Environmental and climatic changes in the central Mediterranean Sea (Siculo–Tunisian Strait) during the last 30ka based on dinoflagellate cyst and planktonic foraminifera assemblages

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Environmental and climatic changes in the central Mediterranean Sea(Siculo–Tunisian Strait) during the last 30 ka based on dinoflagellatecyst and planktonic foraminifera assemblages

Imene Rouis-Zargouni a,b,⁎, Jean-Louis Turon b, Laurent Londeix b, Latifa Essallami a,Néjib Kallel a, Marie-Alexandrine Sicre c

a Faculté des Sciences de Sfax, Laboratoire GEOGLOB, Route de Soukra, BP 802, 3028 Sfax, Tunisiab Université Bordeaux 1, UMR 5805 EPOC, Avenue des Facultés, 33405 Talence cedex, Francec Laboratoire des Sciences du climat et de l'environnement; IPS CNRS/INSU CEA/UVSQ, avenue de la Terrasse, 91198 Gif-sur-Yvette cedex, France

a b s t r a c ta r t i c l e i n f o

Article history:Received 3 February 2009Received in revised form 13 October 2009Accepted 19 October 2009Available online 27 October 2009

Keywords:DinocystsPlanktonic foraminiferaMediterranean regionHeinrich eventsHolocenePresapropelic condition

A high resolution micropalaeontological study of the core MD 04-2797 CQ recovered in the Sicilian–TunisianStrait provides insights into the paleoclimatic history of the Mediterranean Sea at the transition between thewestern and eastern basin over the last 30 ka. Using the analysis of dinoflagellate cyst and planktonicforaminiferal assemblages, we reconstruct the paleoenvironmental changes that took place in this region.High abundances of cold temperate dinocyst species (Nematosphaeropsis labyrinthus, Spiniferites elongatus,Bitectatodinium tepikiense) and the polar planktonic foraminifera Neogloboquadrina pachyderma (left coiling)reveal three major cooling events synchronous with North Atlantic Henrich events 1 and 2 (H1 and H2) andthe European and North Atlantic Younger Dryas event. During the Holocene, the presence of warm dinocystspecies (Spiniferites mirabilis and Impagidinium aculeatum) and planktonic foraminifera (Globorotalia inflataand Globigerinoides ruber), reflects a significant increase of sea surface temperatures in the westernMediterranean basin, but a full warming was not recorded until 1500 years after the onset of the Holocene.Moreover, our results show that the Holocene was interrupted by at least four brief cooling events at ~9.2 ka,~8 ka, ~7 ka and ~2.2 ka cal. BP, which may be correlated to climatic events recorded in Greenland ice coresand in the Atlantic Ocean.

© 2009 Elsevier B.V. All rights reserved.

1. Introduction

The Mediterranean Sea is a semi-enclosed basin, which isconnected to the Atlantic Ocean via the Strait of Gibraltar. This basinhas an anti-estuarine circulation, whichmakes it possible to relate theprocesses of dense water formation and thermohaline circulation(Pinardi and Masetti, 2000). This also enables oceanographic andpaleoceanographic research to be carried out on past climatic changes(Turon and Londeix, 1988; Combourieu-Nebout et al., 1998; Rohlinget al., 1998; Asioli et al., 1999; Paterne et al., 1999; Cacho et al., 1999,2001; Asioli et al., 2001; Sbaffi et al., 2001; Combourieu-Nebout et al.,2002; Rohling et al., 2002; Sprovieri et al., 2003 ; Perez-Folgado et al.,2003, 2004; Sierro et al., 2005; Frigola et al., 2008). In this study,we focus on the Sicilian–Tunisian Strait because it is one of the least

understood areas of the Mediterranean despite being the mainconnection between the deep western and eastern basins and ahigh sedimentation rate for the Holocene and generally throughoutduring the Quaternary (DiStefano, 1998).

Often late Quaternary palaeoclimatic and environmental recon-structions are based on biochemical analyses of organic compoundsand a range of palaeontological proxies including such as planktonicforaminifera, ostracods, radiolarians, coccoliths, pollen and dinofla-gellate cysts. In particular over the last 30years, numerous papershave more demonstrated the value of dinocysts in reconstructingQuaternary paleoenvironments (e.g. Turon, 1981a,b, Turon, 1978;Turon and Londeix, 1988; de Vernal et al., 1992, 1994; Harland andHowe, 1995; Zonneveld, 1996; Combourieu-Nebout et al., 1998;Rochon et al., 1998; Sangiorgi, 2001; Sangiorgi et al., 2002; Eynaudet al., 2004; de Vernal et al., 2005; de Vernal and Hillaire-Marcel,2006). Furthermore, organic-walled dinocysts are resistant to disso-lution and can complement or replace information lost by thedissolution of calcareous and siliceous microfossils.

