sites influenced
by strong lateral transport such as canyon environments or
the northern and central Aegean Sea. Likewise, the applica-
tion of the transfer function to areas outside of the Mediter-
ranean Sea may be biased by contrasting remineralization
rates due to the specific oceanographic conditions, especially
the higher temperatures in the Mediterranean Sea. Further
refinement of this function will require an interdisciplinary
effort including a larger number of direct Corg flux measure-
ments in sediment trap deployments, which can be directly
related to the obtained foraminiferal δ13C signals.

5 Conclusions

The δ¹³C signal of deep-sea benthic foraminifera from dif-
ferent areas of the western, central, and eastern Mediter-
ranean Sea reflects an integration of various environmental
and biological signals. The application of epifaunal benthic
foraminifera as an unbiased proxy for the δ¹³C_DIC of the
surrounding water mass is ambiguous due to possible al-
lochthonous tests, but also due to a slight species-specific
difference in the microhabitat that can result in significant
δ¹³C_Epi shifts. The δ¹³C signal of the strictly epifaunal Plan-
ulina ariminensis should be preferred, in contrast to the δ¹³C
signal of the less strictly epifaunal Cibicidoides pachyder-
mus, which appears to be influenced by pore water DIC and
its δ¹³C value.

The δ¹³C signal of epifaunal taxa lacks ontogenetic ef-
fects supporting results from previous studies (Dunbar and
Wefer, 1984; Corliss et al., 2002; Theodor et al., 2016). Sig-
nificant ontogenetic effects were recorded in the δ¹³C signal
of *Uvigerina mediterranea*. While the ontogenetic increase
of δ¹³C_Umed is more or less comparable (0.11 ± 0.03 ‰
100 µm⁻¹) in the western Mediterranean and the Strait of
Sicily, a stronger increase and even a regional south-north
trend is documented for the Aegean Sea (0.16 ± 0.04 ‰
100 µm⁻¹). In general, the δ¹³C values of *U. mediterranea*
from the Aegean Sea are more negative when compared to
those from the western and central Mediterranean Sea. This
regional contrast cannot be reconciled with different vital and
pore water effects but instead seems to be caused by en-
hanced residence times of bottom waters in the partly iso-
lated small basins within the Aegean Sea. In cases of well-
oxygenated conditions, the δ¹³C_Umed signal, compared to
bottom water, is mainly controlled by regional trophic con-
trasts and related remineralisation rates. The Δδ¹³C_Umed-Epi
are clearly related to the median microhabitat depth, the
depth of the redox boundary (indicating the extent of the
oxygenated layer), and (to a lower extent) to the TOC of
the surface sediment. Based on satellite-derived primary pro-
duction estimates Corg fluxes were calculated and related to
the recorded Δδ¹³C_Umed-Epi values. Comparison with sed-
iment trap data reveals underestimation of satellite-derived
Corg fluxes for the marginal areas of the central and northern
Aegean Sea and the canyon systems of the Gulf of Lion. In
these ecosystems, additional lateral transport of resuspended
and terrestrial OM contributes substantially to Corg fluxes.
Considering these biases, a first estimation for Corg fluxes in
open-ocean settings of the Mediterranean Sea could be es-
tablished.

6 Data availability

The main data source (stable isotope data) of this work is
within the appendix.
Appendix A

List of benthic foraminiferal taxa used in this study

*Cibicides lobatulus* (Walker and Jakob)

*Cibicidoides pachydermus* (Rzehak)

*Planulina ariminensis* (d'Orbigny)

*Uvigerina mediterranea* (Hofker)

*Nautilus lobatulus* (Walker and Jacob, 1798, p. 642, pl. 14, Fig. 36)

*Truncatulina pachyderma* (Rzehak, 1886, p. 87, pl. 1, Fig. 5)

*Planulina ariminensis* (d'Orbigny, 1826, p. 280, pl. 14, Figs. 1–3)

*Uvigerina mediterranea* (Hofker, 1932, pp. 118–121, Fig. 32)
The Supplement related to this article is available online
at doi:10.5194 bg-13-6385-2016-supplement.

*Acknowledgements.* We would like to thank K.-C. Emeis and two anonymous referees for their helpful remarks and suggestions. We thank the ship crews and scientists of the R/V *Meteor* for good collaboration during cruises M40/4, M51/3, and M69/1. We give thanks to Valerie Menke for foraminifera test size measurements and Mareike Paul for selection of epifaunal specimens. We thank David Antoine for suggestions on the GlobColour data set and Jürgen Möbius for support during processing of the TOC samples. Lisa Schönborn and Günther Meyer are thanked for technical support during stable isotope measurements. This study was supported by the Deutsche Forschungsgemeinschaft, grants SCHM1180/16 and MA1942/11.

Edited by: A. Shemesh

Reviewed by: three anonymous referees

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