We present a multidisciplinary approach to the analysis of coreMD04-2797 CQ collected in the Siculo–Tunisian Strait to (1) obtaindinocyst and planktonic foraminifera temporal distribution during the

Palaeogeography, Palaeoclimatology, Palaeoecology 285 (2010) 17–29

⁎ Corresponding author. Université Bordeaux 1, UMR 5805 EPOC, Avenue desFacultés, 33405 Talence cedex, France. Tel.: +33 540 00 8858; fax: +33 556 84 0848.

E-mail addresses: [email protected] (I. Rouis-Zargouni),[email protected] (J.-L. Turon), [email protected] (L. Londeix),[email protected] (L. Essallami), [email protected] (N. Kallel),[email protected] (M.-A. Sicre).

0031-0182/$ – see front matter © 2009 Elsevier B.V. All rights reserved.doi:10.1016/j.palaeo.2009.10.015

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last two isotopic marine stages; (2) attempt a SST reconstruction basedon dinocysts and compare the dinocyst records with those previouslyobtained from planktonic foraminifera and alkenones SST (Essallamiet al., 2007) and (3) provide paleoenvironmental information for theinterpretation of climatic change and improve the dinocyst climato-stratigraphy in the central Mediterranean for the last 30,000 years.

2. Oceanographic setting

The Sicilian–Tunisian Strait is the principal area of exchange ofwaterbetween the Western and Eastern Mediterranean Sea. Through thisStrait, two-thirds of the less saline Atlantic waters, which enter theMediterranean Sea via the Gibraltar Strait as Modified Atlantic Water(MAW), flow to the east. MAW follows a southern path along theTunisian coast and occupies awater column thickness between 150 and200 m in the central part of the Strait and covers the entire photic zone(Astraldi et al., 1999, 2002; Béranger et al., 2004; Gasparini et al., 2005).The more saline Levantine Waters (LIW), flow westward from theeastern basin, via the relatively shallow Siculo–Tunisian channel, wherethey occupy the deep and intermediate parts of the water column(Morell, 1971). The northern path of the LIW can be recognized at thesurface along the southern Sicilian coast (Morell, 1971), where surfacewater temperatures are low throughout the year. The deep LIW mustrise from the deep eastern basin in order to flowalong the bottomof the

Sicily channel,which is only about350 mdeep in its shallowest part. Themain connection between the deepwestern and easternMediterraneanbasins occurs between Cape Bon and Mazzara del Vallo (Fig. 1).

3. Materials and methods

CoreMD04-2797 CQ(36°57N; 11°40E, 771 mwater-depth) (Fig. 1)was collected in the central part of the Sicilian–Tunisian channel duringthe IMAGES cruise in 2004. This 10.3 m Calypso Square (CASQ) core ismainly composed of homogenous greenish grey clays. The highsedimentation rate enables the reconstruction of the paleoceanographyat the transitionbetween the eastern andwesternMediterraneanbasinsduring the late Quaternary. Sampleswere taken every 10 cm in the core.

3.1. Planktonic foraminiferal stable isotopes and POC analysis

A detailed oxygen isotope record, expressed in ‰ versus VPDB(Vienna Pee Dee Belemnite Standard) was taken every 2 cm andobtained on 6 to 20 shells of the planktonic foraminifer Globigerinabulloïdes in the 250–315 µm size range. VPDB is defined with respectto NBS 19 calcite standard (Coplen, 1988). Analyses were performedat Laboratoire des Sciences du Climat et de l'Environnement (LSCE) onFinnigan Delta+ and MAT 251 mass spectrometers. Linearity wascorrected following Ostermann and Curry (2000). The mean external

Fig. 1. Location of the core MD04-2797 CQ (36°57N; 11°40E, 771 m water-depth), Sicilian–Tunisian Strait.

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reproducibility (1σ) of carbonate standards is ±0.05‰, measuredNBS18 δ18O is −23.2±0.2‰ VPDB.

Particulate organic carbon (POC) was measured directly from thedry, powdered and homogenised material, which was sampled every20 cm and at a higher resolution in some sections of particularinterest, using a carbon/sulphur analyser (LECO, CS-125) according toCauwet et al. (1990). Quality was checked by measuring certifiedreference materials (e.g. LECO 501-503) and intercalibrations (e.g.King et al., 1998). Representative subsamples (e.g. 30 mg of dry,powdered and homogenised material) were digested in closed Teflonreactors (savillex) on a heating plate (2 h at 110 °C) using 750 µl HCl(12 M) s.p and 2 ml HF (26 M) s.p and 250 µl HNO3 (14 M) s.p. Afterevaporation to dryness, the residues were completely re-dissolved in150 µl HNO3 (14 M) on a heating plate and after cooling brought to10 ml in volumetric flasks using Milli-Q water (Schäfer et al., 2002).

3.2. Palynological processing and analysis

Dinocyst analysis was performed on the fraction b150 µm. Thepalynological processing followed the procedure described by deVernalet al. (1996) and Rochon et al. (1999), slightly modified at the UMR-EPOC laboratory (www.epoc.u-bordeaux.fr/equipe thématique paléo/Outils). After chemical treatments (cold 10, 25 and50%HCl, cold 40 then70%HF shakenduring24 h), the sampleswere sieved through single use10 µm nylon mesh screens. Acetolysis was not used to avoid thedestruction of polykrikacean and protoperidiniacean cysts (Turon,1984; Marret, 1993). The final residue was mounted in glycerine jellycoloured with fuschin. Dinocysts were counted using a Zeiss Axioscopelight microscope at 400×. An average of 100 to 300 dinoflagellate cystswere identified and counted from each sample. References for dinocystidentification are based on studies from Turon (1984) and de Vernal

Fig. 2. Age model of core MD04-2797 CQ: dotted line represents an interpolation of the five radiocarbon converted into calendar age and control points extrapolated from peak-to-peak correlation of MD04-2797CQ G. bulloides δ18O record with the δ18O record of the dated core MD95-2043 (Cacho et al., 1999).

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et al. (1992). The dinocyst taxonomy is generally in agreementwith thatcited in Fensome and MacRae (1998). Dinocyst concentrations werecalculated using themarker grainmethod (de Vernal et al., 1996) basedon aliquot number of Lycopodium spores.

3.3. Sea surface temperature

The quantitative reconstruction of sea surface temperatures is basedon the analysis of organic-walled dinoflagellate cyst assemblages. Thereference database for dinocysts is composed of the palynological

analysis of 1171 (http:/www.images-pages.org/datamanagement.html) surface samples from Pacific, mid- to high-latitude North Atlanticand Arctic Oceans and adjacent seas. In this study, we use only 401references sites (Fig. 2) (Rouis-Zargouni, in preparation), which wereselectedwithin the latitudinal range of about 25–70°N and longitudinalrange of 15°E–30°W, to avoid over representing Arctic and Pacific sites,in accordance to a previous study of the Mediterranean Sea by Hayeset al. (2005). The North Atlantic core tops are included to provideanalogues for glacial Mediterranean Sea assemblages, which containspecies that do not occur in the Mediterranean today.

To reconstruct sea surface paleotemperatures, we apply theModernAnalogue Technique (MAT) using the software PaleoAnalogs describedin Therón et al. (2004). Estimation is calculated from a set of the best 3modern analogues. The accuracy of the approach is assessed by thecoefficient of dissimilarity or squared chord distance, which measuredthedegree of similarity between each fossil assemblage and the selected3 best analogues. For the fossil samples for which good or acceptablemodern analogues are available in the reference database, dissimilaritycoefficients are generally lower than 0.25 or 0.3 (Prell, 1985). Bycontrast, dissimilarity coefficients higher than 0.3 generally indicatethat there are no close modern analogues in the database and theconstructed SSTs at these levels should be considered with caution.

The Modern Analogue Technique (MAT) was applied on theplanktonic foraminiferal fossil assemblages using a database ofn=252 (Kallel et al., 1997a) to derive SSTs (Essallami et al., 2007).

4. Results

4.1. Stratigraphy and age model

Thirteen radiocarbon ages were determined by accelerator massspectrometry (AMS) by ARTEMIS. A minimum of 8 mg of monospecificforaminifera was sampled for each radiocarbon dating. The precisespatial and temporal patternsof change inpaleoreservoir agearenot fullyunderstood, and therefore we have applied a uniform reservoir cor-rection of−400 years (Bard, 1988; Siani et al., 2001). 14C ages have beenconverted to Calendar years using the calib5.01 program (Stuiver et al.,1998). The age model developed for MD04-2797 CQ (Table 1; Fig. 3) is

Table 1Age model pointers for MD04-2797CQ.

Depth(cm)

Age 14C(yr BP)

Error(yr BP)

Calibrated age(yr BP)

AMS lab Species

0 1105 20 662 ARTEMIS G. inflata 14C controlpoint

40 2508 Control point120 4770 Control point140 5990 Control point160 6764 Control point200 7465 30 7929 ARTEMIS G. ruber 14C control

point330 8965 30 9380 ARTEMIS G. ruber 14C control

point370 10,715 Control point470 12,605 40 14,058 ARTEMIS G. inflata 14C control

point490 15,498 Control point511 13,800 100 15,909 ARTEMIS G. bulloides 14C control

point560 17,080 Control point610 15,590 50 18,617 ARTEMIS G. bulloides 14C control

point700 21,660 Control point910 25,530 Control point960 27,180 Control point970 27,960 Control point1010 28,640 Control point1030 29,110 Control point

Control points (cf. Section 4.1 for explanation).All calibrated ages are calculated using CALIB 501 program.

Fig. 3. Map illustrating the core localities in the North Atlantic portion and the Mediterranean Sea of the calibration “n=401” reference dinocyst database used in this study (takenfrom database presented in http:/www.images-pages.org/datamanagement.html).



based on six radiocarbon ages and correlation between the δ18O curvesfor Globigerina bulloides of our core and the dated core MD95-2043(36°08N;02°37W, 1841 mwater) (Cacho et al., 1999). Mean uncertaintyon ages is estimated to be about 500 years.

4.2. Particulate organic carbon (POC) analysis

The POC curve shows low values ~0.4–0.5% at the base and at thetop of the core (Fig. 4). Between 10.5 and 7.5 ka cal. BP, a slight butsignificant increase of POC values to 0.8% was recorded. This suggestsan occurrence of presapropelic conditions in the western Mediterra-nean Sea in this interval, which is linked and contemporaneouswith the installation of sapropel S1 (9–6 ka cal. BP) in the easternMediterranean basin (Rossignol-Strick, 1985; Kallel et al., 1997a,b;Rossignol-Strick and Paterne, 1999; Ariztegui et al., 2000; Rohlinget al., 2002). Inside this interval, at 8 ka cal. BP, a brief drop of POCvalues was recorded.

4.3. Micropalaeontological results

4.3.1. Dinocyst species associationsDinocyst concentrations are generally more than 1200 cysts/cm3

with a maximum recorded during the Younger Dryas (5–8×103cysts/cm3) and at ~8.4 ka cal. BP (6×103cysts/cm3) (Fig. 4). Dinocystassemblages are based on the identification of 25 species but onlythe most representative are shown in Fig. 5.

4.3.1.1. 30 to 15.5 ka cal. BP (from 1030 to 500 cm). This intervalcorresponds to the end of isotopic stage 3, Heinrich event H2, LastGlacial Maximum (23–18 ka cal. BP) and Heinrich event H1 (Table 2).It is characterised by the dominance of Operculodinium centrocarpumaccompanied by cold microfloral assemblages. The highest percen-tages of the cold taxa have been recorded during Heinrich events:Nematosphaeropsis labyrinthus (10–20%), Bitectatodinium tepikiense(4–8%) and Spiniferites elongatus (5–18%) (Fig. 4).

Nematosphaeropsis labyrinthus has been positively correlated withopen cold water masses and high (winter) nutrient availability(Harland, 1983; Turon and Londeix, 1988; Devilliers and de Vernal,2000). Its modern maximum abundance is recorded on the south-

western Iceland margin where the Irminger Current dominates(Marret et al., 2004). Bitectatodinium tepikiense is mainly distributedfrom temperate to sub-Arctic environments of the North Atlantic,with maximum representation south of the Gulf of St. Laurent(Rochon et al., 1999; de Vernal et al., 2001). This species appears totolerate large seasonal variations in temperature, with very coldwinters (as cold as 1 °C) and mean summer temperatures over 15 °C.B. tepikiense has been observed in Quaternary sediments off thePortugal margin (Eynaud et al., 2000; Turon et al., 2003), where it hasbeen linked to the southward penetration of cold sub-polar watersduring Heinrich Events. In contrast, there is a presence, albeit in lowpercentages (b10%), of Impagidinium aculeatum during H1 and H2.

During the Last Glacial Maximum, we note the occurrence ofthe tropical species Spiniferites delicatus (12–24%) and Spiniferitesmembranaceus (7–13%); they don't reach 5 and 17% respectively ofthe dinocyst assemblages in modern western Mediterranean sedi-ments (Mangin. 2002). Lingulodinium machaerophorum (6–16%),which prefers neritic domains (Wall et al., 1977; Harland, 1983;Turon, 1984) and summer SST more than 10 °C, is also present duringthe Last Glacial Maximum.

4.3.1.2. 15.5 to 13 ka cal. BP (500–430 cm). This interval, the Bölling/Alleröd, is characterised by the dominance of the warm-temperatespecies Spiniferites mirabilis (10–20%) (Turon, 1981a,b; Harland,1983; Turon, 1984; Marret and Turon, 1994; Rochon et al., 1999).Morzadek-Kerfourn (1998a,b) demonstrated its southward tropicalextension (as far as 10°N). The warm species Impagidinium aculeatum(5%), Impagidinium paradoxum (5%) and Spiniferites delicatus (10%)occurred also in this interval.

4.3.1.3. 13–11.5 ka cal. BP (430–390 cm). The Younger Dryas interval isclearly dominated by Nematosphaeropsis labyrinthus (35%) associatedwith an increase of Spiniferites elongatus and a scarcity of such warmwater species as Spiniferites mirabilis and the warm Impagidiniumgroup.

4.3.1.4. From 11.5 ka cal. BP (390 cm to the top). The Holocene intervalis marked by the development of warm dinocyst species: Impagidi-nium paradoxum (5–10%), Impagidinium aculeatum (10–25%) and

Fig. 4. Particulate organic carbon (POC) % and oxygen isotope (δ18O) obtained from Globigerina bulloides of the core MD04-2797 CQ. The brown band indicates the presapropeliccondition period and the other grey bands indicate Heinrich events (H2 and H1) and the Younger Dryas event.



Fig. 5. Relative abundance of main species of dinocysts in core MD04-2797CQ. Grey bands indicate Heinrich events (H2 and H1) and the Younger Dryas event.

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Spiniferites mirabilis (15–50%). We can distinguish also the followingperiods.

The early Holocene (11.5–6 ka cal. BP) characterised by a progressiveincrease of the warm species Impagidinium aculeatum reaching amaximum of 25% at 6 ka cal. BP. Spiniferites mirabilis increasesprogressively and alsoNematosphaeropsis labyrinthuswas still presentthroughout but in low abundance.The Upper Holocene (the last 6 ka cal. BP): the percentage of Impagi-dinium aculeatum (5–12%) decreases and Spiniferites mirabilis reachesits optimum development (40–50%) and becomes the dominantspecies. The highest percentage abundance of S. mirabilis (25–45%)occurs at winter SSTs of between 14 and 15 °C and at summer SSTsbetween 21 and 25 °C in the Atlantic Ocean (http:/www.images-pages.org/datamanagement.html).

4.3.2. Planktonic foraminiferaThe planktonic foraminiferal assemblages are based on the identifi-

cation of 12 species and only the significant species are shown in Fig. 6.

4.3.2.1. 30 to 15.7 ka cal. BP. This interval shows a dominance of coldwater species with Globigerina quinqueloba (10–30%), Neogloboqua-drina pachyderma (right coiling) (20–50%), Globorotalia scitula (5–10%) and N. pachyderma (left coiling) (0–3%) and a scarcity of warmwater species. However, Globigerinoides ruber increases slightly at theend of H2 and during the LGM. N. pachyderma (left coiling) increasesduring the Heinrich Events with percentages higher in H1 (1.5–3%)than in H2 (0.5–2%).

The Bölling/Alleröd is marked essentially by the appearance ofGloborotalia inflata (30%) and Orbulina universa (4%) associatedwith the disappearance of Globorotalia scitula reflecting a rise of seasurface temperature. The subtropical species Globigerinoides ruber(pink and white) shows an increase of up to 20%. Cold species such asGlobigerina quinqueloba and Neogloboquadrina pachyderma (r.c.)decrease considerably.

4.3.2.2. The Younger Dryas. The significant cooling of this period isindicated by the dominance of the sub-polar species Neogloboquadrinapachyderma (right coiling) and Globigerina quinqueloba, and a consider-able reduction in warm water species Globigerinoides ruber and Globoro-talia inflata. At the transition between this event and the Holocene, thereis an abrupt and brief increase of G. inflata of 5 to 40%.

4.3.2.3. The Holocene. At 10 ka cal. BP, a significant increase of Globi-gerinoides ruber (60%) occurs and this remains at ~30% during themid-upper Holocene. From 6.5 ka cal. BP, Globorotalia inflata increasesfrom 10 to 30% and becomes one of the dominant species in thisinterval. A noticeable decrease of G. inflata is concomitant to the slightincrease of the polar species Neogloboquadrina pachyderma (l.c.) at8.2 ka cal. BP (Fig. 8).

4.4. Paleotemperature reconstructions from core MD04-2797CQ proxyrecord (Fig. 7)

The modern analogue technique applied to dinocyst assemblages(c.f. Section 3.3) reveals generally good analogues from the Bölling/

Alleröd to the present. The lack of a good analogue during the glacialperiod could be due to the high percentages of the cosmopolitanspecies Operculolodinium centrocarpum in association with Spiniferitesdelicatus and Spiniferites membranaceus. We also observe highdissimilarity coefficients in the reconstruction of SST based onplanktonic foraminifera during the Last Glacial Maximum possiblydue in part to the presence of Globigerinoides ruber.

Although, the spring SST estimations based on the dinocyst andplanktonic foraminiferal assemblages show equivalent values (9–11 °C) during the glacial period, it appears that the SSTdinocyst (6 °C)are lower than SSTforaminifera (10 °C) during the Younger Dryas.

During the Bölling/Alleröd, the mean U37k SST was 300 °C lower

than the values reconstructed for the Holocene (20 °C). In contrast,SSTdinocyst and SSTforaminifera (17 °C) remain similar to those recordedduring the Holocene. The onset of the highest sea surface tempera-tures derived from the different proxies during the Holocene isobserved only at ~10 ka cal. BP.

5. Discussion

During Heinrich events, the maximal development of cold dinocystspecies Bitectatodinium tepikiense, Spiniferites elongatus and Nemato-sphaeropsis labyrinthus is synchronouswith the appearance of the polarplanktonic foraminifera species Neogloboquadrina pachyderma (l.c.) inour records (Fig. 6). The association of this cold microflora with thisplanktonic foraminifera may be useful for the detection of the responseof the Mediterranean to the North Atlantic Heinrich events. On thePortuguesemargin, Heinrich events (H2 and H1) were characterised bytwo distinct phases: one with N. pachyderma (l.c.) associated withmaximum input of Ice Rafted Debris (IRD), and a second one with thedinoflagellate cyst B. tepikiense (Turon et al., 2003).

Neogloboquadrina pachyderma (l.c.), which thrives today wherethe SST is below 7 °C (Reynolds and Thunell, 1986) was presentduring the upper part of H2 and during H1 but only in low percentages(1–3%). However, in the Alboran Sea, this species reached 12% duringH2 (Cacho et al., 1999) but was absent in the Cretan basin (EasternMediterranean) (Geraga et al., 2005). Thismay indicate that cooling inthe eastern Mediterranean during Heinrich events was not as intenseas that in the western Mediterranean (Geraga et al., 2005).

In spite of the lack of good analogues (c.f. Section 3.3), thequantitative approach based on dinocysts and planktonic foraminifera(Fig. 7) shows lowSST values duringHeinrich events in accordancewiththe alkenones record (Essallami, 2007). This cooling is observed also inthe Tyrrhenian Sea (Kallel et al., 1997b; Sbaffi et al., 2001), in the Gulf ofLions (Melki et al., 1999) and in the Alboran Sea (Cacho et al., 1999;Perez-Folgado et al., 2003; Voelker et al., 2006) allowing us to confirmthat the Mediterranean Sea was cool during Heinrich events. Pollenrecords from the Adriatic Sea (Combourieu-Nebout et al., 1998; Guintaet al., 2003), Alboran Sea (Combourieu-Nebout et al., 1999, 2002;Fletcher and Sanchez-Goni, 2008) and Iberian margin (Sanchez Goniet al., 2002; Turon et al., 2003) have shown that the Heinrich eventswere represented by dry and cold climates on the neighbouringborderlands of the Mediterranean Sea. This reveals the sensitivity ofthe marine environment and atmospheric conditions in the westernMediterranean tomassive iceberg discharge and associated paleoenvir-onmental changes in the North Atlantic at these times (Cacho et al.,1999, 2000;Moreno et al., 2002; Perez-Folgado et al., 2003; Colmenero-Hidalgo et al., 2004; Sierro et al., 2005; Cacho et al., 2006; Voelker et al.,2006; Fletcher and Sanchez-Goni, 2008).

The Bölling/Alleröd is characterised by a contemporaneous devel-opment of the warm microflora and microfauna species Spiniferitesmirabilis and Globorotalia inflata respectively. Pollen records show acontraction of Artemisia-rich steppe areas and well-developed forestvegetation in the Adriatic Sea (Combourieu-Nebout et al., 1998; Guintaet al., 2003), in the Alboran Sea (Fletcher and Sanchez-Goni, 2008;Fletcher et al., 2009) and in the Iberianmargins highland (Fletcher et al.,

Table 2Average age of Heinrich events (H1 and H2) (Elliot et al., 1998; Bard et al., 2000).

Age(ka cal. BP)

Heinrich event 1 (H1) 15.5–18Heinrich event 2 (H2) 23–26

The Last Glacial Maximum (23–18 ka cal. BP) was limited by H2 and H1 in agreementwith EPILOG Workshop (1999) and MARGO (Mix et al., 2001; Kucera et al., 2005).



Fig. 6. Relative abundance of main species of planktonic foraminifera in core MD04-2797CQ. Grey bands indicate Heinrich events (H2 and H1) and the Younger Dryas event.

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2007). These simultaneous responses of the vegetation, the microfloraand the microfauna confirm that the Bölling/Alleröd is warm and lessdry than the Heinrich events. In contrast, the alkenones SST (Essallami,2007) show a less pronounced warming than that recorded by thedinocysts and planktonic foraminifera.

The cooling of the Younger Dryas, revealed by the maxima of cooldinocyst and planktonic foraminifera species between 13 and 11.5 kacal. BP, is contemporaneous to that recorded in the Tyrrhenian Sea(Kallel et al., 1997a), the Gulf of Lions (Melki et al., 1999) and in theAlboran Sea (Cacho et al., 1999). Also during this event, an expansionof semi-desert vegetation is recorded in the Alboran Sea (Fletcheret al., 2009) and in the Adriatic Sea (Combourieu-Nebout et al., 1998)

suggesting a dry climate on the neighbouring borderlands of theMediterranean Sea as during Heinrich events.

Themaximum dinocyst concentration observed during the YoungerDryas suggests an increase of nutrients in the Sicilian–Tunisian Straitduring this period. Therefore, this situation is confirmed by the optimaldevelopment of Nematosphaeropsis labyrinthus in this interval, which iscurrently associated to cold water masses with high (winter) nutrientsavailability (Turon and Londeix, 1988; Devilliers and de Vernal, 2000).

The onset of the Holocene was marked not only by the increase ofwarm dinocyst species and planktonic foraminifera but also with thepersistence of cool microflora and microfauna species. Our datasuggests that the warming is strongest at about 10 ka cal. BP. In fact,

Fig. 7. Results of MD04-2797CQ SST reconstructions, (A) MAT Spring SST dinocyst (B) dissimilarity coefficient relative to estimation of SST based on dinocyst assemblages. Thisparameter measures the degree of similarity between each fossil sample and three best modern analogues. (C) MAT Spring SST planktonic foraminifera (D) dissimilarity coefficientrelative to estimation of SST based on planktonic foraminifera assemblages. This parameter measures the degree of similarity between each fossil sample and five best modernanalogues. (E) U37

k index SST. Grey bands indicate Heinrich events (H2 and H1) and the Younger Dryas event.



the first occurrence of Pistacia in the Mediterranean forest, whichsuggests a notable rise in winter temperature on the continent, isfrequently detected only some 1500 years after the beginning ofHolocene (Willis, 1994; Rossignol-Strick, 1995; Watts et al., 1996;Combourieu-Nebout et al., 1998, 1999, 2002). Contemporaneouspollen records from the Adriatic Sea show a large expansion of foresttaxa (Combourieu-Nebout et al., 1998; Guinta et al., 2003) withincreases of Quercus and then progressively all the temperate trees.Such improvement of climatic conditions was synchronous with thebeginning of the Holocene Climatic Optimum, which occurredbetween 9 ka and 6 ka cal. BP in the North Atlantic Ocean (Eynaudet al., 2004; de Vernal and Hillaire-Marcel, 2006; Turon et al., 2008).

At this period, the increase of the POC values suggests theestablishment of presapropelic conditions in the bottom water ofthe Sicilian–Tunisian Strait (Fig. 4) at about the same time as thedeposition of sapropel S1 in eastern Mediterranean (Rossignol-Strick,1985; Kallel et al., 1997b; Rossignol-Strick and Paterne, 1999;Ariztegui et al., 2000; Rohling et al., 2002).

However, this period is interrupted by a brief decrease in warmdinocyst species Impagidinium aculeatum and Spiniferites mirabilis, andthe planktonic foraminiferaGloborotalia inflata at ~8.2 ka cal. BP (Fig. 8),contemporaneouswith the brief decrease of the POC values (Fig. 4) andthe S1 temporary interruption in the Eastern Mediterranean (de Rijk etal., 1999). At this time, a forest decline is recorded in pollen data fromthe Alboran Sea, suggesting a brief cold and dry event (Fletcher et al.,2009). This distinctive climatic shift has been linked to a high-latitudecooling event detected in δ18O and accumulation signals in Greenlandice cores know as the ‘8.2 event’ (Thomas et al., 2007) and associated, inaddition to the coolingevent recorded at9.2 ka cal. BP,withLakeAgassizmeltwater discharges (Teller et al., 2002).

Another brief cold event between ~6.5 ka and 7 ka cal. BP isrevealed by reconstruction of SST based on the different proxies that

may correlate with the Ungava and Labrador Lakes meltwaterdischarges (Jansson and Kleman, 2004).

During the late Holocene, the abundance of Impagidinium aculea-tum decreases while that of Spiniferites mirabilis increases. In spite ofthis change in microflora abundance, the SST estimations based ondinocyst assemblages and the other proxies remain almost the sameas those recorded during the early Holocene. At ~2.2 ka cal. BP, a slightcooling contemporaneous to that observed in Norwegian Sea (Calvoet al., 2002) is recorded only by dinocyst assemblages.

6. Conclusion

The evolution of the dinocyst and planktonic foraminiferalassemblages observed in the deep-core MD04-2797CQ exhibits asequence of paleontological associated with climatic change. Theoccurrence of sub-polar and temperate dinocyst and foraminiferalspecies respectively in the Sicilian–Tunisia Strait, characterises theresponse of the Mediterranean to North Atlantic Heinrich eventsH2 and H1 confirming the sensitivity of the region to climatic eventsoccurring during the last glacial cycle in the Northern Hemispherehigh latitudes.

During the Bölling/Alleröd, the U37k SST was 3 °C lower than the

values reconstructed for the Holocene (20 °C). In contrast, SSTdinocystand SSTforaminifera remain similar to those recorded throughout duringthe Holocene; these discrepancies remain poorly understood.

Although different seasonswere involved in the SST reconstructions,whichwere based onmicropaleontological and biogeochemical proxies(dinocyst, planktonic foraminifera and alkenones), a significant coolingevent synchronous to the establishment of dry conditions in theMediterranean is recorded during the Younger Dryas. Also, a maximumdinocyst concentration was observed suggestive of an increase innutrient availability in the Sicilian–Tunisian Strait at that time.

Fig. 8. Relative abundance of main warm species of dinocyst (S. mirabilis and I. aculeatum) and planktonic foraminifera (G. inflata) with relative abundances of main cold species ofdinocyst (B. tepikiense, N. labyrinthus) and planktonic foraminifera (N. pachyderma l.c) during the marine isotopic stage 1.



Around 1500 years after the onset of the Holocene, SST recon-struction based on dinocyst, planktonic foraminifera and alkenonesreveals the warmest values observed in the sequence. During the lateHolocene, percentages of the dinocyst Spiniferites mirabilis and theplanktonic foraminifera Globorotalia inflata increase but those of thedinocyst Impagidinium aculeatum and the planktonic foraminiferaGlobigerinoides ruber decrease. While these microflora and microfau-na change, SST values remain equal to those recorded during the earlyHolocene suggesting no evidence of a Hypsithermal in the Mediter-ranean Sea as observed in the Boreal to Atlantic European margin.

Abrupt decreases of the percentages of warm microflora andmicrofauna species at brief cooling events are recorded as 9.2 ka,~8.2 ka, between 6.5 and 7 ka and ~2 ka cal. BP. The ‘8.2 cooling event’is contemporaneous to the presapropelic condition in the Sicilian–Tunisian Strait and the S1 temporary interruption in the EasternMediterranean Sea (de Rijk et al., 1999). The first two cooling episodes(9.2 ka, ~8.2 ka) may be correlated to the meltwater discharges ofLake Agassiz (Teller et al., 2002) but the third (6.5–7 ka cal. BP) maybe linked to the meltwater discharges of the Ungava-Labrador Lakes(Jansson and Kleman, 2004).

Acknowledgements

The authors thank the Institut National des Sciences de l'Univers(INSU) of the Centre National de la Recherche Scientifique (CNRS), theRV Marion-Dufresne officers and crew, the IMAGES program and theInstitut Paul Emile Victor (IPEV) for support and organisation of thecoring cruises. We would finally like to thank B. Lecoat for isotopeanalyses, ARTEMIS for the radiocarbon age measurements and M.H.Castera (UMR-EPOC5805) for preparing the dinocyst samples.

I. R-Z. and N.K. gratefully acknowledge the financial supportprovided by the French–Tunisian CMCU (Comité Mixte de Coopéra-tion Universitaire) joint project 05-S-1004.

We also gratefully thank Eynaud, F. and Flatcher, W. for theirhelpful comments on this manuscript.

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Documents

Environmental and climatic changes in the central Mediterranean Sea (Siculo–Tunisian Strait) during the last 30ka based on dinoflagellate cyst and planktonic foraminifera assemblages